Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico 9781501734618

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Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico

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Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico RICHARD H. FOOTE Agricultural Research Service U.S. Department of Agriculture (Retired)

F. L. BLANC California Department of Agriculture (Retired)

ALLEN L. NORRBOM Agricultural Research Service U.S. Department of Agriculture

COMSTOCK PUBLISHING ASSOCIATES

a division of Cornell University Press Ithaca and London

Copyright © 1993 by Cornell University All rights reserved. Except for brief quotations in a review, this book, or parts thereof, must not be reproduced in any form without permission in writing from the publisher. For information, address Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850 First published 1993 by Cornell University Press. Printed in the United States of America

@ The paper in this book meets the minimum requirements of the American National Standard for Information Sciences— Permanence of Paper for Printed Fibrary Materials, ANSI Z39.48-1984.

Library of Congress Cataloging-in-Publication Data Foote, Richard H. (Richard Herbert), 1918Handbook of the fruit flies (Diptera-.Tephritidae) of America north of Mexico / Richard H. Foote, F. L. Blanc, Allen L. Norrbom. p. cm. Includes bibliographical references and indexes. ISBN 0-8014-2623-5 1. Tephritidae—United States—Classification. 2. Tephritidae— Canada—Classification. 1. Blanc, F. L. (Francis Louis), 1916— . II. Norrbom, Allen L. (Allen Lee), 1957- . 111. Title. QL537.T42F66 1993 595.77'4—dc20 92-52844

To Alan Stone Whose encouragement and cooperation early in the project were indispensable

Digitized by the Internet Archive in 2018 with funding from The Arcadia Fund

https://archive.org/details/handbookoffruitfOOfoot

CONTENTS

ACKNOWLEDGMENTS

XI

Introduction

1

Adult Morphology

3

Head, 3

Thorax, 5

Abdomen, 10

General Features, 13

Biology

14

Classification of the Tephritidae Relationships to Other Families of Diptera, 17 Genera, 18

The U.S. and Canadian

A Classification of the Genera Occurring in America North

of Mexico, 20

Current Status of Taxonomic Studies Nearctic Region, 34

Neotropical Region, 35

Afrotropical Region, 35

Oriental Region, 36

34 Palearctic Region, 35 Australasian-Oceanic

Region, 36

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Vlll

Contents

Techniques for Study Mounting, 37

Specimens in Fluids, 38

Maceration (Clearing), 39

Immature Stages, 38

Aculeus, 39

Labeling, 38

Male Genitalia, 41

Wings, 41

About This Handbook Keys, 42

Illustrations, 42

Genera, 43

Data, 44

Distribution, 45

Hosts, 45

Species, 43

Literature, 43

Terminology, 45

Type

Nomenclature, 46

Key to the U.S. and Canadian Genera

Systematic Treatment of the Genera Genus Acidogona Loew, 65 Aciurinu Curran, 69

Schiner, 83

Genus Acinia Robineau-Desvoidy, 67

Genus Acrotueniu Loew, 81

Genus Buctvocevci Miacquart, 112

Genus Cecidocbarella Hendel, 122

Benjamin, 154

Hering, 202

Genus Dcicus Fabncius, 132

Genus Dyseuaresta Hendel, 137

Genus Euaresta Loew, 141

Genus Euleia Walker, 161

Eutreta Loew, 179

Genus Cdlldchuci Aldrich, 120

Genus Chetostoma Rondani, 128

Genus Cvyptotvctu Blanc and Foote, 130

Epocbra Loew, 140

Genus Anustrepba

Genus Ceratitis Macleay, 123

Genus Gbaetostomella Hendel, 126

Genus Dioxyna Frey, 133

Genus Hexacbaeta Loew, 205

Genus

Genus Euarestoides

Genus Eurosta Loew, 165

Genus Gonioxyna Hendel, 200

Genus Icterica Loew, 209

Genus Metatepbritis Foote, 213

Mylogymnocarena Foote, 214

Genus Myolejd Rondani, 216

Neaspilota Osten Sacken, 222

Genus Neotepbritis Hendel, 243

terellia Foote, 270

Genus Paracantba Coquillett, 262

Genus Paroxyna Hendel, 274

Genus Procecidocbares Hendel, 303 Genus Rbagoletis Loew, 322

Genus

Genus Para-

Genus Peronyma Loew, 301

Genus Procecidocbaroides Foote, 318

Genus Rbagoletotrypeta Aczel, 365 Genus Stenopa Loew, 369

Strauzia Robineau-Desvoidy, 373

Genus Tepbritis Latreille, 387

Trupanea Schrank, 416

Robineau-Desvoidy, 457 Xantbaciura Hendel, 481 Xenocbaeta Snow, 490

Genus Genus Oxyna

Genus Rbynencina Johnson, 368

Tomoplagia Coquillett, 410

Genus

Genus

Genus Orellia Robineau-Desvoidy, 253

Robineau-Desvoidy, 259

Genus

Genus Gymnocarena

Jamesomyia Quisenberry, 211

Oedicarena Loew, 247

Genus

Genus Genus

Genus Toxotrypana Gerstaecker, 414

Genus Trypeta Meigen, 448

Genus

Genus Uropbora

Genus Valentibulld Foote and Blanc, 475 Genus Xantbomyia Phillips, 487 Genus Xonosemata Benjamin, 493

Genus

Genus

Contents

REFERENCES CITED INDEX OF INSECT NAMES INDEX OF PLANT NAMES

499 553 564

ACKNOWLEDGMENTS

This project was initiated in 1954 in that part of the USDA’s Systematic Entomology Laboratory located in the Smithsonian’s National Museum of Natural History in Washington, D.C. The identification of large numbers of fruit flies from North America and abroad and the presence in the museum of a massive tephritid reference collection presented an unparalleled opportunity to gather the kinds of data required to produce a book such as this. Shortly after 1954, F. L. Blanc, who was engaged in survey and identification activities in the California Department of Agriculture in Sacramento, joined the project. A. L. Norrbom succeeded R. H. Foote in Washing¬ ton, and as his expertise in the systematics of the family grew, he was asked to join the project as well. Our institutions, the USDA-ARS Systematic Entomology Laboratory (SEL) and the California Department of Agriculture, have made the most important contribu¬ tions to our project by providing salaries, facilities, a great amount of time by formalizing our activities as a research project, and additional funds for the comple¬ tion of our maps and illustrations. We are especially indebted to W. H. Anderson, R W. Oman, R. I. Sailer, L. V. Knutson, R. W. Hodges, R M. Marsh, and D. R. Miller, who, in their various administrative capacities, made all these advantages possible. We hereby acknowledge with gratitude the contributions made by our colleagues associated with the Diptera Unit, SEL, with whom we have been closely associated: R. J. Gagne, B. V. Peterson, C. W. Sabrosky (retired), G. C. Steyskal (retired), A. Stone (retired), F. C. Thompson, W. W. Wirth (retired), N. E. Woodley, and W. N. Mathis (Smithsonian Institution). Numerous institutions made type specimens and other study material and data available during the 39-year life of this project. Likewise, many individuals, associ¬ ated or not with those institutions, graciously extended their support in a variety of ways. To the following and their associated organizations, and to anyone we may have inadvertently omitted either here or in the text, we extend our heartfelt apprexi

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Acknowledgments ciation: M. L. Aczel, L. A. Andres, P. H. Arnaud, Jr., D. Azuma, S. W. T. Batra, H. D. Blocker, P. Boldt, G. L. Bush, G. W. Byers, L. E. Carroll, D. S. Chandler, R. Contraras-Lichtenberg, K. Corwin, J. R. Coulson, R. Danielsson, H. A. Denmark, G. Dodson, R. A. I. Drew, D. K. Faulkner, E. M. Fisher, B. A. Foote, K. E. Frick, S. I. Frommer, J. K. Gelhaus, G. Gordh, D. Grimaldi, D. H. Habeck, H.-Y. Han, W. J. Hanson, D. E. Hardy, P. Harris, V. Hernandez, E. R. Hoebeke, C. L. Hogue, B. R. Ibrahim, J. Jenkins, L. Matile, J. F. McAlpine, G. C. McGavin, K. C. McGiffen, C. E. Miller, E. Mitchell, F. W. Muesebeck, H. K. Munroe, J. A. Novak, M. O’Brien, W. A. Palmer, N. Papavero, L. Papp, V. T. Phillips, R. J. Prokopy, H. Schumann, J. L. Sharp, L. J. Shervis, G. E. Shewell, R. E. Snodgrass, C. E. Stegmaier, Jr., R. Stewart, W. B. Stoltzfus, D. W. S. Sutherland, C. E. Turner, W. J. Turner, K. R. Valley, J. K. Wangberg, M. S. Wasbauer, H. V. Weems, F. G. Werner, R. L. Wescott, R. A. Wharton, N. P. Wyatt, P. Wygodzinsky, C. W. Young, M. Yue, R. S. Zack, and H. Zwolfer. Other individuals are acknowledged in the text. The major collaborative contributions of our colleagues A. Freidberg, R. D. Goeden, D. L. Hancock, G. Steck, and I. M. White must be highlighted. In addition to providing critical study material and data, they offered substantial suggestions throughout the course of the research and during manuscript preparation. Their intimate knowledge of fruit flies has saved us from many grievous errors. We most gratefully acknowledge the indispensable help of those who relieved us of much tedious work on the project. N. Malikul handled thousands of specimens and made slide mounts of wings. R. W. Carlson eased us through the many complex¬ ities associated with program conversions and the efficient use of our computers. P. W. Iglehart, assisted by S. D. Diaz, E. B. Jamison, and V. M. Lee, spent untold hours carefully retyping the manuscript, treating it as if it were her own. H. B. Norton printed most of the photographic negatives used in the book. T. B. Griswold executed all the line drawings and was primarily responsible for making up and labeling the plates. M. Kohn devoted many hours to map design, L. H. Lawrence and E. R. Hodges guided us through the complexities of map making, Suzanne Shute applied most of the map symbols, and E. Lockett was instrumental in preparing the maps for reproduction. Finally, we are deeply grateful to H. Maddux and M. Quinto, whose patience and superb editing brought much-needed order to our rambling text. Our current addresses are as follows: Richard H. Foote, HC 75 Box 166, Locust Grove, VA 22508; F. L. Blanc, 5309 Spilman Ave., Sacramento, CA 95819; Allen L. Norrbom, Systematic Entomology Laboratory, USDA, c/o U.S. National Museum NHB 168, Washington, D.C. 20560. R. H. F. F. L. B. A. L. N.

INTRODUCTION

The family Tephritidae includes some of the most biologically interesting and most economically important species of flies. Commonly known as fruit flies, although sometimes called peacock flies (Borror et al. 1976) because of their coloration and habit of waving their wings, tephritids are among the most striking of insects. The wings of the brightly colored adults almost always are patterned with dark spots and bands on a clear field or clear spots on a dark background. The family Tephritidae occurs virtually everywhere in the temperate, subtropical, and tropical parts of the world except in desert and arctic areas where plant life is scarce or absent. The greatest diversity of species occurs in the tropics, but except in a few instances, those areas of the world are least explored. It is expected that many undescribed taxa will be discovered as these areas become more accessible for collect¬ ing. About 4,200 species in about 500 genera are currently known throughout the world. Of these, 300 species in 56 genera, all of which are discussed in this hand¬ book, have been found to be native to, or recently introduced into, the United States and Canada. The geographic area treated in this book is North America north of Mexico, including the continental United States, Canada, Greenland, Bermuda, and their associated islands; it excludes the Bahama and Commander Islands. The body lengths of individual flies range from about 2 mm to more than 20 mm, and mature larvae, especially some of those found in fruit, may be up to 15 mm long. The wings of most tephritids are marked in some way, usually with patterns charac¬ teristic of a genus or species. The head, body, and legs of some species also bear markings that can be used as identification characters. In the past, color patterns alone were used for identification; indeed, in many cases they have been found to be quite reliable for that purpose. On the other hand, biosystematists today have dem¬ onstrated that such characters in some groups are highly variable and thus cannot be depended upon for accurate recognition. Although we have attempted to restrict our recognition characters to those that are relatively easy to see, the user will find it 1

2

Introduction necessary to dissect the ovipositors of a limited number of species to achieve accurate identifications. With very few exceptions, none of which occur in North America, members of the family are intimately associated with and breed in living plants. In the typical devel¬ opmental cycle, the female inserts her eggs into plant material through an eversible ovipositor, the terminal segment of which is sclerotized and often needle- or trowel¬ like. Eggs'hatch in the plant tissue, and the resulting larvae progressively undergo three molts, developing an inactive fourth instar within the puparium. Pupation takes place either at the feeding site or in the ground below, then the emerging adults feed, attain sexual maturity, and mate, thus completing the life cycle. Most temperate zone tephritids undergo diapause, but some subtropical and most tropical species do not. Agriculturally, the Tephritidae is the most important of all fly families. Several species of Anastrepha and Rbagoletis are highly destructive to commercially grown fruit and vegetable crops in North America north of Mexico, and the accidentally introduced Mediterranean fruit fly and species of Bactrocera and Dacus discussed in this handbook are the subject of massive survey and eradication procedures when¬ ever they are detected. Agricultural quarantines must be rigorously enforced against the possibility of introduction of many other potentially harmful species not yet found here. On the other hand, some tephritid species have proved to be beneficial in the control of noxious weeds. No fewer than eight species of Uropbora have already been introduced from Europe into the United States and/or Canada for this purpose (White and Clement 1987), and efforts are being made to identify potentially valu¬ able members of other tephritid genera that might be valuable in controlling harmful plants in North America. For instance, Cbaetorellia australis Hering (Maddox et al. 1990) already has been released in the western United States for the control of starthistle (White and Marquardt 1989, Wood 1989), and Cbaetorellia acrolopbi White and Marquardt and Terellia virens (Loew) are being considered for introduc¬ tion to Canada from Europe (Groppe and Marquardt 1989a, 1989b). (These genera have been investigated too recently for inclusion in this handbook.) Species of the New World genera Acinia, Euaresta, Eutreta, Procecidocbares, Tetreuaresta, and Xantbaciura have been and are being introduced to Hawaii and other Pacific islands,

Australia, Europe, and the former USSR for control of weeds, and research on this phase of biological control continues today. Comments on the economic importance of individual genera and species are given on ensuing pages of this handbook.

ADULT MORPHOLOGY

HEAD (Figs. 1-3) The tephritid head exhibits many characters of taxonomic significance. Of primary importance is its shape in lateral view, the configuration of the face and frons, and the setation. Our terminology is shown in figs. 1—3 and associated legends. In profile (fig. 1), the head is higher than it is long in most North American species, but in a few species it is elongated. Whenever this character is used, it must be measured carefully because appearances are sometimes deceiving. The compound eye (comp eye) in all cases is oval or elongate-oval and offers few characters except in live specimens where color may be important (Steyskal 1957). In most species, the frons (fr) meets the face (fc) at an angle of 90-110° or more, but in those with an elongated head, this angle may be less than 90°. The face is variously convex, straight, or concave; it is occasionally so sharply concave that its medial part, often produced as a carina, cannot be seen beyond the parafacial (pafc). The latter varies in width and has been used taxonomically. The postgena (pgn) is invariably expanded to varying degrees. The height of the gena (fig. 2, gn) (the distance between the lowermost margin of the compound eye and the edge of the head capsule directly beneath it) in proportion to the eye height often is taxonomically significant. The proportion of length to width of the first flagellomere (1st fig) sometimes is useful for distinguishing species, and occasionally the configuration and color of the arista (ar), which is at most very short-haired in North American species, also are valuable. In top or front view (fig. 2), the width of the frons in relation to the width of the compound eye and the proportion of length (measured from vertex to lunule) to width (usually measured near the vertex at the greatest distance between the eyes) often are important. The palpi (fig. 2, pip) are not of any known taxonomic value. The proboscis is generally of two types; the most commonly encountered is the usual muscoid type with a padlike labellum (lbl), as shown in fig. 1. In not a few genera 3

i vt b

o b

Figures 1—3. Side, front, and rear views, respectively, of head of Anastrepha ludens (Lw.), showing abbreviations designating bristles and sclerites. Terms in parentheses are those used by some other authors, ar, arista; clyp, clypeus; comp eye, compound eye (eye); f b, frontal bristle (lower frontoorbital seta); fc, face; 1st fig, 1st flagellomere (3rd antennal segment, flagellum, postpedicel); fr, frons; gn, gena; gn b, genal bristle (seta); gn grv, genal groove; gn s, genal setulae; i vt b, inner vertical bristle (seta); lbl, labellum; lun, lunule; m ocp si, median occipital sclerite; o b, orbital bristle (upper fronto-orbital seta); oc, ocellus; oc b, ocellar bristle (seta); oc tr, ocellar triangle; ocp, occiput; ocp for, occipital foramen; ocp s, occipital setae; o vt b, outer vertical bristle (seta); pafc, parafacial; ped, pedicel (2nd antennal segment); pgn, postgena; pgn b, postgenal bristle (seta); pgn s, postgenal setulae; pip, palpus; poc b, postocellar bristle (seta); pocl s, postocular setae; ptil fis, ptilinal fissure; pv b, postvertical bristle (seta); scp, scape (1st antennal segment); spc s, supracervi¬ cal setae; vrt, vertex.

5

Adult Morphology having an elongate head, especially Oxyna, Paroxyna, Gonioxyna, Dioxyna, and some Uropbora, and in a few species belonging to other genera, the labellum is clearly elongated and retroflexed (geniculate) (see figs. 19, c; 59, b). This condition is seen clearly from a ventral view if the mouthparts are folded close to the head. The setation of the head is frequently of critical importance. Most notable is the number of frontal bristles (f b), which separates large groups of genera as shown in the key. These bristles (2-5 pairs) are always present and are usually strong and prominent; some of them are occasionally whitish as in Oxyna and Paracantha. In several genera (e.g., Myoleja, Strauzia), some of the species have greatly enlarged and blackened frontals, especially in the males. The orbital bristles (o b), always present in 1-3 pairs in North American species, are invariably separated as a set from the frontals. In males of the introduced Mediterranean fruit fly, the anterior orbital is forwardly directed (proclinate) and terminates in a flat, diamond-shaped expan¬ sion. In the flower-feeding tephritids, the posterior pair of orbitals is almost always a light color in contrast to the darker anterior pair. In several genera, the posterior orbitals are convergent; this is a character common to members of the tribe Terelliini (Chaetostomella, Orellia, and Neaspilota in North America) and several other unre¬ lated species or genera. The ocellar bristles (oc b) are strong, except in isolated cases such as Anastrepha, Bactrocera, Dacus, Toxotrypana, some species of Myoleja and Trypeta, and certain other genera outside North America, in which they are absent or relatively minute. The postocular setae (pocl s) in most of the frugivorous genera are black and slender, whereas in the flower-feeding genera they are usually whitish, fewer in number, and somewhat thickened, or are a mixture of the two types. The genal bristle (gn b) and genal setae (gn s) vary in size but are always present; the genal setae may be numerous anteriorly, forming a small tuft or congregation of longer, heavier hairs. The remaining head bristles and setae, including those of the palpi, remain little-studied. The color of the head is variable but usually is similar to that of the thorax. The frons occasionally is a color different from the remainder of the head capsule, whereas the face is most often of a lighter hue. In a few genera (e.g., Paracantha and some Eutreta), the face and parafacial (pafc) may have distinctive black markings, and in some species the occiput (fig. 3, ocp) and postgena (pgn) are darkened, sometimes in a distinctive pattern (best seen when viewed from behind) (fig. 3). The first flagellomere in some species is darkened as well, although rarely consistently enough to be of taxonomic value.

THORAX (Figs. 4, 5) The configuration of the thorax is generally comparable to that found in most other acalyptrate flies. Except for the scutellum, the physical structure, shape, and inter¬ relationships of sclerites are rarely used, possibly because they have been little studied in the family. Setation and color provide most of the taxonomic characters of value. Our terminology is illustrated and explained in figs. 4 and 5. The nature and position of bristles are of critical taxonomic importance. Especially significant is the position on the scutum of the dorsocentral bristles (dc b, prs dc b);

6

Adult Morphology scap b

scap b

pp I

pp I b

npl b

spal b

dc b

acr b

sett b sett sbsctl mtg

Figures 4, 5. Top and side views, respectively, of thorax of Anastrepha ludens (Lw.), showing names for bristles and sclerites. Note that presutural dorsocentral bristles, not present in A. ludens, have been added for illustrative purposes. Terms in parentheses are those used by other authors, a sctl b, anterior scutellar bristle (seta); abd, abdomen; acr b, acrostichal bristles (setae); anepm, anepimeron (pteropleuron); anepm b, anepimeral bristle (seta); anepst, anepisternum (mesopleuron); anepst b, anepisternal bristles (setae); as, anterior spiracle; cx 1, 2, 3, coxae of fore, middle, and hind legs; dc b, dorsocentral bristle (seta); hit, halter; ial b, intra-alar bristle (seta); kepst, katepisternum (sternopleuron); kepst b, katepisternal bristle (seta); mtg, mediotergite (postnotum, metanotum); npl, notopleuron; npl b, notopleural bristle (seta); p sctl b, posterior scutellar bristle (seta); pal b, postalar bristle (seta); pp 1 postpronotal lobe (humerus); pp 1 b, postpronotal lobe bristle (humeral bristle, seta); prs b, presutural supra-alar bristle (seta); prs dc b, presutural dorsocentral bristle (seta) (not present in Anastrepha spp.); sbsctl, subscutellum (postscutellum); scap b, scapular bristle; set, scutum; sctl, scutellum; sctl b, scutellar bristle; sctsctl sut, scutoscutellar suture; spal b, supra-alar bristle (seta); trn sut, transverse suture; wg b, wing base.

,

their position is one of the primary divisions in the key to genera. In a large group of genera, the dorsocentrals are inserted very close to the acrostichal bristles (acr b) as shown in fig. 4, in others very close to and behind the transverse suture (trn sut), and in others this position varies between the two extremes. In several genera, an addi¬ tional pair of dorsocentrals (prs dc b) (figs. 4; 53, a) arises anterior to the suture, most often when the posterior pair of dorsocentrals is relatively close to the suture. Among the genera found in North America, only in Bactrocera and Dacus are the bristles of the postpronotal lobe (pp 1, pp 1 b) and the dorsocentrals absent, although they are weak in Toxotrypana. The notopleural bristles (npl b) shown in figs. 4 and 5 are always present, and the posterior pair in most flower-feeding species is a lighter hue than the anterior pair. The presence and number of anepisternal bristles (anepst b) are sometimes critical. In the genus Procecidochares and in a few species in other genera, often-used taxonomic characters are the number and arrangement of numer¬ ous, short, whitish, expanded setae distributed over the scutum. The occurrence and nature of other scutal, pleural, and sternal bristles have not been used as taxonomic characters in North American tephritids.

7

Adult Morphology The characters of the scutellum (sctl) and associated structures are important taxonomically. In dorsal view (fig. 4), the scutellum is essentially triangular; its apex is rounded or pointed and of varying sharpness. In several genera, the scutellum is swollen and the dorsal surface is rounded and polished, and in Peronyma it is divided longitudinally by a shallow depression. Otherwise, the scutellum is rather flat with rounded lateral edges. The subscutellum (sbsctl) and mediotergite (mtg) are not haired, but various color patterns on these two sclerites may be significant; they can be viewed best from behind. In only one genus occurring in North America (Hexachaeta) does the scutellum bear three pairs of scutellar bristles (sctl b); otherwise only one or two pairs are present. The basal pair is always strong, but occasionally it may be difficult to decide whether a second, apical pair is present because they may be extremely small or whitish and expanded. The disk or margin of the scutellum sometimes bears additional small setae. Color patterns of the thorax can be significant (for example, see figs. 157—163). They consist either of contrasting pigmentation in the form of stripes or spots, especially in light-colored species, or are caused by the nature of the tomentum on the surface of the cuticle. In the latter case, especially on the scutum, the pattern is most easily observed at a low angle from behind with the light source also at a low angle directed toward the observer.

Wing (Fig. 6)

Taxonomic characters found in the tephritid wing are of primary importance in the identification of most genera and species. These reside not only in the pattern but also in the arrangement of the veins and shapes of the cells. The general wing shape, however, is of little taxonomic value in North American Tephritidae. The terminology is shown in fig. 6 and its legend. As mentioned elsewhere, one of the principal recognition characters for the family Tephritidae is the nature of the subcostal vein (Sc), which is more or less sharply bent anteriorly near its apex, changing its course beyond the bend to join the costa at a nearly right angle. Vein Sc almost always becomes very weak beyond the bend, and in some species it almost completely disappears along the base of the pterostigma. Whenever reference is made to cell sc, only that part of the cell basad of the bend is intended (i.e., the elongated, very narrow portion of the cell between the longitudinal part of vein Sc and vein R,). In this handbook, we have used the term pterostigma for the cell apicad of the bend in vein Sc. For the recognition of genera within the family, other venational characters are of value. Vein R2+^ may be sinuate, and in at least two genera, there are one or more supernumerary crossveins extending transversely between the costa and that vein. The positions of crossveins r-m and dm-cu relative to each other is sometimes tax¬ onomically significant. The terminations of these veins on vein M range from quite close together to distinctly farther apart than the length of either crossvein. Most veins extend more or less straight to the wing margins except in the genus Anastrepha, in which vein M curves forward apically, in most species gradually merging with the apex of the costa (fig. 24, a). Generally speaking, the arrangement of veins and cells in the basal half of the wing is constant except for the shape of cell cup. That cell may be cut off apically by the transverse or outwardly convex nature of vein

8

Adult Morphology

Figure 6. Right wing of Anastrepha ludens (Lw.), showing abbreviations for veins and cells. Terms in parentheses are those used by other authors. Cells and crossveins appear in lowercase letters, longitudinal veins in capitals. Aj, 1st anal vein; Aj + CuA2, 2nd branch of anterior cubital vein and 1st anal vein (anal vein, 1st anal vein, Cu2 + 2nd A, 6th longitudinal vein); alula, axillary lobe; anal lobe, anal lobe (anal cell, 4th posterior cell, 2nd anal cell, 2nd A, axillary lobe); be, basal costal cell (1st costal cell, 1st C); bm, basal medial cell (M, 2nd basal cell); bm-cu, basal medial-cubital crossvein (M3, anterior basal cross vein); br, basal radial cell (R, 1st basal cell); C, costa (costal vein); c, costal cell (2nd costal cell, 2nd C); cuaj, anterior cubital cell (apical cubital cell, 3rd posterior cell, Cu,); CuAj, 1st branch of anterior cubital vein (cubitus, Cu, M3 + Cu, Cu2; 5th longitudinal vein); CuA2, 2nd branch of anterior cubital vein (Cu2, cu-an, posterior basal cross¬ vein); cup, posterior cubital cell (basal cubital cell, anal cell, 1st anal cell); dm, discal medial cell (1st M2, discal, discoidal cell); dm-cu, discal medial-cubital crossvein (posterior crossvein, hind poste¬ rior crossvein, vein im, vein m, median crossvein, lower crossvein); h, humeral crossvein (basal crossvein); M, medial vein (M1+2, discoidal vein, 4th longitudinal vein); m, medial cell (apical medial cell, 2nd M2, 2nd posterior cell); pterostigma, expanded end of subcostal cell (stigma, mediastinal cell); r-m, radial-medial crossvein (anterior crossvein, Ta, median crossvein, upper crossvein); Rs, sectoral branch of radial vein (1st radial vein, 1st longitudinal vein); R1? sectoral branch of radial vein (1st longitudinal vein) r„, anterior radial cell (marginal cell, subcostal cell); R2 + 3, sectoral branch of radial vein (R3, 2nd longitudinal vein); r2 + 3, sectoral radial cell (r3, submarginal cell, cubital cell); R4 + 5, sectoral branch of radial vein (R5, 3rd longitudinal vein); r4 + 5, sectoral radial cell (r5, 1st posterior cell); Sc, subcostal vein (auxiliary vein, mediastinal vein); sc, subcostal cell.

CuA2 (see fig. 17, b), as in all genera belonging to the subfamily Myopitinae. In other genera, there is an elongation of the cell along vein Aj 4- CuA2 of varying extent (figs. 13, c; 18, a). The greatest elongations are exhibited by the frugivorous species, whereas the shortest are featured in the flower feeders. Setation of the veins varies and is of some taxonomic value. Vein Rj is haired in nearly all species; several genera possess dorsal setulae at least at the branching of veins R2+3 and R4+5, the setulae sometimes extending well past crossvein r-m on the latter vein. In general, the flower-inhabiting species lack or have reduced setation on vein R44-5. None of the other longitudinal or cross veins possesses setulae in Ameri¬ can species occurring north of Mexico. All the significant wing setulae are found on the dorsal (upper) side of the wing.

9

Adult Morphology The wing membrane is covered in whole or in part by minute microtrichia; their color contributes to the nature of the wing pattern, and their distribution may be species-specific. The microtrichia have not been studied or used as taxonomic characters in North American species. In a few species, the wing membrane is thickened and sometimes specially colored in a small area in cells r2+3 and r4+5. The function of this area, the bulla (see fig. 309, a), is not known. The color pattern of tephritid wings is relied upon heavily for the identification of genera and species, and in some genera its use is the only means for making conve¬ nient and rapid identifications. The pattern is of two principal types: a series of more or less transverse or oblique dark bands on an essentially hyaline background, or a collection of numerous hyaline or yellowed spots on a dark background. In at least one genus, Acrotaenia (fig. 106), a combination of both kinds of pattern is found. Although many exceptions occur, generally the banded type is characteristic of frugivorous species, whereas the spotted condition is more commonly found among the flower-inhabiting ones. The nature of the markings is sometimes characteristic of all or most of the species in a given genus. For example, Anastrepha species typically show a costal band, an S-band, and a V-band (see fig. 126); in most Tomoplagia species the bands are distinctively oblique (fig. 86); in many Trupanea species, one finds a characteristic “stellate” pattern (figs. 446-472); in most Paroxyna species (fig. 335, a), there is a rounded hyaline spot in the center of the pterostigma. Legs In almost all North American Tephritidae, the three pairs of legs are of nearly equal size, the forelegs shortest and the hind legs longest. The fore femur is enlarged in circumference only in males of Euaresta, one of the recognition charac¬ ters for that genus in North America. South of the United States, many other species still unplaced to genus possess that character as well. The tibiae of all legs are slender, and the tarsomeres, five on each leg, are relatively short. In only one genus studied so far for this character (Neaspilota), the fifth tarsomere of males of all but one species shows asymmetry that usually involves the claws, pulvilli, and tarsal comb (see figs. 275-282). Taxonomic characters other than color that may be present in the coxae and trochanters generally have been ignored and are largely unstudied in the Tephritidae. The fore femur of nearly all North American tephritids is the most heavily haired, and most species possess a loose “comb” of quite long, prominent anteroventral setae in addition to other hairs. The bristling of the middle legs is generally in¬ conspicuous. The hind femur of the genus Tephritis has a single anterodorsal preapical bristle, whereas most species of Tephritidae have two to several dorsal bristles, the apical-most pair side by side. The posterior tibiae of both sexes in such genera as Rhagoletotrypeta, Oedicarena, and some other frugivorous species possess a comb of dorsal or anterodorsal bristles of varying extent, as does at least one known species of Tephritis. The leg color is generally yellowish or concolorous with the body. Any darkening present usually occurs on the femora or on the tarsomeres and may vary from inconspicuous to coal black. Coloration of the legs is often highly variable intraspecifically and cannot be fully depended upon taxonomically except in rare cases. In

10

Adult Morphology a few species (e.g., Paracantha) (figs. 314, 315), discrete black femoral spots may be present.

ABDOMEN (Figs. 7—11)

The configuration of the tephritid abdomen offers many valid taxonomic characters, especially in details of the genitalia (the so-called postabdomen) in both males and females. The characters, illustrated in figs. 7—11, are explained in the accompanying legends. Female (Figs. 7—9)

The female abdomen proper (the “preabdomen”) consists of

six visible segments, each comprising a tergite (T1—T6) and a sternite (SI—S6) connected by membranes. Invariably, the abdomen from dorsal view is widest at the

snm

Figures 7—9. Generalized views of female abdomen. 7, Top view, ovipositor fully extended. 8, Retracted tephritid ovipositor. 9, Side view, ovipositor fully extended. Terms in parentheses are those used by other authors, acu, aculeus (ovipositor, abdominal segments 8 and 9, gynium); acu t, aculeus tip; ev m, eversible membrane (eversible ovipositor sheath, ovipositubus, conjunctiva of segment 7); fl, genital flaps (8th sternite); ovscp, oviscape (ovipositor sheath, segment 7, syntergosternite 7); S1-S6, sternites of abdominal segments 1-6; sctl, scutellum; snm, sternum of preabdomen; t, taenia; tgm, tergum of preabdomen; th, teeth (rasper); T1-T6, tergites of abdomi¬ nal segments 1—6.

11

Adult Morphology second segment (T2), whereas the first tergite (Tl) is reduced in size. It is fused to T2 and is difficult to discern. Posteriorly, the abdomen narrows to the smallest segment (T6), followed by the tergite and sternite of segment 7, which are fused to form the oviscape (ovscp). This contains, at rest, the eversible membrane (ev m) and the aculeus (acu), as shown schematically in figs. 7-9. In preserved specimens, the oviscape may be either flattened or rounded and tube- or funnel-like. Its length relative to the preceding abdominal tergites is often used as a taxonomic character, but we regard this ratio as an unreliable character because some of the base of the oviscape may be hidden under the sixth tergite, and the tergites may not always be distributed evenly along the dorsum of the abdomen in preserved specimens. In some species and species groups, the length of the oviscape may be quite variable, not corresponding fully to the length of the aculeus contained in it. The eversible mem¬ brane is entirely membranous except for the two pairs of lightly sclerotized taeniae (t), arising basally, and sclerotized scales or teeth which often surround the tube and extend more apically than shown in figs. 7 and 9. The shape and distribution of these teeth, especially the large dorsobasal ones (th) found in some Trypetinae, often contain excellent taxonomic characters; they must be examined under a compound microscope. The aculeus, termed the “ovipositor” by many North American work¬ ers, is connected to the apical end of the eversible membrane; it is a sclerotized tube provided with ventral flaps (fl) that cover the genital opening and through which the egg is extruded. Especially in the case of the Anastrepha and Urophora species discussed in this handbook, the details of the shape and ornamentation of the aculeus tip (acu t) are invaluable in identification at the species level, but a compound microscope sometimes must be used to reveal necessary detail. The ovipositors of all North American species have not been studied in detail, but such studies often show relatively constant morphological differences at the species level. Careful anatomical and taxonomic studies also reveal that the spermathecae, visible only after clearing and mounting the specimens, often exhibit differences at the genus and species level. In addition to shape and surface structure, the number of spermathecae is taxonomically significant. Oedicarena have four (Norrbom et al. 1988), whereas other Trypetinae have two or, more commonly, three, and Dacinae and the Tephritinae have two (Hancock 1986b). Normally the aculeus rests within the oviscape, surrounded in turn by the eversible membrane as shown in schematic fig. 8, so it is not usually visible in dried specimens. When ready to oviposit, the female extends her aculeus with her body fluids and inserts the tip into the host plant tissue where the egg can be embedded and pro¬ tected. The procedure for preparing useful slide mounts of the terminalia is detailed in the Techniques section of this handbook. Male (Figs. 10, 11) As in the female, the widest part of the abdomen in dorsal view is the second segment (T2), but in contrast to the female, the male has only five visible segments. In the Tephritidae, the remaining abdominal segments are termed the postabdomen. The individual parts of these hidden segments are largely unrecog¬ nizable without clearing and dissection (fig. 11). The male fifth tergite is markedly

Adult Morphology

Figures 10, 11. Generalized views of male abdomen. 10, Side view. 11, Detail of genitalia. Terms in parentheses are those used by other authors, a a, aedeagal apodeme (fultella); bph, basiphallus (phallotheca); dph, distiphallus (aedeagus, glans, penis, phallosome); ej a, ejaculatory apodeme; ep, epandrium (tergite 9); hyp, hypandrium (genital ring, sternite 9); p a, phallic apodeme; pr, protandrium (syntergosternite 6-8); prg, proctiger; prs, prensisetae; SI—S5, sternites of abdominal segments 1—5; sctl, scutellum; snm, sternum of preabdomen; sp p, sperm pump; sst, surstylus; tgm, tergum of preabdomen; Tl—T5, tergites of abdominal segments 1—5.

longer than the preceding ones and often extends over the postabdomen. Few useful taxonomic characters have yet been discovered in the structure of segments 1—5 of the male abdomen in North American species. Fig. 11 shows a stylized male postabdomen to illustrate the details of its various components. The epandrium (ep) is often visible from posterior view as an “appen¬ dage” of the fifth segment in dried specimens, although more often it is hidden beneath the fifth tergite. It is generally horseshoe-shaped. The outer surstyli (sst), which may be short or long and lobelike, are fused, often indistinguishably, to the epandrium ventrally. A second pair of lobes, the inner surstyli, are closely associated with the surstyli and may be indistinguishable from them without careful examina¬ tion. Each of these bears one or two short, blunt, usually very dark teeth called prensisetae (prs). The proctiger (prg), which bears the anal opening, is soft and varies in shape and size. The aedeagus consists of the distiphallus (dph), borne on a long, curved basiphallus (bph), which in turn is connected to the sperm pump (sp p) and the ejaculatory apodeme (ej a), the latter a muscle attachment for moving sperm to the distiphallus. The aedeagal apodeme (a a) and hypandrium (hyp) assist in the movements of the basiphallus by means of appropriate muscle attachments. This entire assemblage is joined to abdominal segment 5 by a series of narrow, partially fused sclerites (and associated membrane) representing reduced segments 6, 7, and 8. The distiphallus itself is usually a vastly complicated, partly sclerotized organ surrounded by a membranous sheath; it appears to offer valuable taxonomic charac¬ ters when studied properly. Flowever, the convolutions of its internal structure have never been adequately explained, making comparisons of a taxonomic nature diffi-

13

Adult Morphology cult at best. With our present knowledge, comparisons must be made on a side-byside basis with proper rotation, much as in the double microscope techniques used to determine bullet riflings. This technique is made doubly difficult by the springy nature of the basiphallus. In this handbook, we have purposely avoided referring to any but the most easily viewed genitalic structures because most users will be un¬ prepared to deal with the difficulties in dissecting and preparing these minute parts.

GENERAL FEATURES

The setation of the visible abdominal segments in both sexes is little used in North American species, but we suspect that characters, useful for identification but not yet studied extensively, may be present. For instance, Freidberg and Mathis (1986) have shown the value of the color of dorsal abdominal setae in distinguishing among species of Neaspilota. Color characters on the visible abdominal segments may be of considerable taxonomic value, especially in light-colored species in which various dark spots are present. In some of the flower-feeding species, dark stripes and sugges¬ tions of abdominal spots are characters inherent in the tomentum rather than pig¬ ments in the cuticle itself.

BIOLOGY

Information about the biology of the Tephritidae is widely scattered in the literature; recent summaries for the family as a whole are rare. Christenson and Foote (1960) and Bateman (1972) provide reviews of information about general tephritid biology; Wasbauer (1972) has summarized host information for the Tephritidae north of Mexico; and White (1988), although emphasizing the British fauna, provides an excellent overview of this subject. The interested reader will find many review papers on various aspects of tephritid biology in two recent books: Fruit Flies of Economic Importance (Cavalloro 1983), and Fruit Flies: Their Biology, Natural Enemies and Control (Robinson and Hooper 1989). Much valuable information also can be found by consulting the references we have listed and annotated in the synonymy of each taxon. Despite the common name fruit flies, tephritids as a group attack a wide variety of host tissues. Fruit-breeding species may feed on the pulp or the seeds. Other species mine leaves and stems or breed in flowers, young shoots, or buds. Most species in the subfamily Tephritinae form flower, stem, or root galls, or feed in developing flower heads of the plant family Asteraceae. In a few taxa of this group, including some species of Tephritis, different generations have been found to switch from gall forma¬ tion to flowerhead feeding (Goeden 1988, Jenkins and Turner 1989). Species in no fewer than 14 North American genera of Tephritinae in the tribes Dithrycini, Eutretini, Myopitini, and Tephritini form galls on various species of Asteraceae. Certain morphological features appear to be associated with gall forma¬ tion. Adults of gall formers tend to be relatively robust and have a broad frons; a short, broad thorax; a robust and conical oviscape; and often a tendency to become greasy after being killed and pinned. Freidberg (1984) has reviewed gall-making by the Tephritidae in considerable detail on a worldwide basis. The host relationships of fruit flies is one of the most interesting aspects of their biology. G. L. Bush (1966a, 1969a, 1974) and his students have used the genus 14

Biology Rhagoletis, particularly R. pomonella (Walsh) and its host races, as a model of sympatric speciation. Tephritids are also a main component of the European thistle fauna, the subject of extensive resource partitioning studies by Zwolfer and associ¬ ates (Zwolfer 1983, 1988). Many other fruit flies would be fruitful subjects for the evolutionary biologist. Among wasp mimics are Toxotrypana (Landolt 1984a), many Dacinae, and some Anastrepha species. Others, for example some Rhagoletis and Zonosemata species, have wing patterns that simulate, and therefore protect them from, their jumping spider predators (Whitman et al. 1988). Aggression between males and courtship and mating behavior often involve elaborate signaling such as wing waving and other repetitive actions (Burk 1981). Males frequently produce pheromones to attract females, and some species are thought to form aggregations that attract females. Most Dacini males have a stridulatory structure on the wing and abdomen (Hardy 1973), and some Anastrepha males make sounds by vibrating their wings. During courtship in some species such as Paracantha gentilis, there is a “kiss,” in which the male apparently feeds the female a nutritive secretion; the male in other species, including those of Eutreta, Icterica, and Stenopa, makes a froth mass on the host plant to attract the female (Freidberg 1981). Many tephritid females mark fruit in which they have laid eggs with a pheromone to deter oviposition by other females (Averill and Prokopy 1989). The study of tephritid biology also contributes materially to our ability to recog¬ nize species groups and classify them taxonomically and ecologically. Zwolfer (1983) distinguishes two major classes of substrate used by fruit flies. One of these, the pulp of fleshy fruits of a wide variety of host plants, corresponds to a group of tephritids that we often call “trypetines” because most of the forms belonging to the subfamily Trypetinae use this part of the plant for their development; this is also true of the Dacinae. In the temperate regions of the world, most species develop within vegetative structures of the plant other than fleshy fruit. We call those species “tephritines” for the same reason. These terms are used strictly for convenience because the currently recognized subfamilies do not strictly correspond to this sub¬ division of host habitats. The reader is referred to the discussion of classification elsewhere in this handbook for an account of suprageneric categories in the family. The distinctions between “trypetine” and “tephritine” are helpful to the taxono¬ mist in that certain morphological characters also correspond to these two broad biological groups. One of these is the nature of the larva, which in trypetines is maggotlike and well adapted to moving about actively in the flesh of the host fruit. In so doing, it gains continual access to suitable feeding substrate and is able to escape from the fruit at pupation time. On the other hand, larvae of the tephritines spend their lives in a very small area close to where the egg is deposited, such as in a stem or flower ovary, thus do not require great mobility. These larvae usually are ovoid and pupate at the feeding site, having lost some locomotive powers. Furthermore, certain morphological features of the adult appear to be characteris¬ tic for each major host division, although some of the characters overlap and cannot be used alone as diagnostic for the group. Usually the wings of trypetines are hyaline and adorned with dark markings in various shapes and combinations, whereas the wings of tephritines are more often dark with light-colored or hyaline spots and

16

Biology bands. Other morphological characters associated with these two types of wing patterns also are characteristic of the particular group. In the trypetines, the basal cubital cell (fig. 13, c) is usually elongated along the anal vein; the dorsocentral bristles (fig. 50, b) are situated no farther anteriorly than the level of the wing base; and the postocular setae (fig. 61, b) are usually all slender, dark, and sharply pointed. In contrast, the basal cubital cell of tephritines usually has only a very small point (fig. 18, a) or none at all (fig. 17, b); their dorsocentral bristles are usually located anterior to the wing base (fig. 51, b); and at least some of their postocular bristles are usually whitish, short, and blunt. Characters such as these (and others not discussed above) often enable the taxono¬ mist to make rapid preliminary judgments about the taxonomic position of a partic¬ ular specimen and provide some predictive value in forecasting its biological traits in lieu of strictly taxonomic identification through the use of keys, illustrations, and descriptions.

CLASSIFICATION OF THE TEPHRITIDAE

RELATIONSHIPS TO OTHER FAMILIES OF DIPTERA

The following table summarizes the higher classification of Diptera and is presented to demonstrate the taxonomic position of the Tephritidae relative to other flies. It roughly reflects the evolutionary classification of the order, with the most primitive taxa listed first and the succeeding ones reflecting a successively higher degree of development. This arrangement corresponds to that of Hennig (1973) as modified somewhat by Griffiths (1972), Steyskal (1974), and McAlpine et al. (1981b). The last-named publication and McAlpine (1981b, 1989) present this information in considerable detail. As shown in this outline, the Tephritidae belong to the subgroup Acalyptratae of the infraorder Muscomorpha. Suborder Nematocera

Suborder Brachycera Infraorder Tabanomorpha Infraorder Asilomorpha Infraorder Muscomorpha Aschiza Schizophora Acalyptratae Calyptratae

17

7 infraorders and 16 superfamilies, including mosquitoes, midges, gnats, crane flies, black flies 2 superfamilies, including horse and deer flies and relatives 3 superfamilies, including robber flies and relatives 3 superfamilies, including flower flies and relatives 9 superfamilies, including many diverse forms and Tephritidae 3 superfamilies, including the house fly, blow flies, bot and warble flies

18

Classification of the Tephritidae The classification of the acalyptrates has a long and turbulent history, for this group, which is questionably monophyletic, presents the most confusing and difficult problems within the Diptera. In many cases, the differentiation of its member taxa is not always clear, individuals are often small and difficult to identify, and the number of distinguishable suprageneric categories is large. The tephritids share many taxonomic characters with other acalyptrate flies, espe¬ cially the other families of the superfamily Tephritoidea, many of which also have spotted or striped wings. Tephritids most closely resemble the Otitidae, Pyrgotidae, Richardiidae, and Platystomatidae in general appearance, but according to Griffiths (1972), they are most closely related to the Platystomatidae, Tachiniscidae, and Pyrgotidae mainly on the basis of their similar male genitalia. McAlpine (1989) considers the latter two families the tephritids’ closest relatives. Fortunately, almost all of the world’s tephritids may be distinguished from members of any other acalyp¬ trate family by the nature of the subcostal vein (fig. 6, Sc). This vein, situated in the basal half of the wing immediately behind the anterior margin, parallels the wing margin, then bends forward abruptly, often forming a minute enlargement at the bend. The subcosta usually becomes weaker between the bend and the anterior wing margin (i.e., the costa), sometimes disappearing completely at its junction with that vein. This character is in distinct contrast to that of all other acalyptrates, in which the subcosta is either complete (not weak apically), markedly incomplete along part of its length, curved gradually forward before joining the costa, or joined to other veins basad of that point.

THE U.S. AND CANADIAN GENERA Historically, there has been little universal agreement among authorities as to an arrangement of genera that reflects the true evolutionary development of the Tephritidae. Among the earliest of the students of this family to propose a su¬ prageneric classification, Loew (1862b) suggested two main groups, the Dacina and the Trypetina, primarily on the basis of a study of the Palearctic fauna. Later, he placed all of the endemic North American genera into his Trypetina without any subdivisions (Loew 1873). Among the first to demonstrate subdivisions within the family was Wulp (1899-1900), who segregated the Neotropical genera in a key without applying group names. The first grouping of the nondacine genera is be¬ lieved to be that of Bezzi (1910), who later revised his own ideas (Bezzi 1913, 1924a, 1924b). Cresson (1914a, 1914b), in tentatively assigning the North American genera to the two subfamilies and three tribes suggested by Bezzi (1913), was the first American to concern himself with the possible phylogeny of New World tephritids. Hendel (1927) based his review of the Palearctic fruit flies on Bezzi’s concepts but further divided the Dacinae, Trypetinae, and Tephritinae in an attempt to express morphological relationships in more detail. Additional divisions were subsequently proposed by authors who studied the faunas of significant geographical areas or revised certain groups of genera. Most prominent among them were Shiraki (1933) for Japan; Zia (1937) and Zia and Chen (1938) for China and North China, respec-

19

Classification of the Tephritidae

tively; Hering (1941a, 1941b) for Peru and East Africa, respectively; Munro (1947) for Africa; and Collin (1947) for Britain. An ever-increasing complexity in the taxonomy of the family, resulting from the increase in knowledge of its component parts and the overall absence of attempts at correlating many of these newer concepts, opened the way for Hering’s (1947b) synthesis of the advances made to that date. In the form of a key, Hering presented an arrangement of subfamilies and tribes for the fruit flies of the world. His close acquaintance with the world literature establishes this work as a landmark summary of the progress made to that date, modified by him on the basis of his own familiarity with the world fauna. It is interesting that, after an intervening 46 years of work on this aspect of fly taxonomy, most of Hering’s concepts are being used today in one way or another to designate suprageneric taxa within the family, although some of his categories have been redefined and a few new ones have been devised. The status and interrelationships of most of the currently used tephritid su¬ prageneric categories continue to be revised as a result of current studies. Most notable among the many post-Hering works dealing with this subject are those of the following authors: Aczel (1949, 1952a), Hardy (1955, 1973, 1974, 1977, 1980, 1983, 1986a, 1986b, 1987, 1988a, 1988b), Drew (1972), Foote (1980), Cogan and Munro (1983), Ito (1983-1985), Munro (1984), Hancock (1984, 1985a, 1895b, 1986a, 1986b, 1990), and Hardy and Foote (1989). The problem of achieving universal agreement among fruit fly taxonomists is yet to be resolved. Most reviews presented since 1947 have been based on studies within a particular geographical region or on a particular fly taxon, as is readily seen in reviewing the titles of those works referenced above. To be completely fair to the authors of those studies, we should note that their knowledge of fruit flies is very probably more widely based than the particular geographical or zoological area in which they are acknowledged specialists. However, many factors have prevented them from conducting studies on a worldwide basis or on flies belonging to other, related taxa, to attain a universal understanding of across-the-board relationships. Until recently, there has also been little use of rigorous phylogenetic methods, and most subfamilies and tribes have not been demonstrated to be monophyletic. An¬ other factor contributing to this unstable situation is the absence of broadly based biological studies that would otherwise affect conclusions so far based on mor¬ phological studies. The phylogeny of the Tephritidae is intimately interwoven with the phylogeny and distribution of its host plants, yet few studies demonstrating the effects of this connection are available, nor do they include studies of genetics, physiology, or ecology that undoubtedly apply to these problems. Part of the problem also lies with the flies themselves. Not surprising for a rela¬ tively recent, rapidly radiating group such as the Tephritidae, homoplasy (convergent evolution) appears to be common in many morphological characters that have been the main basis of classification. External characters such as chaetotaxy and body and wing color patterns, which are extremely useful at the species level, appear to be especially subject to convergence. The growing body of biological data, especially concerning host plants, mode of feeding, and recent studies incorporating analyses of male and female genitalic structures, have led to improvements in the classification

20

Classification of the Tephritidae

(e.g., Freidberg 1985; Korneyev 1985, 1989; Hancock 1986b; Norrbom 1987; White and Korneyev 1989), but until a comprehensive generic analysis based on characters of many kinds has been achieved, the relationships of many taxa will probably remain uncertain. North America, or at least that part of it north of Mexico, is among the best known major areas at the species level for the Tephritidae. Nonetheless, relationships among the Nearctic genera have never been discussed comprehensively. Some genera that also occur in the Neotropical Region were treated in the classification of Foote (1980), which forms the major basis for this discussion. With the reader thus forewarned, we present a classification of the Nearctic fruit fly genera. Our mam intent is not to define higher level taxa on a worldwide basis or to justify their rankings as given here, but to summarize what is known about the phylogenetic relationships of our fauna. Where possible, we discuss hypotheses of monophyly; that is, we suggest what taxa are most closely related to each Nearctic genus. Wfe hope that pointing out relationships among groups of genera, however small, will eventually lead to the resolution of relationships at higher levels.

A CLASSIFICATION OF THE GENERA OCCURRING IN AMERICA NORTH OF MEXICO Family Tephritidae Newman Subfamily Dacinae Loew Tribe Dacini Bactrocera Macquart Dacus Fabricius Tribe Ceratitini Bezzi Ceratitis Macleay Subfamily Trypetinae Loew Tribe Euphrantini Hering Epocbra Loew Tribe Toxotrypanini Munro Anastrepha Schiner Toxotrypana Gerstaecker Tribe Trypetini Subtribe Carpomyina Norrbom Rhagoletis Loew Rhagoletotrypeta Aczel Zonosemata Benjamin Subtribe Trypetina Euleia Walker Strauzia Robineau-Desvoidy Trypeta Meigen Unplaced genera of Trypetini Cbetostoma Rondani Myoleja Rondani Oedicarena Loew Raraterellia Foote Unplaced genus of Trypetinae Hexacbaeta Loew

21

Classification of the Tephritidae

Subfamily Tephritinae Tribe Terelliini Hendel Chaetostomella Hendel Neaspilota Osten Sacken Orellia Robineau-Desvoidy Tribe Myopitini Bezzi Rbynencina Johnson Urophora Robineau-Desvoidy Tribe Acrotaeniini, new tribe Acrotaenia Loew Tomoplagia Coquillett Tribe Dithrycini Hendel Subtribe Cecidocharina Hering Callacbna Aldrich Cecidocbarella Hendel Procecidocbares Hendel Procecidocbaroides Foote Stenopa Loew Subtribe Eurostina, new subtribe Aciurina Curran Eurosta Loew Valentibulla Foote and Blanc Unplaced genus of Dithrycini Peronyma Loew Tribe Eutretini Munro Acidogona Loew Cryptotreta Blanc and Foote Eutreta Loew Jamesomyia Quisenberry Metatepbritis Foote Paracantba Coquillett Xantbomyia Phillips Xenocbaeta Snow Tribe Tephritini Acinia Robineau-Desvoidy Dioxyna Frey Dyseuaresta Hendel Euaresta Loew Euarestoides Benjamin Gonioxyna Hendel Neotepbritis Hendel Oxyna Robineau-Desvoidy Paroxyna Hendel Tepbritis Latreille Trupanea Schrank Unplaced genera of Tephritinae Gymnocarena Hering Icterica Loew Mylogymnocarena Foote Xantbaciura Hendel

22

Classification of the Tephritidae

Subfamily Dacinae Because of its economic importance, large number of species, and distinctiveness, most systematists have ranked what is here called the tribe Dacini as a separate subfamily. The Ceratitini have been included in the Trypetinae in most classifications, but we follow Hancock (1986a), who suggested their closer relationship with the Dacini. Hypothesized synapomorphies include the ridge across the large caudal tubercles below the hind spiracles in the larva and the shape of the surstyli, which usually are elongate with an anterior and posterior lobe. Hancock (1986a) also considered the loss of one of the spermathecae an autapomorphy for the Dacinae or a synapomorphy for it and the Tephritinae, which also have only two. However, this loss has occurred a number of times in the Trypetini (e.g., within the genus Rbagoletis). He also noted that in many dacines and ceratitines, the lobe of cell cup is narrower at the base than farther apically. No members of this subfamily are native to the New World, although Ceratitis capitata has been introduced to many parts of Central and South America, and one species of Bactrocera has become established in Surinam. None is currently estab¬ lished in America north of Mexico, although species of Dacus, Bactrocera, and Ceratitis are frequently intercepted and infestations have occurred. The Dacini and Ceratitini breed in fruit, seed pods, or flower buds.

Subfamily Trypetinae

The Trypetinae is the most diverse subfamily of Tephriti¬

dae. It is recognized on the basis of plesiomorphic (primitive) characters, and its monophyly is uncertain; some taxa now included might be more closely related to the Dacinae or Tephritinae than to other trypetines. The group is especially diverse in the Old World tropics, which may be the center of origin of the family. Various subtaxa have been proposed, some of which have been ranked as separate sub¬ families, but until relationships among them are clarified, arguments about their ranking are meaningless.

Tribe Euphrantini

The tribe Euphrantini is restricted to the Paleotropics except

for Eupbranta Loew, which is widespread in the Palearctic Region, and Epocbra, the only New World representative (A. Freidberg, personal communication). The group is characterized mainly by the presence of long, fine setae on the anatergite and the absence of a sclerotized area below the base of the abdomen (Hardy 1983). This group is poorly separated from the tribe Adramini and likely will be combined with it eventually. Epocbra, like most euphrantine genera, lacks presutural supra-alar and ocellar bristles. It possesses acrostichal and three frontal bristles as does Eupbranta (Rbacocblaena Loew), which appears to be the most closely related taxon. Only one species has been known from North America, but a second, undescribed species occurs in Mexico. Both breed in fruit of Ribes species.

Tribe Toxotrypanini The tribe Toxotrypanini includes only the genera Anastrepba and Toxotrypana and is the only trypetine tribe restricted to the New World (Hancock 1986b). It is mainly Neotropical; species in these two genera occur only in southern Texas and Florida in our area. Anastrepba species breed mainly in fruit or

23

Classification of the Tephritidae

seed pods of a wide range of plant families (Norrbom and Kim 1988b). Toxotrypana species also breed in fruit or seed pods but appear to be specialized on latex-bearing plants including Caricaceae, Asclepiadaceae, and Apocynaceae. These two tephritid genera share a number of unique derived characters (synapomorphies), including a T-shaped apical sclerite on the distiphallus, lateral flangelike lobes on the base of the oviscape, and the eversible membrane with an expanded basal area bearing enlarged scales dorsally (Norrbom 1985). Immunological data for A. suspensa and T. curvicauda also support this relationship (Kitto 1983, Sarma et al. 1987). Although the monophyly of these two genera is well supported, how they are related to other fruit fly taxa is not clear. Tribe Trypetini The tribe Trypetini is not well delimited; it is recognized by the lack of derived characters of the other tribes. Nevertheless, there are groups of clearly related genera within it. Rhagoletis, Rhagoletotrypeta, and Zonosemata belong to the subtribe Carpomyina, a mainly Holarctic and Neotropical group of fruit¬ breeding species. These are characterized by an area of weak sclerotization at the apex of the oviscape in the female (Norrbom 1990) and stomal guards, sclerotized projections lateral to the mandibles on the larvae (L. E. Carroll, personal communi¬ cation). Oedicarena and Paraterellia are perhaps related to this group (Berlocher and Bush 1982, Norrbom et al. 1988). Trypeta and Euleia (both widespread in the Old World) and probably Strauzia (endemic to the Nearctic Region) belong to the subtribe Trypetina, a mainly Old World group of genera in which the larvae mine leaves (in the case of Strauzia, stems) of species of Asteraceae or Apiaceae (Hancock 1986b, White 1988). At least Acidia Robineau-Desvoidy, Aciuropsis Hardy, Aischrocrania Hendel, Cryptaciura Hendel, Hoplandromyia Bezzi, Matsumuracidia Ito, and Vidalia Robineau-Desvoidy also appear to belong to the Trypetina, although the exact limits of the group have not been determined, and a number of genera or species that mine stems or leaves in other plant families should perhaps be included. In the three Nearctic genera, as in the Palearctic taxa treated by White (1988), the aculeus tip is broad, flat, and finely serrate, and the distiphallus has minute scalelike sculpturing in two areas. The cuticle often has tiny black dots, especially on the thorax. The puparia of Trypeta angustigena and the Mexican species T. concolor (Wulp) have the middle opening of the hind spiracle projecting spinelike laterally, similar to that shown in White (1988, fig. 220), who stated that this occurs in certain Palearctic leaf-mining species (genus not specified). This spinelike projection does not occur in Euleia fratria or the various Strauzia species we examined, which have more or less typical hind spiracles. The subtribe Trypetina is being revised by H.-Y. Han, currently a doctoral student at Pennsylvania State University. The relationships of Cbetostoma and Myoleja are not well understood, but they and the Neotropical genus Parastenopa Hendel are probably part of another mainly Old World group that is possibly related to the Trypetina. Cbetostoma is mainly Holarctic, with a few species entering the Oriental Region. Myoleja is probably the most widespread trypetine genus because it occurs in every zoogeographic region except Antarctica (Foote 1980, Hancock 1986b, Hardy 1987), but it may be poly-

24

Classification of the Tephritidae

phyletic. White (1988) indicated some Old World species that mine leaves may belong in the group of leaf-mining genera discussed above, but none of the six Nearctic species of Myoleja and Chetostoma appears to belong in the latter group. Only M. limata, which breeds in fruit of various Ilex species (Aquifoliaceae), has known hosts, but in the other species (except possibly M. unifasciata, of which the female is unstudied), the tip of the aculeus is flattened in the sagittal plane, indicating frugivory. Two Neotropical species in this group are also associated with Ilex: Anomoia ogloblini (Blanchard) mines young shoots of I. paraguaiensis, and Parastenopa elegans (Blanchard) mines psyllid leaf galls on the same plant. A third species, P. marcetiae (Bezzi and Tavares), was reared from galls, probably of a cecidomyiid, on Marcetia (Melastomataceae) (Blanchard 1929, Aczel 1955c). The shape of the aculeus tip varies in the Neotropical species but is never broad as in Trypeta, Euleia, or Strauzia. The distiphallus in all New World species we examined has fine sculpturing. Hexachaeta is a mainly Neotropical genus that, within our area, has been collected only from southern Texas. It is poorly known biologically, although a few species have been reared from the fruit or twigs of Moraceae and Verbenaceae. The classifi¬ cation of Hexachaeta is uncertain, as is that of Blepharoneura Loew and Ceratodacus Hendel, the other New World trypetine genera with three pairs of scutellar bristles. Hancock (1986b) excluded them from the Acanthonevrini, most genera of which also have three pairs of scutellar bristles, because they have well-developed ocellar bristles (actually these are often weak in Hexachaeta), and the subapical sensilla on the aculeus are not long and setalike. These three genera, unlike most other New World tephritids, do resemble certain Old World taxa in their male genitalia: the lateral sclerites are reduced or absent (or completely fused to the hypandrium), and the arms of the aedeagal apodeme articulate with the hypandrium near its apex and are usually fused basally. The fusion of the arms of the aedeagal apodeme is common in the Acanthonevrini, Euphrantini, and Phytalmiini, although Hardy (1986b) ques¬ tioned its phylogenetic significance. As these are mainly Paleotropical groups, how¬ ever, the taxa most closely related to Hexachaeta are probably from that region. The genera most closely related to Blepharoneura are the Afrotropical Baryglossa Bezzi and the Oriental Hexaptilona Hering (Munro 1957). Subfamily Tephritinae The subfamily Tephritinae is probably best defined by its biology. Except for the Old World tribe Tephrellini (= Aciurini, see Hancock 1990), most species in this group breed in flower heads or form flower, stem, or root galls in species of Asteraceae. This is true of all the New World species except at least two Neotropical species of Eutreta that form galls in Verbenaceae. Some tribes of the Tephritinae, such as the Tephrellini (= Aciurini), Oedaspidini, Myopitini, and Terelliini, often have been ranked as separate subfamilies. Because relationships among them are unresolved, we prefer to include all of them in one subfamily. Defining the Tephritinae in terms of synapomorphies has been an elusive goal. Many tephritine species, including most of those treated in this handbook, have certain head bristles (especially the postoculars or the posterior orbital) or other bristles or setulae (especially on the scutum) that are broad and whitish, but in some

25

Classification of the Tephritidae

genera such as those in the Myopitini and Tephrellini (= Aciurini), all setae are slender and unicolorous as in the Trypetinae. Other species such as those in the Terelliini and Acrotaeniini have intermediate (i.e., slightly swollen) postoculars. Per¬ haps the best chaetotaxic character to separate the subfamilies is the scapular bris¬ tles. In most Trypetinae, they are well differentiated and are located consistently in the same spot. Except for some Tephritini (Freidberg 1987), some Schistopterini, and some Myopitini, species of Tephritinae usually lack scapular bristles. Sometimes larger setae occur near the anterior margin of the scutum that may be scapulars, but if present, they are variable in size, number, and location within species. The reduc¬ tion or loss of the scapular bristles may be a synapomorphy for the Tephritinae, but it is a subtle character, difficult to use for identification purposes. The bodies of most Tephritinae are also densely microtrichose, giving them a matte grayish appearance. These microtrichia also obscure the internal suture in the middle of the anepisternum (mesopleural suture of Hancock 1986a), but its visibility is also affected by cuticle color and is not consistent across the subfamily. Many Tephrellini (= Aciurini), Myopitini, and some genera of other tribes are relatively sparsely microtrichose, whereas some Trypetinae and Ceratitini are moderately microtrichose. Hancock (1986b) noted that all Tephritinae, like the Dacinae, have only two spermathecae. Many taxa within the Trypetinae also have only two, so it is either a synapomorphy at a higher level, or there is considerable homoplasy within this character. Another perhaps significant character is the shape of the epandrium and surstyli. There is considerable variation within the Tephritinae, but in all tribes there are at least some taxa with a typical shape that does not occur in the Trypetinae or Dacinae. In this type, the epandrium and surstyli are more or less oval in posterior view. The margin of the surstylus often projects posteriorly as a dorsal lobe (see Jenkins and Turner 1989, fig. 1, or Munro 1957, figs. 160—192). This type of genitalia may be the ground plan condition for the subfamily. The type of sex-determining method also appears to have phylogenetic significance in the Tephritinae. Whereas all Trypetinae and Dacini that have been studied are male heterogametic, as are most Diptera, the investigated species of Cecidocharina (Cecidocharella, Procecidochares), Eutretini (Rhachiptera Bigot), and Tephritini (.Acinia, Chrysotrypanea Malloch, Tephritis) have a heterogametic female, or in the case of some Tephritini (Trupanea, Trypanaresta Hering), both male and female sex chromosomes are isomorphic (Bush 1966b; Bush and Taylor 1969; Bush and Huettel 1970; Frias 1981, 1985, 1988). An exception to this rule may be Tephritis arnicae (L.), which is reportedly XO in the male (Bush 1966b). Female heterogamety may be a synapomorphy for the subfamily with a subsequent change to isomorphic sex chromosomes in some Tephritini, but until species from other tribes have been sampled, the level at which this character is a synapomorphy remains uncertain. Considerable further research is needed to determine the relationships among the higher groups of Tephritinae. Tribe Terelliini The tribe Terelliini is a well-defined group. Freidberg (1985) listed the lyre-shaped pattern on the scutum and the inclinate posterior pair of orbital bristles as synapomorphies. The New World genera are Cbaetostomella, Neaspilota

26

Classification of the Tephritidae

(which is endemic), and Orellia, and plans are being formed to release species of Cbaetorellia Hendel and Terellia Robineau-Desvoidy in North America for the bio¬ logical control of weeds (Groppe and Marquardt 1989a, 1989b). The New World species of Orellia have been transferred recently to Terellia, but this change came to our attention too late to modify the taxonomic arrangement of this book. Tribe Myopitini

The tribe Myopitini is defined by the following synapomor-

phies: (1) posterior orbital bristles absent (this character occurs independently in several other taxa); (2) cell cup without a posterior lobe (i.e., vein CuA2 convex or straight); and (3) abdominal pleura darker than terga or sterna, usually with a granular appearance (especially obvious when the abdomen is dissected and cleared). In all but a few species (none from the New World), the spermathecae are weakly sclerotized. The myopitines have retained the plesiomorphic type of postocular bris¬ tles (slender and dark) and for that reason were often classified in the Trypetinae. Freidberg and Norrbom (in preparation) are currently revising the generic classifi¬ cation of this group, and White and Korneyev (1989) and Korneyev and White (1991) are revising the Palearctic fauna. These revisions will result in the breakup of the large genus Uropbora, which currently contains the large majority of species in the tribe, including all of the New World species except one. Of the species occurring in North America, only the introduced Palearctic species, which all breed in plants of the tribe Cardueae (White and Clement 1987), actually belong in JJropbora. Except for U. claripennis, which is part of a mostly Neotropical group, the native species of the western United States, which all breed in plants of the subtribe Solidagininae, tribe Astereae (Goeden 1987a), form a natural group. Rbynencina is currently monotypic, but several Neotropical species placed in Uropbora by Steyskal (1979) belong there. This genus appears to be most closely related to the Old World genera Myopites Blot and Stamnopbora Munro. Tribe Acrotaeniini

The tribe Acrotaeniini is proposed to include Acrotaenia,

Tomoplagia, and the following Neotropical genera: Acrotaeniacantba Hering, Baryplegma Wulp (= Pseudacrotaenia Hendel, new synonymy), Caenoriata Foote, Euarestopsis Hering, Neotaracia Foote, Polionota Wulp, Pseudopolionota Lima, and Tetreuaresta Hendel. The differences between the type species of Baryplegma and Pseudacrotaenia, limited mainly to spot shape and the predominant color of the wing pattern, are not significant enough to maintain them in separate genera. Diagnostic characters of this tribe include: head usually not quadrate, with angle between face and frons obtuse or rounded (close to 90° in some Acrotaenia, Polionota, and Tetreuaresta); three or more pairs of frontal bristles; postocular setae sometimes mixed large and small but unicolorous yellowish (except small ones black in Acro¬ taeniacantbar); scutellum with two pairs of bristles; vein R4+5 densely setose dorsally to beyond apex of R2+3; cell cup with length of posteroapical lobe two-thirds or more apical width of main part of cell; epandrium and surstyli oval, outer surstylus without dorsal lobe (see Norrbom 1987, fig. 2); distiphallus with endophallus vari¬ able in shape but always somewhat tubelike and exposed or only partially enclosed by a dorsal sclerite (see Norrbom 1987, fig. 5); and the aculeus tip straight or

27

Classification of the Tephritidae

dorsally curved. Except for the distiphallus, the characters listed above are probably plesiomorphic for the Tephritinae. Most of them are variable within the other tribes; the Acrotaeniini are therefore remarkable in that all of those characters are constant. Foote (1980) included many of these genera in the tribe Platensinini, an otherwise Paleotropical group whose species breed in Acanthaceae (Hancock 1986a). We have examined only six genera of Platensinini, and although they resemble the Acrotae¬ niini in head shape and the number of frontal and scutellar bristles, all differ in having mixed (large white and small dark) postocular setae, a shorter lobe on cell cup, and a different type of distiphallus. Vein R4+5 often is more sparsely setulose, and some species have a parafacial spot. Most genera have an elongate, very slender, subapically expanded and/or ventrally curved aculeus tip. These differences, especial¬ ly in the genitalia, suggest that the Acrotaeniini at least is a separate monophyletic group; they cast doubt about how closely related it is to the Platensinini. Hancock (1986a) suggested instead that the Acrotaeniini may be related to Hendrella Munro and related genera (= Tephrellini of Hancock 1986a; see Hancock 1990), which he distinguished from the Tephritini mainly by their predomi¬ nantly dark wing pattern and lightly microtrichose abdomen with only fine, dark setulae. This diagnosis does not work well for the New World Tephritini, especially the extent of abdominal microtrichia. Some genera (e.g., Euaresta) also vary in the color and shape of the abdominal setulae, although in the Acrotaeniini they are consistently slender. We have seen only a few species of the Afrotropical genera Hancock (1986a) included in this group. Spatbulina Rondani in most characters seems like a typical Tephritini. Parafreutreta regalis Munro resembles the Acrotae¬ niini in chaetotaxy (except vein R4 + 5 is more sparsely setose) and in head shape, but the lobe of cell cup is smaller, there is a strong dorsal lobe on the outer surstylus, and the distiphallus is more complex. Like the Platensinini, it does not appear especially closely related to the Acrotaeniini. From the limited host data available, a fairly broad range of Asteraceae and several parts of the plants appear to be used by Acrotaeniini. Two species of Acrotaenia breed in flowers of Heliantheae; one Polionota forms galls on an unknown plant; species of Tomoplagia have been reared from flowers or galls on species of Liabeae, Mutisieae, and Vernonieae (records from non-Asteraceae, fleshy fruits are doubtful); and Tetraeuaresta obscuripennis (Loew) has been introduced to Hawaii and other Pacific islands as a biological control agent for Elepbantopus (Vernonieae). Tribe Dithrycini The New World taxa we place here have usually been classified in the Oedaspidini, but Dithrycini (= Ditrichini) has priority over that name and Cecidocharini. The relationships among Oedaspis Loew and related genera (i.e., subtribe Oedaspidina) and the New World subtribe Cecidocharina are not well established, but the Palearctic Ditbryca Rondani appears to be more closely related to them than to the New World genera that are placed here in the Eutretini but previously were classified in the Dithrycini. The only characters from the diagnosis of Hendel (1927) or the key of Hering (1947b) that differentiate Ditbryca (or at least its type species guttularis (Meigen)) from typical Oedaspidina or Cecidocharina are its reticulate, nonbanded wing pattern (a few Afrotropical Oedaspis also have reticulate

28

Classification of the Tephritidae

wings [A. Freidberg, personal communication]) and the more strongly pointed anten¬ nal flagellomere. Like them, Ditbryca is a gall former (on species of Anthemideae). It has a swollen scutellum, broad frons, short antenna, and a basally swollen oviscape, characters that are common in gall-forming tephritids (Freidberg 1984). Flowever, it also has two pairs of dorsocentral bristles as do many Oedaspidina and Cecidocharina and often has clusters of white setulae on the scutum as do many Cecidocharina. The rounded shape of the scutellum also resembles that of most Cecidocharina. None of the latter three characters occurs in the Eutretini, except for the swollen scutellum in some species of the Noeeta Robineau-Desvoidy group. Foote (1980) relied mainly on the presence of a parafacial spot and both dark and white postocu¬ lar setae to group the Neotropical genera here placed in the Eutretini with Ditbryca. The latter character is common in the Platensinini (e.g., Platensina), Schistopterini, and Tephritini (e.g., Paroxyna, Tepbritis) and occurs in some Cecidocharina (Dracontomyia, Stenopa, some Cecidocbares) and Eurostina (Eurosta); it is thus either derived independently many times or is a synapomorphy at a higher level. The parafacial spot appears to be highly subject to homoplasy; for example, it is present in only some species of Dyseuaresta and absent in a few species of Eutreta; it is also present in some Platensinini. Although not present in Oedaspidina or Cecidocharina, a strong parafacial spot is present in Peronyma and a faint one occurs in some species of Aciurina and Eurosta. A careful analysis of male terminalia might help to clarify the relationships of these taxa. We have examined few species, but the shapes of the epandrium and surstyli are at least superficially similar in Ditbryca, several species of Oedaspis, and various species of Cecidocharina. Whatever their higher relationships, the New World genera comprise several prob¬ ably monophyletic subgroups. The first, the subtribe Cecidocharina, comprises Callacbna, Cecidocbarella, Procecidocbares, Procecidocbaroides, Stenopa, and the Neotropical Cecidocbares Bezzi, Dracontomyia Becker, Hetscbkomyia Hendel, and Neorbagoletis Hendel. Almost all species whose biology is known are stem or flower gall formers on Astereae, Heliantheae, Eupatorieae, or Senecioneae, although Cecidocbarella borricbia Bush and Huettel, Procecidocbares penelope (Osten Sacken), and at least one species of Cecidocbares feed in flowers without making a gall (Bush and Huettel 1970; A. L. Norrbom, unpublished data). In species of these genera, usually the mesonotal setulae occur in distinctive patterns, often in clusters on the scutum posteriorly or on the scutellum. Except in Dracontomyia and Stenopa, there are usually postsutural lateral areas without setulae and often without microtrichia. Usually (less so in Stenopa and Procecidocbaroides), the scutellum is broadly swollen and more spherical than in other Dithrycini we have seen, except Ditbryca. The wing pattern divides the subtribe into two groups, one of which is proba¬ bly paraphyletic: Callacbna, Cecidocbares, Hetscbkomyia, Procecidocbares, and Procecidocbaroides with one dark band crossing both veins r-m and dm-cu; and Cecidocbarella, Stenopa, Dracontomyia, and Neorbagoletis with those crossveins separated by a hyaline band. Callacbna gibba (Loew) is probably a species of Pro¬ cecidocbares with several distinctive autapomorphic characters; although it usually has supernumerary anterior orbital bristles, the posterior orbital bristle is absent or

29

Classification of the Tephritidae

minute as in Procecidochares. Stenopa probably is most closely related to Dracontomyia. The Neotropical Ostracocoelia Giglio-Tos, Gerrboceras Hering, and Pyrgotoides Curran, at least the first two of which are gall formers, probably also belong in the Cecidocharina. Although their chaetotaxy (all bristles slender, dorsocentral bristles posterior to supra-alar bristles) might suggest that they belong to the Trypetinae, their genitalia and biology indicate that they are certainly Tephritinae. Ostracocoelia has a wing pattern similar to those of Cecidocharella, Dracontomyia, Neorhagoletis, and Stenopa. Another monophyletic subgroup, the Eurostina, includes Eurosta, Aciurina, and Valentibulla, which form galls mainly on species of Astereae, the former exclusively on Solidago. Their monophyly is indicated mainly by the shape of their elongate surstyli (Norrbom 1985, fig. 1). Peronyma is tentatively included in the Dithrycini because of its biology and swollen scutellum. The single species forms stem galls on a species of Astereae. Further study is required to understand its relationships. Tribe Eutretini Among the taxa placed here, several probably monophyletic groups can be recognized, but whether they are closely related to each other is uncertain. Most of them were previously classified in the Dithrycini, but the type genus of that tribe appears to be more closely related to genera in the Oedaspidina and Cecidocharina (see above). With the exception of a few species of Eutreta (Uncaculeus) that lack a parafacial spot, the group includes those genera having both a parafacial spot and mixed black and white postocular setae but lacking derived features of the Dithrycini or Schistoptenni. The tribe may then be paraphyletic, or because (as noted above) both the parafacial spot and postocular setae appear highly subject to homoplasy, it may be polyphyletic. The tribe Schistopterini comprises a few endemic Old World genera (e.g., Scbistopterum Becker, Eutretosoma Hendel, and Rbabdocbaeta de Meijere) which often have preocellar bristles, a distinctive pattern of wing bullae, and/or an incision in the costal margin of the wing at the apex of the subcostal vein (Hancock 1986a). Several characters occur sporadically within it and the Eutretini that may indicate the relationship of at least some of the genera. For example, in some Schistopterini, in the Noeeta group, in an undescribed species probably belonging in Dictyotrypeta Hendel, and in Paracantba, Laksyetsa Foote, and Neorbabdocbaeta Malloch, there are erect scutellar setae in addition to the marginal bristles. Their number and location vary, however, suggesting that they may be the result of convergence. Para¬ cantba and Neorbabdocbaeta have a pattern of wing bullae similar to that in many Schistopterini, Neorbabdocbaeta and Laksyetsa have a medial facial ridge as do many schistopterines, and Neorbabdocbaeta has a costal incision and extremely acute antennae. The fact that Paracantba, Laksyetsa, and Neorbabdocbaeta share synapomorphies with other Neotropical genera that lack the above characters indi¬ cates that the irregular distribution of these characters within the Eutretini and Schistopterini involves considerable homoplasy. Inferences about phylogenetic rela-

30

Classification of the Tephritidae

tionships (i.e., whether these characters evolved through convergence or are synapomorphies at a higher level that were lost in some taxa) are thus difficult to make at this time. A little-investigated character that may have phylogenetic significance for this group and the Dithrycini is eye color pattern, which normally is visible only in living or freshly killed specimens. In dried specimens, it usually fades within a few months or years but sometimes remains visible. In most Tephritinae (e.g., Tomoplagia in the Acrotaeniini, Rhynencina and Urophora in the Myopitini, Euaresta and Euarestoides in the Tephritini, Neaspilota and Orellia in the Terelliini), the eyes are entirely bright green or reddish with a green sheen which changes depending on the angle of view (Steyskal 1957; A. L. Norrbom, unpublished data). The species of Cecidocharina, Eutretini, and Schistopterini in which eye color has been seen in fresh specimens have eyes with various types of red and green patterns. Procecidochares atra and two species of Cecidochares have a single, broad, vertical green band (Steyskal 1957; unpublished data), whereas in the four species of Eutreta and three species of Dictyotrypeta for which eye color has been observed, there are three narrow horizontal green stripes or a single narrow, medial green stripe, respectively, with the rest of the eye red. Some Schistopterini also have patterned eyes, including a species of Eutretosoma that has a striped pattern similar to that in Eutreta (A. Freidberg, personal communication). Five species of Paracantba, the single species of Laksyetsa, and two species of Rbachiptera Bigot that have been seen fresh have eyes that are mostly green with two rows of large spots. Unfortunately, eye color has not been studied in many genera, but the presence of a pattern or certain type of pattern would seem to be phylogenetically significant. As noted above, the Paracantba group is most similar to the Schistopterini. It includes Paracantba and the Neotropical genera Laksyetsa, Neorbabdocbaeta, Strobelia Rondani, and Rbacbiptera (Foote 1980; Norrbom, in preparation). As noted above, all species of these genera of which we have seen fresh material have a distinct eye pattern of two rows of large spots. They also have the face and frons shiny yellow (without microtrichia) with a similar pattern of black spots. Rbacbiptera and Strobelia lack erect scutellar setae, but the first three genera have three pairs of large, erect setae on the disk rather than on the margin of the scutellum. They also have two or three pairs of orbital bristles, all of which are white, further indicating their close relationship. Species of Paracantba and Laksyetsa breed in flower heads of Heliantheae or Cynareae; those of Rbacbiptera and Strobelia form galls on Baccbaris and other Astereae. Eutreta, Metatepbritis, and the Neotropical Pseudeutreta Hendel probably form another monophyletic group. All three have an apical or subapical hyaline crescent on the wing (lacking only in the males of two Eutreta species). Erect scutellar bristles are lacking. All the species of Eutreta of which fresh specimens have been seen (all in the subgenus Eutreta) have three stripes on the eye as in the schistopterine Eu¬ tretosoma. All species that have been reared are gall formers. Species of Metatepbritis and Eutreta (Uncaculeus) breed in Artemisia (Anthemideae). Other Eutreta species breed in a wide variety of Asteraceae, or in the case of two Neotropical species, in several genera of Verbenaceae, the only non-Asteraceae host of any New World

31

Classification of the Tephritidae

Tephritinae. Two species of Pseudeutreta were bred from Baccbaris (Astereae). The sole species of Metatephritis may be only an abberant species of Eutreta (Uncaculeus), in which case Metatephritis would be the senior synonym of Uncaculeus. The most distinctive synapomorphy for Uncaculeus species is the transverse row of dorsal spines on the aculeus in the female but, unfortunately, females of Meta¬ tephritis are unknown. The head shape and lack of facial spots are similar in Meta¬ tephritis and Uncaculeus. The gall-forming Neotropical genus Polymorphomyia Snow might also be related to this group of genera. It has mixed black and white postocular setae and a wing pattern similar to those of Eutreta and Pseudeutreta in having an apical or subapical hyaline band, but the parafacial spot is absent. The Neotropical genus Dictyotrypeta is also included in this tribe. One un¬ described species that is tentatively placed here has large, erect scutellar bristles, but in most species, they are small. At least three species have a single, medial stripe on the eye. Four species have been reared from flowers of Heliantheae. Acidogona and Xenochaeta are most closely related to the Palearctic Carphotricha Loew, Noeeta Robineau-Desvoidy, and Paracanthella Hendel. Considering the rela¬ tively few species involved, all these taxa should perhaps be synonymized as one genus. Species of these genera have distinctive male genitalia, with the inner surstylus elongate and well separated apically from the outer surstylus, and the distiphallus is very slender. The scutellum is often swollen and shiny and has at least three pairs of erect bristles on the margin (sometimes small in Acidogona). Jamesomyia, which has a short inner surstylus and irregularly located and small (although often erect) scutel¬ lar bristles, has a similarly shaped distiphallus and outer surstylus and is probably related to these genera. The species of Acidogona, jamesomyia, Noeeta, and Xeno¬ chaeta whose biology is known breed in flowers of Lactuceae, and Carphotricha crepidis (Hering) was collected in numbers on a species of Crepis (Lactuceae). The swollen scutellum in most species in this group of genera suggests their possible relationship with some Dithrycini, but we have included them here because of the presence of the erect scutellar bristles. Cryptotreta and Xanthomyia are tentatively included in the Eutretini. A parafacial spot is present, but the scutellum lacks large, erect bristles. The eye color and the host plants of these genera are unknown. The relationships of Cryptotreta are poorly studied. Xanthomyia is most closely related to the eastern Palearctic genus Paranoeeta Shiraki, which probably should be synonymized with Xanthomyia. Both genera have a synapomorphy on the distiphallus, the apical third of which is mem¬ branous except for a distinctive, somewhat triangular, pointed apical sclerite. The outer surstylus is of the typical tephritine shape with a dorsal lobe present. Tribe Tephritini

Tephritini is the largest tribe of Tephritinae and it may be

paraphyletic. All of the Nearctic genera included have some bristles or setulae white, the head more or less quadrate, the thoracic microtrichia very dense, vein R4+5 not or only sparsely setose dorsally, and the lobe of cell cup short, in addition to lacking the derived features of other tribes. Dioxyna, Gonioxyna, Paroxyna, and Oxyna, which are mainly Holarctic or more cosmopolitan in distribution, have been considered related by authors studying Old

32

Classification of the Tephritidae

World species (Munro 1957, Freidberg 1987, Korneyev 1989). Species of these gen¬ era usually have small spines on the aedeagus just basal to the distiphallus (not present in Dioxyna), and an elongate, geniculate proboscis. The Nearctic species of

Dioxyna and Paroxyna appear to fit the classification of the authors cited above, but Paroxyna, in the sense used in this handbook, has been divided into several genera within the Palearctic Region. Without a consideration of genitalic features, which are important in the classification of this group, many of our species of Paroxyna will run to Campiglossa Rondani or Whiteina Korneyev in Korneyev’s (1989) key to the eastern Palearctic genera. The taxonomic status of the New World species thus needs reevaluation with consideration of the characters used by the authors cited above.

Gonioxyna, whose type is an Old World species, was recently synonymized with Campiglossa (Korneyev 1989). This change came to our attention too late in the production of this handbook to modify our taxonomic arrangement. Of the Nearctic species, at least G. fuscata Foote does not appear to be a Campiglossa; the classifica¬ tion of it and the other New World species needs further study.

Trupanea is one of the largest and most widespread genera of Tephritidae. It probably forms a monophyletic group with the Neotropical Celidospbenella Hendel, the endemic Hawaiian Phaeogramma Grimshaw, and the Old World Acantbiophilus Becker and Urelliosoma Hendel (and perhaps other genera), all of which have a large, strongly sclerotized, hooklike structure at the base of the distiphallus. Most species of this group have only one pair of scutellar bristles (two in Acantbiopbilus) and three or more frontal bristles. The wing typically has a stellate pattern but may be quite different. Trupanea as presently constituted is probably paraphyletic.

Tepbritis is mostly an Old World genus. It is not known to occur south of Mexico in the Americas. South American species formerly placed here (Foote 1967) belong in Trupanea or Trypanaresta.

Dyseuaresta is a mainly Neotropical genus. Its closest relatives are Lamproxynella Hering and Pseudoedaspis Hendel, which are restricted to South America. In all three genera, the outer surstyli are elongate and strongly curved posteriorly, and there is one pair of scutellar bristles.

Euarestoides is probably most closely related to Trypanaresta Hering and Plaumannimyia Hering. These three genera are restricted to the Americas, the latter two to the Neotropical Region. They have similar chaetotaxy (usually three frontal bris¬ tles, two pairs of scutellar bristles), and many species have stellate or evenly reticulate wing patterns; typically, there is a small hyaline spot at the apex of cell rl5 and the hyaline marginal spot beyond the apex of vein R2+3 is usually wedge-shaped rather than rounded as in Trupanea and Tepbritis. Trypanaresta and Plaumannimyia differ from Euarestoides in lacking the subapical pair of dorsal setae found on the hind femur in most tephntids. The larvae of all three genera breed in flower heads. All known hosts of Trypanaresta and Plaumannimyia are Astereae, especially Baccbaris.

Euarestoides dreisbacbi Foote and E. flavus (Adams) also breed in Baccbaris, but E. acutangulus (Thomson) and E. abstersus (Loew) have hosts in other tribes of Asteraceae. Except for species introduced elsewhere, Euaresta is also an endemic New World genus. Non-American species have been, or should be, transferred. The genus is

33

Classification of the Tephritidae

defined by two synapomorphies in the male: the epandrium is very broad, with posterior striations (fig. 93); and the fore femur is swollen. All species of Euaresta whose biology is known breed in the closely related plant genera Ambrosia and Xanthium. Neotephritis is probably most closely related to the Andean genus Homoeothrix Hering. Neotephritis has an interesting distribution; most species are American, but there are two native species in Hawaii (Hardy and Delfinado 1980). The relationships of Acinia are not well understood. Aczel (1958) suggested that the American and Palearctic species might be phylogenetically distinct groups, but no further research has been pursued in this area. Other New World genera belonging to this tribe include Ensina RobineauDesvoidy, Lamproxyna Hendel, Protensina Hendel, and Rhithrum Hendel (Foote 1980). Unplaced Tephritinae Their Asteraceae host plants as well as their morphologi¬ cal characters indicate that certain genera belong in the subfamily Tephritinae, but their tribal placement is uncertain. Gymnocarena and Mylogymnocarena are closely related. A distinctive synapomorphy is the presence of long, spinelike or scalelike structures on the apical half of the distiphallus. Foote (1980) placed Gymnocarena in the Trypetinae because the postocular setae are not as strongly swollen as in most Tephritinae, but the male genitalia and hosts indicate that both genera belong in the Tephritinae (Norrbom 1987). In retaining a number of plesiomorphic characters such as slightly swollen postocular setae, a long lobe on cell cup, vein R4+5 setose dorsally, and two scutellar and three or more frontal bristles, they resemble the Acrotaeniini, but the distiphallus is much different and the outer surstylus sometimes has a dorsal lobe. Icterica is an endemic Nearctic genus of two species that breed in flowers of Bidens (Heliantheae). The Palearctic species previously placed in it have been transferred to Orotava Frey (Korneyev 1989). Icterica retains some of the plesiomorphic features also found in Gymnocarena and Mylogymnocarena, such as slightly swollen post¬ ocular setae, two scutellar and three frontal bristles, and the lobe on cell cup moder¬ ately large, although the distiphallus is different. It might be related to them, al¬ though its closest relatives may be Palearctic genera. Xanthaciura has an “aciurine” wing pattern, but it does not belong in the Tephrellini (= Aciurini) because some of its bristles and setulae are white and expanded and its hosts are Asteraceae rather than Acanthaceae, Labiatae, or Verbenaceae. The type and only species of Cbrysaciura Aczel, C. bipunctata Aczel, is closely related to the species of Xanthaciura that have a short vein R2+3, and it differs from species such as X. speciosa Hendel and X. excelsa Aczel mainly in having the wing pattern yellow rather than brown. Thus, we consider Cbrysaciura a subjective synonym of Xanthaciura (new synonym). The Neotropical genus Lilloaciura Aczel also remains unplaced in our classifica¬ tion of the Tephritinae. We have not seen specimens of its only known species. Aczel (1953) discussed some characters in which it resembles Xanthaciura.

CURRENT STATUS OF TAXONOMIC STUDIES

Taxonomic investigations of Tephritidae around the world appear to be undertaken most aggressively on groups of economic importance. For example, the genus Bactrocera, which contains generally the world’s most economically important spe¬ cies, is well known and is being actively investigated taxonomically in every zoo¬ geographic region, and the genus Anastrepha in the New World has been and is being investigated as thoroughly as any of the other genera known to exist there. Perhaps a larger percentage of genera occurring in the United States and Canada than any¬ where else in the world have been recently revised, one of the factors that has made this handbook possible. Taxonomic catalogs of Tephritidae for all of the zoogeographic regions are now available. Although substantive changes have already been made in some of them, the compilation of a catalog of tephritids of the world now seems to be within reach. Thompson (1989) described a computerized biosystematic data base for entomol¬ ogy, and Thompson and Carroll (1991) described details of an expert system, using Tephritidae as the test group, that has been initiated and is being funded by the Agricultural Research Service of the USDA. We present here some highlights of taxonomic progress for each of the world’s zoogeographic regions.

NEARCTIC REGION Of all zoogeographic regions, the tephritid fauna in the Nearctic Region is probably the best known. Except for Mexico, where the fauna is poorly known but for the genus Anastrepha, many of the genera have recently been revised, abundant distribu¬ tional information is available, and a considerable amount of biosystematic work is 34

35

Current Status of Taxonomic Studies taking place. McAlpine (1981b) and Teskey (1981) included the Tephritidae in their keys to the North American dipterous families (adults and larvae, respectively). References to recent generic revisions will be found in the text of this handbook. Foote (1965a) presented a catalog of Tephritidae occurring north of Mexico, and Foote and Steyskal (1987) keyed the genera.

NEOTROPICAL REGION The collection of tephritid study material in Central and South America and the West Indies has been sporadic except in a few isolated areas or for a few specific genera. Anastrepha, through the taxonomic revisions by Stone (1939a, 1939b, 1942a) and more modern systematic studies by Norrbom (1985, and manuscripts in prepara¬ tion) and Norrbom and Kim (1988a), is perhaps the best known genus, mainly because of its actual and potential economic importance. Most geographical areas in the Neotropical Region are relatively unknown, especially for the subfamily Tephritinae. Foote (1967b) summarized the then available literature for the region in his catalog and provided a means for identifying the genera (Foote 1980).

PALEARCTIC REGION Because the earliest collections of Tephritidae were made in the Palearctic Region, and because a great many entomologists have participated in the accumulation of study material over the years, this region is recognized as the “grandfather” location of tephritid taxonomy. It is especially important to the taxonomist because of the immense size of the literature and the large number of type specimens located in museums throughout the region. Ito (1983-1985) completely revised the Japanese tephritid fauna. Various aspects of the tephritid fauna, especially of Europe and the Near East, are being brought up to date through the work of White, Freidberg, Korneyev, and others, and White (1988) provided an effective guide to the Tephriti¬ dae of Britain. A taxonomic catalog by Foote (1984), amended by White (1986), treats the tephritid taxa of the Palearctic Region.

AFROTROPICAL REGION Because many geographic areas are inaccessible and because few taxonomists have had the interest, much remains to be done in the Afrotropical Region. Pioneer taxonomic work is reflected in the many publications of Munro, especially his faunal revisions (Munro 1947, 1957) and his 1984 monograph revising the genus Dacus as the family Dacidae. Cogan and Munro (1983) produced a comprehensive catalog of the Tephritidae of this region, and more recent investigations have been and will continue to be reported by Freidberg, Hancock, and others.

36

Current Status of Taxonomic Studies

ORIENTAL REGION The Tephritidae of the Oriental Region are fairly well known, primarily through the immense amount of detailed taxonomic research reported by Hardy. This research culminated in a catalog published in 1977 and is being continued by him for Indo¬ nesia and certain geographical areas in the Australasian Region (see References Cited, which lists Hardy’s most significant contributions to our knowledge of the region).

AUSTRALASIAN-OCEANIC REGION The Tephritidae of the Australian-Oceanic Region have been long unknown. In a recent series of publications dealing in part with New Guinea and the Bismarck and Solomon Islands, however, Hardy has remedied this situation. The catalog by Hardy and Foote (1989) is a major review of the fauna of this region (Evenhuis 1989a). The subfamily Dacinae has been the subject of intensive research by R. A. I. Drew through a number of publications in the 1970s and 1980s, and his latest revision (1989) summarizes virtually all the known taxonomic information concerning the subfamily in the region.

TECHNIQUES FOR STUDY

Many sources may be consulted for the care and handling of insect specimens. Perhaps foremost among these is the recent manual by Steyskal et al. (1986), which describes techniques for collection and preservation of insects and mites in consider¬ able detail. Although tephritids may be treated in the same way as many other kinds of flies, the study of some of their features is enhanced by the use of somewhat specialized techniques. We describe here those that have proved useful for our studies of this fly family.

MOUNTING Always mount dry specimens so they will not rest on a surface or be touched during handling and observation. Whenever possible, mounting should be done as soon as the specimens are killed and while they are still flexible, because drying makes bristles and appendages brittle and easily broken or lost. Do not try to pin specimens that are completely dry because internal fluids must be present to act as a fixative in holding specimens in place during and after drying. Dry specimens should be either held in a relaxing jar for a day or two before pinning or mounted on points with glue. Every attempt should be made to preserve all characters on at least one side of the center line, regardless of the mounting method. If the specimen is large enough, a No. 1 or 2 pin may be inserted on the right side of the center line at midscutum and pushed straight through to the desired height, being careful also to preserve the left side of the venter. With smaller specimens, a minuten pin fastened to a piece of cork or plastic on a regular pin may be used. The minuten should stand away to the left of the pin and be inserted into the specimen through the thorax. The specimen also may be glued to a cardboard point; in this case the point should project from the pin to the left, and the tip should always be bent down at 90° so the specimen can be glued 37

38

Techniques for Study by its right side, facing away from the observer, scutum up, with wings and legs positioned at 45° to each other. Use an adhesive that will not become brittle with age; small amounts are indicated with today’s modern glues. After mounting, arrange appendages so that all possible morphological characters may be viewed easily.

SPECIMENS IN FLUIDS Adult specimens mounted in the various ways described above are generally preferred over those preserved in alcohol. They are less subject to damage, more conveniently handled and labeled, and easier to store. In addition, specimens in alcohol gradually become brittle over time and eventually will never yield to clearing. On the other hand, certain morphological characters appear to be easier to see in “alcoholic” specimens, and often it is not necessary to macerate genitalia for certain types of study. For example, when live females are killed in 70% alcohol, the aculeus often is ejected from the eversible membrane far enough so that the free tip can be examined in detail without further treatment. If long-term use is not indicated, specimens should be stored in ethyl alcohol no more dilute than 70% in vials with leakproof closures. To minimize damage to specimens when containers are being moved, confine the specimen(s) to a small space at the bottom of the vial with a wad of nonfibrous material such as high-quality facial tissue.

IMMATURE STAGES Normally all immature stages of tephritids (except puparia) are stored and studied in 70% alcohol; they are virtually useless as dried specimens because they shrink and become malformed. Some of the useful characters of larvae and pupae in liquid can be seen clearly enough with a dissecting microscope. Often it is necessary to view extremely small papillules at the posterior end of larvae, in which case a specimen may be raised out of the fluid along the side of a dish, allowed to dry temporarily, and studied in a low angle of light. Eggs can most often be found in gravid females whose abdomens have been cleared as described elsewhere in this section. No universally used technique has been developed for the detailed study of eggs, but their shapes are often quite clearly outlined in slide-mounted abdomens. As taxonomic studies of these stages progress, other characters requiring the use of a compound microscope will be found. In those cases, taxonomists will very probably describe the techniques required for the study of those structures.

LABELING Regardless of whether specimens are to be archived, they should be labeled in such a way that the data can be separated from them only with difficulty. This is especially

39

Techniques for Study true of specimens that are difficult or impossible to identify, as these are ones most likely to require further study. Locality (in as much detail as feasible, preferably country, state or province, county, and locality), full date, and collector are the minimum data required with an individual specimen. With plant-inhabiting flies such as the Tephritidae, it is also highly desirable to append host data which should specify the host status as well (i.e., “swept from . . . ,” “reared from . . . ,” etc.). Simply writing a plant name on a label may cause confusion about whether it is a true host or merely the plant on which the fly was collected. If parts of specimens such as appendages, genitalia, and wings are separated from the specimen and mounted separately, always use a unique number or symbol to associate the separate parts. Male genitalia should not be slide-mounted; they should be stored in glycerol in a microvial, the stopper of which is impaled on the pin containing the pertinent specimen. Female genitalia also may be stored in this way rather than slide-mounted. Permanent slides are useful because they can be examined rapidly, but the genitalia cannot be viewed in different positions or placed close to other specimens for comparison.

MACERATION (CLEARING) When required, we clear specimens or parts thereof in fresh 10% KOH. This chemi¬ cal is used at room temperature (or heated for only a few minutes) to give us control over the end-point of the clearing process; on average, an overnight treatment will suffice, but more time is required for large specimens and those having very dark or black pigment. If the specimen appears to resist clearing, siphon off the old fluid and add fresh, and gently press hollow structures a few times to introduce fresh clearing agent to the interior. The clearing process must be stopped at the appropriate time by immersing the specimens in water. If balsam is to be used as a mounting medium, the specimens should be immersed in an alcohol series of 50, 70, and 95% and then in Cellosolve for about an hour before transferring them to the balsam. Further direc¬ tions and alternative methods of mounting are presented by Steyskal et al. (1986).

ACULEUS At times it becomes necessary to examine details of the aculeus, especially of genera such as Anastrepha and Urophora, in which accurate identification of species is otherwise not possible. To assure that all the organs associated with the terminalia are preserved properly, it is necessary to remove the entire abdomen from a dry specimen. Usually this can be achieved by grasping the abdomen base with fine forceps and breaking it away from the metathorax with a twisting motion. If the forceps are dipped beforehand in alcohol, the abdomen will stick to them, thus preventing it from suddenly flying off to a location where it can be found only with extreme good fortune. The abdomen should be macerated as described above. The maceration process

Techniques for Study gradually softens the nonsclerotized parts so they can be moved relative to one another, making it possible to squeeze any air bubbles gently from the interior for faster and more uniform processing. After all the interior sclerotized parts have been made clearly visible through the integument but before they lose their color and definition, remove the abdomen and oviscape from the clearing medium and transfer them to water. The procedure for liberating the aculeus from the eversible membrane as described below is made easier by using a relatively flat surface because needles can be held horizontally rather than vertically, preventing tearing of tissues with sharp points. A microscope slide is ideal because either temporary or permanent mounts can be made in situ. If a temporary mount will suffice, use a drop of glycerine for the operation; if a permanent slide is required, the operation can be performed directly in the ultimate mounting medium (i.e., balsam). The procedure is as follows:

(a) Carefully separate the oviscape from the terminal abdominal segment with a pair of fine needles or forceps, gently drawing the spermathecae free of the interior of the abdomen, and place them on the slide. (b) Position the oviscape ventral side up. (c) With a fine needle, hold the dorsal side of the oviscape (the side next to the slide) in place against the slide surface at its proximal end. (d) Insert another needle into the open “mouth” of the oviscape and gently engage the needle tip against the proximal end of the aculeus. (e) While holding the oviscape in place, gradually and gently push the aculeus free of the oviscape until at least half of the aculeus tip is exposed. (f) Withdraw the pin used for this pushing operation, position it on the freed part of the aculeus at a point well removed from the tip (or grasp the free part with fine forceps), then gently pull the aculeus away from the oviscape until it is lying com¬ pletely free. (g) If a view of the eversible membrane is required, continue pulling as in (f) above until the oviscape, the membrane, and the aculeus itself lie in a line as shown in fig. 7.

An alternative method for freeing the aculeus from the oviscape is to simply squeeze the abdomen and subsequently the base of the oviscape of a fresh specimen with watchmaker’s forceps to force the aculeus free. Occasionally the abdomen and oviscape are broken, and the aculeus often is pulled partially back into the oviscape as the specimen dries, but it is an effective method for making at least the tip of the aculeus clearly visible. For a permanent mount, process the abdomen and transfer it to the slide, then complete the slide-making procedure in the usual way. It is convenient to orient the aculeus under the cover slip with the tip toward the observer, the usual position in which illustrations of these parts are shown in publication, the opposite if these structures are to be observed more often with a compound scope, which inverts the image.

41

Techniques for Study

MALE GENITALIA We have purposely avoided using the characters of male genitalia wherever possible in this handbook because of their complexity and the difficulty in dissecting and studying them. When removed for study, they are usually macerated, appropriately dissected, then stored in glycerine in a microvial. The vial is kept with the adult by inserting the pin holding the specimen through the stopper of the vial.

WINGS The general aspects as well as details of tephritid wing patterns should be observed against a white or very light background, which transmits some light through the wing. Some reflected light also is required so that the setulae on the veins and the character of the surface can be studied. The setation of veins may often be seen best at a very small angle to the disk. We often slide-mount wings to obtain more precise measurements of patterns, vein positions, and the relationships between the two; the plane of the wing needs to be as flat as possible for this purpose. In this case, usually the right wing of a specimen is taken. With fine forceps, grasp the wing base close to the thorax, breaking away as much of the wing base as possible. Wash the wing in a Cellosolve bath with a gentle motion to dislodge adhering foreign particles, which become quite bothersome when viewed in transmitted light under a microscope. The wing may be transferred directly from Cellosolve to balsam on a slide, leaving as much as possible of the Cellosolve behind. During orientation on the slide (wing tip to the right, dorsal side up) any air bubbles may be teased out from under the wing disk; other small bubbles will disappear automatically during the hardening process, and the Cellosolve transferred with the wing will gradually dissolve in the balsam, resulting in a perfectly clear mount. The brittle wings of old specimens must be handled with extreme care.

ABOUT THIS HANDBOOK

This publication is designed as a guide to the identification of the adults of the Tephritidae known to occur in the United States and Canada at the present time. The information contained herein is limited to that which might assist the user in identi¬ fying a particular specimen of fruit fly, so coverage is not monographic in any sense of the word. The following list of features may assist the user in making the most efficient use of the contents.

KEYS

A key to the genera introduces the taxonomic treatment, and we have included a key to the species of every genus that contains two or more species. As noted in the discussions of the genera Bactrocera, Ceratitis, Dacus, and Urophora, we have in¬ cluded in the keys only those exotic species that have actually been introduced into the United States and Canada; the keys will not distinguish any of them from other members of these genera occurring elsewhere.

ILLUSTRATIONS

We have attempted to illustrate every important key character; illustrations appear in the order they are mentioned in the keys. The discussions of the various species also refer to these figures. We have included an illustration of the wing pattern of nearly every species discussed in this handbook.

42

43

About this Handbook

GENERA The genera are arranged alphabetically. Within each genus the following information is provided: author; references to the genus as a whole in the form of a “synonymy,” which also presents all names under which the genus has been known in the North American literature; a recognition section, which discusses the taxonomic characters that separate that genus from all other Tephritidae in the region; and a discussion setting forth other pertinent information.

SPECIES The species are arranged alphabetically within each genus. For each species we have provided the following information where available: author; common name(s); refer¬ ences to figures and maps; a synonymy and literature review as described for the genus; type data and depositories; a recognition section; a distribution section, in most cases including a map of the U.S. and Canadian range and comments on extraterritorial occurrences; a generalized account of hosts; and a general discussion of the species where appropriate. Most of the common names have been taken from Auclair et al. (1964), Sutherland (1978), and Stoetzel (1989).

LITERATURE Content Our literature review includes a reference to the original description of every taxon known to occur in the United States and Canada, including those that have been introduced. We believe we have provided a thorough coverage of the early references to these taxa. On the other hand, coverage since about 1980 may be sporadic because, in many cases, we have included only references to critical tax¬ onomic research. Format The format of the review for each taxon is patterned after that found in most formal taxonomic publications (i.e., in a synonymy). The names under which the taxon has been known are given in chronological order. For each year and author, the order is by page number; and if on the same page, the order is alphabetical by genus or species name. The literature pertaining to the species under a particular name is chronological by year, then alphabetical by author for each year. Each reference consists of the name of the author, the year of publication, and the page number. The full citation appears in the bibliography concluding this volume. Annotations

An annotation indicating the subject matter of each reference also is

included, often in very general terms; for example, the term biology is used in cases in which a single reference includes many aspects of that subject, and taxonomy is used for a reference that may pertain to a combination of nomenclature, description,

44

About This Handbook systematic position, or other taxonomic activity. A state abbreviation in an annota¬ tion refers to the appearance of the name in a list. We make a distinction between the terms revision and review, in that the former refers to a formal taxonomic presenta¬ tion, whereas the latter represents a more generalized treatment.

TYPE DATA Each generic name is accompanied by a reference to its original description, the name and original combination of the type species, the method of designation, and the valid name if applicable. Following the reference to the original description of a species, we give the data for the primary type, depository of the primary type speci¬ men^), and the type locality in parentheses in the following format: (type data, depository; type locality). The list of type depositories and their abbreviations follows: AMNH ANSP BMNH CAS CD FA CMP CNC CSUFC CUI DEI

INHS IOC LACM LCL MCZ MHNL MNHN MZLS NMNH NMW NRS PDAH RNHL SMT

American Museum of Natural History, New York, N.Y. Academy of Natural Sciences, Philadelphia, Pa. Natural History Museum (formerly British Museum [Natural History]), London, England California Academy of Sciences, San Francisco California Department of Food and Agriculture, Sacramento Carnegie Museum, Pittsburgh, Pa. Canadian National Collection, Ottawa, Canada Colorado State University, Ft. Collins Cornell University, Ithaca, N.Y. Institut fur Pflanzenschiitzforschung der Akademie der Landwirtswissenschaften, Abteilung Taxonomie der Insekten (for¬ merly Deutsches Entomologisches Institut), Eberswalde, Ger¬ many. Illinois Natural History Survey, Urbana Instituto Oswaldo Cruz, Rio de Janeiro, Brazil Los Angeles County Museum, Los Angeles, Calif. Linnaean Collection, Linnaean Society, London, England Museum of Comparative Zoology, Harvard University, Cam¬ bridge, Mass. Musee d’Histoire Naturelle, Lille, France Museum National d’Histoire Naturelle, Paris, France Museo Zoologico “la Specola,” Florence, Italy National Museum of Natural History (formerly U.S. National Museum), Smithsonian Institution, Washington, D.C. Naturhistorisches Museum, Vienna, Austria Naturhistoriska Riksmuseet, Stockholm, Sweden Pennsylvania Department of Agriculture, Bureau of Plant Indus¬ try, Harrisburg Rijksmuseum van Natuurlijke Histoire, Leiden, Netherlands Staatliches Museum fur Tierkunde, Dresden, Germany

45

About This Handbook SUS UCD UCR UKL UMO UMSP WSUP ZMHU ZMUC

Stanford University, Stanford, Calif. University of California, Davis University of California, Riverside University of Kansas, Lawrence (Snow Museum) University Museum, Oxford University, Oxford, England University of Minnesota, St. Paul Washington State University, Pullman Zoologisches Museum, Humboldt Universitat, Berlin, Germany Zoologiske Museum, Universitets Copenhagen, Denmark

DISTRIBUTION Since 1954, R. H. Foote and F. L. Blanc have recorded information on distribution in detail for a great many species encountered in their work on the Tephritidae, and F. L. Blanc has accumulated these data in a 5" x 8" card file. The user of this handbook may consult Mr. Blanc for details omitted here. Rather than including all of this distributional information in detail, we show the ranges of nearly all species on maps. All distribution outside the United States and Canada is discussed briefly in the text; if no information on a species distribution is presented in the Distribution section, the species is not known to occur outside our area.

HOSTS Despite the great importance of host information in the recognition of tephritids, we have not been able to present a formal host list. The most comprehensive host information for species occurring in North America north of Mexico, on which we have relied heavily, was compiled by Wasbauer (1972). For information beyond that date, the user should rely on information in the host sections and on references annotated as “host” that may appear in the literature sections of individual species. The plant names and associated text page numbers that appear in the index of plant names at the end of this book may be used as a substitute host list. If a large number of plant species are known as hosts of a given tephritid species, we have listed only the plant genera.

TERMINOLOGY With a few exceptions, we have adhered to the terminology for adult flies proposed by McAlpine (1981a). Otherwise, we have used the simplest terms possible to en¬ courage the use of this manual by the nonprofessional taxonomist. Consult the legends for figs. 1-11 to identify alternative terms that are often encountered in the literature.

46

About This Handbook

NOMENCLATURE The following nomenclatural changes, some of which do not normally appear in a handbook such as this, have been made for the first time or are brought to the reader’s attention because the literature in which they first appeared may not be universally available. See the respective sections noted below for additional details. Classification (Introduction). The new tribe Acrotaeniini and the new subtribe Eurostina are proposed. Pseudacrotaenia Hendel is made a new synonym of Baryplegma Wulp, and Cbrysaciura Aczel is made a new synonym of Xanthaciura Hendel. Acrotaenia. A lectotype is designated for Trypeta testudinea Loew. Anastrepha. A lectotype is designated for Trypeta ludens Loew. Anastrepha latbana Stone is now a synonym of Anastrepha ludens, as explained in detail by Hernandez (1991). Dacus and Bactrocera. In accordance with changes made by Drew (1989b), we regard all but one of the accidentally introduced species formerly assigned to Dacus as belonging to the genus Bactrocera. Dacus bivittatus remains untransferred. We strongly urge readers to adopt this change in view of the probability that Drew’s conclusions will become universally accepted by authorities on the Dacinae. Eurosta. Lectotypes are designated for Trypeta comma Wiedemann and Acinia solidaginis Fitch. In accordance with Ming (1989), Eurosta conspurcata Doane and Eurosta reticulata Snow are now synonyms of Eurosta cribrata Wulp, Eurosta elsa Daecke is now a synonym of Eurosta comma (Wiedemann), the varieties fascipennis Curran and subfasciata Currran of solidaginis are now treated as subspecies, and subfasciata is now a synonym of fascipennis. Eutreta. Eutreta navajorum Blanc is made a new synonym of Eutreta intermedia Stoltzfus. Paracantba. Tepbritis marginepunctata Macquart, which appeared as an “un¬ known species” of Paracantba in Foote (1965a), is made a new synonym of Paracantba culta (Wiedemann). Paroxyna. In accordance with Jenkins (1985), the species long known as Tepbritis opacipennis Foote is now recognized as a member of the genus Paroxyna. Peronyma. The species long known as Peronyma sarcinata (Loew) is given the name Peronyma quadrifasciata (Macquart) via the examination of the type of the latter by A. L. Norrbom. Rbagoletis. Rbagoletis meigenii (Loew), never before recognized as being present in North America, has been added by virtue of its discovery in New Hampshire. Uropbora. In accordance with recent taxonomic investigations by White and Korneyev (1989), names of the introduced species Uropbora affinis and Uropbora quadrifasciata are now to be used at the subspecies level because of the discovery of additional subspecies in the Old World. Xantbaciura. Cbrysaciura bipuncta Aczel is transferred to Xantbaciura (see the discussion of Xantbaciura), thereby becoming a new combination.

KEY TO THE U.S. AND CANADIAN GENERA

1. Cell bm (fig. 13, a) much wider than cell cup (fig. 13, b); ocellar (fig. 14, a), dorsocentral (fig. 14, d), katepisternal (fig. 14, c), and usually postpronotal lobe bristles (fig. 14, b) lacking .2 Cells bm and cup approximately equal in width (fig. 17); ocellar and dorso¬ central bristles at least weakly present; bristle of postpronotal lobe and katepisternal bristles usually present.3 2. Abdominal tergites (fig. 7) fused .Dacus Fabricius 1 sp. adventive in California; many Afrotropical species Abdominal tergites articulate, not fused .Bactrocera Macquart 6 spp. adventive in California; many species worldwide 3. Apex of cell cup (fig. 17, b) closed by transverse or convexly bent vein, the cell never drawn out into a point, however short, in lower apical corner; only 1 pair of orbital bristles (fig. 1); all postocular setae (fig. 1) and scutal setulae dark and slender .4 Apex of cell cup closed by a concavely bent vein, resulting in at least a slight elongation of cell in lower apical corner (figs. 13, c; 18, a); usually 2 pairs of orbital bristles; postocular setae and scutal setulae variable, sometimes whitish and swollen .5 4. Face strongly produced at anterior oral margin (fig. 19, a); proboscis strongly geniculate, labellum slender and longer than head (fig. 19, c); anepisternum (fig. 5) and scutellum tomentose, dull. .Rhynencina Johnson 1 sp., Massachusetts to Georgia Face not strongly produced at anterior oral margin (fig. 20, a), proboscis at most short-geniculate or moderately long (fig. 20, b); labellum shorter than head, or if slightly longer, anepisternum or scutellum or both not tomentose, shining.Urophora Robineau-Desvoidy 8 spp. native to U.S.; several Palearctic species introduced for weed control

47

Figures 12, 13. Right wings. 12, Toxotrypana curvicauda Gerst.; 13, Bactrocera dorsalis (Hend.). Figure 14. Dorsal view, head, thorax, abdomen, Bactrocera dorsalis (Hend.) (missing bristles shown by dotted lines). Figure 15. Dorsal view, abdomen, Toxotrypana curvicauda Gerst. (figs. 14 and 15 shown to same scale). Figure 16. Side view, abdomen and oviscape, Toxotrypana curvicauda Gerst. Figures 17,18. Right wings. 17, Urophora caurina (Doane); 18, Euaresta aequalis (Lw.). Figures 19, 20. Lateral view, head. 19, Rhynencina longirostris Johns.; 20, Urophora caurina (Doane).

48

Figure 21. Right wing, Zonosemata electa (Say). Figures 22, 23. Generalized scutum, showing position of dorsocentral bristles. Figures 24, 25. Right wings (color pattern not shown). 24, Anastrepha dentata (Stone); 25, Euaresta bullans (Wied.). Figure 26. Dorsal view, scutellum, Hexachaeta seabrai Lima. Figures 27, 28. Front view, head. 27, Paraterellia immaculata Blanc; 28, Zonosemata vittigera (Coq.). Figures 29, 30. Right wings. 29, Callachna gibba (Lw.); 30, Ceratitis capitata (Wied.).

49

50

Key to the U.S. and Canadian Genera

5. Dorsocentral bristles (fig. 22, b) not arising farther forward than midpoint between supra-alar (fig. 22, a) and acrostichal (fig. 22, c) bristles, or if farther forward, posterior orbital bristles reclinate and wing with dark bands covering veins r-m and dm-cu forming a V which encloses a small dark costal spot (fig. 21, a) .6 Dorsocentral bristles arising anterior to midpoint between supra-alar and acrostichal bristles (fig. 23, a, c), or if not (some species of Myoleja), then posterior orbitals minute or absent.11 6. Vein M curved forward apically (fig. 24, a), gradually merging into costa .Anastrepha Schiner 20 spp., California, Florida, Texas, Bermuda; many others southward Vein M not curved forward apically (fig. 25, a), meeting costa at a distinct angle.7 7. Scutellum with 3 or more pairs of bristles (fig. 26) ... Hexachaeta Loew 3 spp., Texas; several others southward Scutellum with no more than 2 pairs of bristles.8 8. Vein R2+3 undulate with at least 1 short appendix (fig. 12, a); oviscape much longer than abdomen (figs. 15, a; 16, a); scutum with medial longitudinal depression; abdomen petiolate . .Toxotrypana Gerstaecker 1 sp., Florida, Texas; several others southward Vein R2+3 not undulate and without appendix (see fig. 6); oviscape shorter than abdomen (fig. 14, e); scutum flat or evenly convex; abdomen not petiolate .9 9. Posterior orbital bristles convergent (fig. 27, a); wing without intercalary band (fig. 21, a), or if present, fused to other bands. .Paraterellia Foote 4 spp., western U.S. Posterior orbital bristles divergent or reclinate (fig. 28, a); wing with iso¬ lated intercalary band (fig. 21, a) .10 10. Scutum tomentose, with 2 pairs of small bare spots at the inner ends of the parts of the transverse suture and at the bases of the dorsocentral bristles; 1st flagellomere rounded apically (as in fig. 33, a); postocular setae poor¬ ly differentiated.Oedicarena Loew 3 spp., western U.S. Scutum shiny, without tomentum; 1st flagellomere drawn to a point apically (fig. 34, a); postocular setae differentiated . .Zonosemata Benjamin 2 spp., eastern Canada, widespread U.S., several species southward 11. Scutellum inflated, usually polished or bilobate (fig. 36, a); wing with dark bands on hyaline background (fig. 29), the pattern not reticulate. ...12

Scutellum not distinctly swollen or polished; wing sometimes with reticulate pattern .jg 12. Lower outer corners of face with white setae (fig. 33, b); costa and vein ^2+3 connected by transverse supernumerary vein distad of pterostigma (fig. 29, a); wing pattern as in fig. 29.Callachna Aldrich 1 sp., widespread Lower outer corners of face without white setae; costa and vein R, , , other¬ wise . 23

37

Figure 31. Middle (left) and hind (right) legs, Oedicarena latifrons (Wulp). Figure 32. Base of right wing, Ceratitis capitata (Wied.). Figures 33-35. Lateral view, head. 33, Callachna gibba (Lw.); 34, Zonosemata vittigera (Coq.); 35, Ceratitis capitata (Wied.)., male. Figure 36. Lateral view, scutellum, Callachna gibba (Lw.). Figure 37. Posterior view, scutellum, Peronyma quadrifasciata (Macq.). Figures 38, 39. Right wings. 38, Cecidocharella borrichia Bush and Huettel; 39, Procecidochares atra (Lw.).

51

52

Key to the U.S. and Canadian Genera

13. Wing proximad of pterostigma with dark streaks and spots in middle of cells (fig. 30, a); apical extension of cell cup narrower at base than at middle (fig. 32, a); male with anterior orbital bristle apically flat and diamond-shaped (fig. 35, a); wing pattern as in fig. 30 . .Ceratitis Macleay 1 sp., adventive in California, Florida, Texas, Bermuda; widespread in world’s semitropics Wing with dark color proximad of pterostigma not as streaks and spots in middle of cells; apical extension of cell cup gradually tapering; orbital bristles simple.14 14. Scutellum bilobed, with shallow mesal depression dorsally (fig. 37, a) ... .Peronyma Loew 1 sp., southeastern U.S. Scutellum evenly rounded dorsally, without mesal depression .15 15. Vein r-m situated near middle of cell dm and approximately half-way be¬ tween veins bm-cu and dm-cu, not in same dark band that covers vein dm-cu (figs. 38, a; 40, a) .16 Vein r-m well apicad of middle of cell dm and closer to vein dm-cu, situated in same dark oblique band that covers vein dm-cu (figs. 39, a; 41, a) ..17 16. Wing with only 1 hyaline mark crossing cell r1 distad of pterostigma, com¬ pletely crossing wing, or, if interrupted, in line with spot in cell cua1 (fig. 38, b); anepimeron (fig. 5) swollen, whitish ... Cecidocharella Hendel 1 sp., Texas; others southward Wing with 2 hyaline marks crossing cell apicad of pterostigma, the more apical one extending posterad of cell dm and apicad of hyaline mark crossing cell cua1 (fig. 40, b); anepimeron black .Stenopa Loew 2 spp., widespread southern Canada and U.S. 17. One or 2 pairs of orbital bristles present, posterior pair minute if present (fig. 42, a); hyaline area between 2nd and 3rd wing bands without a dark mark (fig. 39, b); 1-2 pairs of dorsocentral bristles. .Procecidochares Hendel 11 spp., southeastern Canada, widespread U.S. Two pairs of orbital bristles present (fig. 43, a), the posterior pair well developed, often convergent (as in fig. 44, a); at least a faint mark or stripe always present in hyaline area between 2nd and 3rd bands of wing pattern below middle of cell dm (fig. 41, b); 2 pairs of dorsocentral bristles .Procecidocharoides Foote 4 spp., southeastern Canada, widespread U.S. 18. Posterior orbital bristles distinctly convergent (fig. 44, a); 1st flagellomere rounded at apex dorsally; scutum usually with lyre-shaped dark gray pattern .19 Posterior orbital bristles reclinate or absent; scutum without lyre-shaped pattern .21 19. Anterior oral margin with a few well-developed bristles larger than adjacent setae (fig. 45, a) .Chaetostomella Hendel 1 sp., western U.S.; several Palearctic species Anterior oral margin with only equally small, short setae (fig. 44, b) _ .20

Figures 40, 41. Right wings. 40, Stenopa vulnerata (Lw.); 41, Procecidocharoides penelope (O.S.). Figures 42, 43. Lateral view, upper head. 42, Procecidocbares flavipes Aid.; 43, Procecidocharoides pullatus Foote. Figure 44. Front view, head, Orellia palposa (Lw.). Figure 45. Lateral view, head, Chaetostomella undosa (Coq.). Figure 46. Right wing, Valentibulla munda (Coq.). Figure 47. Base, right wing, Orellia occidentalis (Snow). Figures 48, 49. Right wings. 48, Orellia palposa (Lw.); 49, Neaspilota verrtoniae (Lw.).

53

Figures 50, 51. Generalized scutum, showing position of dorsocentral bristles. Figure 52. Lateral view, head, Chetostoma rubidum (Coq.). Figure 53. Generalized scutum, showing position of presutural bristles. Figure 54. Lateral view, head, Epochra canadensis (Lw.). Figures 55-58. Right wings. 55, Strauzia longipennis (Wied.); 56, Ensina sonchi (L.); 57, Rhagoletis indifferens Curr. (pattern not shown); 58, Icterica seriata (Lw.) (pattern not shown). Figure 59. Lateral view, head, Ensina sonchi (L.).

54

55

Key to the U.S. and Canadian Genera

20. One to several setae present on node of vein Rs (fig. 47, a); apex of cell r4+5 broadly brown (fig. 48) .Orellia Robineau-Desvoidy 3 spp., Alaska, Canada, and U.S. Node of vein Rs without setae; apex of cell r4 + 5 broadly hyaline (fig. 49) .Neaspilota Osten Sacken 19 spp., Alaska, widespread southern Canada and U.S.; also Mexico 21. Dorsocentral bristles (fig. 50, b) closer to level of supra-alar bristles (fig. 50, c) than to transverse suture (fig. 50, a) .22 Dorsocentral bristles closer to transverse suture than to level of supra-alar bristles (fig. 51, b); if doubtful, bulla present in cell r4+5 (fig. 46, a) .. .38 22. Lower margin of gena with several well-developed setae anterior to genal bristle (fig. 52, b).Chetostoma Rondani 2 spp., southern Canada, widespread U.S. Lower margin of gena without such large setae in addition to the genal bristle (fig. 54, c) .23 23. Only 1 pair of orbital bristles present (fig. 54, a).24 Two to 4 pairs of orbital bristles present (fig. 52, a) .26 24. Presutural bristles lacking (fig. 53, a); anatergite with long, fine setae; an¬ tenna extending to, or nearly to, oral margin (fig. 54, b). .Epochra Loew 1 sp., widespread southern Canada, northern U.S. Presutural bristles present (fig. 53, a); anatergite without long setae; anten¬ na shorter, extending only slightly below middle of face (figs. 59, a; 60, a) .25 25. Head higher than long, rounded below (fig. 60, b); proboscis not geniculate (fig. 60, c); wing with pronounced banded pattern (fig. 55) . .Strauzia Robineau-Desvoidy 7 spp., widespread southern Canada and U.S. Head longer than high, straight below (fig. 59, c); proboscis geniculate (fig. 59, b); wing pattern nearly absent or consisting of a few spots (fig. 56) .Ensina Robineau-Desvoidy 1 sp., nearly cosmopolitan, eventually may be found in North America 26. Vein r-m situated approximately at middle of cell dm (fig. 57, a).27 Vein r-m situated rather close to vein dm-cu and distinctly apicad of middle of cell dm (fig. 58, a).29 27. First flagellomere pointed at dorsal apex (fig. 61, a). .Rhagoletis Loew 23 spp., widespread North and South America and Palearctic Region First flagellomere rounded apically (figs. 59, a; 62, a) .28 28. Background color of body yellow, sometimes with a few dark markings; wings as in figs. 475-481 .Trypeta Meigen 7 spp., widespread Alaska, Canada, and U.S.; widespread in Palearctic Region Background color of body dark brown to black; scutum with yellowish median streak.Rhagoletotrypeta Aczel 2 spp., northeastern U.S., Texas 29. All postocular setae dark brown to black (figs. 65, b; 66, b).30 Some or all postocular setae whitish or yellowish (fig. 62, b) .31

Figures 60-62. Lateral view, head. 60, Strauzia longipennis (Wied.); 61, Rhagoletis pomonella (Walsh); 62, Dioxyna sororcula (Wied.). Figures 63, 64. Right wings. 63, Gymnocarena diffusa (Snow); 64, Mylogymnocarena apicata (Thomas). Figures 65, 66. Lateral view, head. 65, Euleia fratria (Lw.); 66, Myoleja limata (Coq.). Figures 67, 68. Dorsal view, scutellum. 67, Eurosta s. solidaginis (Fitch); 68, Acidogona melanura (Lw.). Figures 69, 70. Lateral view, head. 69, Eurosta cribrata (Wulp); 70, Acidogona melanura (Lw.).

56

57

Key to the U.S. and Canadian Genera

30. Facial carina strongly developed, protuberant (fig. 65, a). .Euleia Walker 2 spp., widespread Alaska, Canada, and U.S. Facial carina not strongly developed (fig. 66, a) .Myoleja Rondani 4 spp., widespread southern Canada and U.S. 31. Surface of frons bare; all head and body bristles yellowish to brown. .32 Surface of frons with fine setae; head and body bristles yellowish, brown, or black.33 32. Cell r, with 2 hyaline areas apicad of pterostigma (fig. 63, a); femora of all 3 legs noticeably enlarged.Gymnocarena Hering 3 spp., western Canada, central and northwestern U.S. Cell rj with only 1 hyaline area apicad of pterostigma (fig. 64, a); femora not noticeably enlarged .Mylogymnocarena Foote 2 spp., western U.S. 33. Apical pair of scutellar bristles (fig. 67, a) lacking or distinctly shorter than basal pair; 2-4 pairs of frontal bristles present (fig. 69, a); 1st flagellomere (fig. 69, b) shorter than V2 height of eye . .Eurosta Loew 9 spp., widespread southern Canada and U.S. Apical pair of scutellar bristles about as long as basal pair (fig. 68); frontal bristles and antennae variable .34 34. First flagellomere at least 2h> as long as height of eye and more or less acute at dorsal apex (fig. 70, b); 3 or 4 pairs of frontal bristles present (fig. 70, a) .Acidogona Loew 1 sp., Canada south to Florida First flagellomere less than V2 as long as height of eye (as in fig. 69, b); 2-4 pairs of frontal bristles present .35 35. Body and bristles wholly yellowish; wing disk yellowish, varied with irregu¬ lar brown markings and with border of apical half distinctly darkened; apex of cell r4 + 5 with transverse brown band (fig. 71, a) . .Icterica Loew 2 spp., widespread southern Canada and U.S. Body, bristles, and wing pattern otherwise .36 36. Two pairs of frontal bristles present, both black (fig. 75, a); inner vertical bristle black (fig. 75, b); apex of cell r4+5 with small, round hyaline spot (fig. 72, a) .Xanthomyia Phillips 2 spp., widespread Alaska, Canada, and northern U.S. Three or 4 pairs of frontal bristles present (fig. 76, a); otherwise differing .37 37. All frontal and orbital bristles usually whitish to light brown (fig. 76, a, b), not dark brown to black; apex of cell r4+5 usually with a very narrow transverse hyaline band (fig. 73, a) .Jamesomyia Quisenberry 1 sp., eastern Canada, eastern half of U.S. All frontal and orbital bristles dark brown to black; apex of cell r4+5 dark brown, with or without hyaline spot (fig. 74, a) .... Xenochaeta Snow 2 spp., far western U.S. 38. Width of frons at vertex less than or about equal to width of one eye (fig. 77); acrostichal bristles (fig. 78, a) in line with or anterior to anterior

Figures 71-74. Right wings. 71, lcterica seriata (Lw.); 72, Xantbomyia platyptera (Lw.); 73, Jamesomyia geminata (Lw.); 74, Xenocbaeta dicbromata Snow. Figures 75, 76. Lateral view, head. 75, Xantbomyia platypera (Lw.); 76, Jamesomyia geminata (Lw.). Figure 77. Top view, head, Xantbaciura insecta (Lw.). Figure 78. Generalized scutum, dorsal view, showing position of acrostichal bristles. Figures 79, 80. Dorsal view, scutellum. 79, Trupanea wbeeleri Curr.; 80, Tepbritis araneosa (Coq.).

58

59

Key to the U.S. and Canadian Genera edge of wing base (fig. 78, b) .Xanthaciura Hendel 4 spp., southern U.S., Bermuda Not as above.39 39. Only 1 pair of scutellar bristles present (fig. 79) .40 Two pairs of scutellar bristles present, the posterior pair (fig. 80, a) some¬ times quite small .43 40. Cell r4+5 without a bulla (fig. 81); dark color of wing usually forming a stellate pattern in apical half of disk (fig. 81, a) and with proximal half of wing usually hyaline; apex of cell r4 + 5 (fig. 81, b) usually hyaline. .Trupanea Schrank 21 spp., widespread Canada and U.S.; worldwide Cell r4+5 usually with a bulla (fig. 82, a); if bulla indistinct, apex of cell r4 + 5 usually dark brown; apical half of wing disk with or with-out stellate pattern .41 41. Abdomen tomentose; wing hyaline at extreme base (fig. 83, a); apical half of wing with stellate pattern (fig. 83, b); apex of cell r4+5 hyaline (fig. 83, c) .Dyseuaresta Hendel 1 sp., Florida, Texas; other species southward Abdomen polished, at least in part; wing dark or hyaline at extreme base, with or without apical stellate pattern; apex of cell r4+5 hyaline or not .42 42. Apex of cell r4+5 hyaline between arms of a dark V-shaped mark (fig. 84, a) .Valentibulla Foote and Blanc 6 spp., southwestern Canada, widespread western U.S. Apex of cell r4 + 5 dark brown to hyaline without V-shaped mark (fig. 85, a) .Aciurina Curran 11 spp., widespread western U.S. 43. Only 1 pair of dark frontal bristles present (fig. 87, a), usually accompanied by several adjacent white bristles nearly as long as blackish one (fig. 87, b); labellum elongate (fig. 87, c).Oxyna Robineau-Desvoidy 3 spp., widespread western Canada and western U.S. Frontal bristles otherwise .44 44. Three pairs of white to yellowish orbital bristles present (figs. 88, a; 89, a); 3 pairs frontal bristles present (anterior bristle short and pale, 2 posterior bristles strong, widely separated and black) (figs. 88, b; 89, b); face with central black spot (fig. 89, c) .Paracantha Coquillett 5 spp., southern Canada, widespread U.S.; other species southward One or 2 pairs of orbital bristles present, the anterior pair often darker than the posterior pair; otherwise differing .45 45. Wing hyaline with 4 strongly oblique yellow and brown bands (fig. 86); scutum with pair of dark lateral prescutellar spots . .Tomoplagia Coquillett 2 spp., widespread U.S.; many species southward Wing and scutum not thus marked.46 46. Two pairs frontal bristles present (fig. 92, a); if more, then light areas of wing distinctly yellowed, not white hyaline spots that contrast strongly with dark areas.47 Three pairs of frontal bristles present.51

Figures 81-86. Right wings. 81, Trupanea bisetosa (Coq.); 82, Aciurina bigeloviae (Ckll.); 83, Dyseuaresta mexicana (Wied.); 84, Valentibulla thurmanae Foote; 85, Aciurina maculata (Cole); 86, Tomoplagia cressoni Aczel. Figures 87, 88. Lateral view, head. 87, Oxyna palpalis (Coq.); 88, Paracantha culta (Wied.). Figure 89. Front view, head, Paracantha cultaris (Coq.).

60

Figures 90, 91. Right wings. 90, Tephritis araneosa (Coq.); 91, Gonioxyna fuscata Foote. Figure 92. Lateral view, head, Euaresta festiva (Lw.). Figure 93. Posterior view, anal region, male, Euaresta festiva (Lw.). Figure 94. Lateral view, head, Tephritis dilacerata (Lw.). Figures 95, 96. Lateral view, hind leg. 95, Tephritis subpura (Johns.); 96, Paroxyna genalis (Thoms.). Figures 97-100. Right wings. 97, Dioxyna sororcula (Wied.); 98, Paroxyna albiceps (Lw.); 99, Acinia picturata (Snow); 100, Neotephritis finalis (Lw.).

61

62

Key to the U.S. and Canadian Genera

47. Lower facial margin weakly projecting (fig. 92, b); male fore femur swollen; male terminalia broadened, with conspicuous striations in anal region (fig. 93).Euaresta Loew 8 spp., widespread southern Canada and U.S. Characters not as above .48 48. Proboscis never geniculate (fig. 94, a); pterostigma usually mostly dark, never with a central round hyaline spot (fig. 90, a); hind femur usually with 1 preapical dorsal seta (fig. 95, a).Tephritis Latreille 19 spp., Alaska, widespread Canada and U.S.; worldwide Proboscis geniculate, labellum usually elongate (fig. 101, a); hind femur with more than 1 preapical dorsal seta, the distal 2 side-by-side (fig. 96, a), or none at all present.49 49. Costa bowed anteriorly between subcostal break and apex of vein + causing cell r, to be distinctly wider than cell r2+3 at level of apex of pterostigma (fig. 91, a).. Gonioxyna Hendel 1 sp., western Canada, western U.S,; other Neotropical and Palearctic species Costa not distinctly bowed anteriorly between subcostal break and apex of vein R2+35 cells r, and r2+3 about equal in width at level of pterostigma .50 50. Head usually longer than high (fig. 101); pterostigma (fig. 97, a) dark centrally, hyaline at extreme base and apex; basiphallus (fig. 11) without setae.Dioxyna Frey 2 spp., widespread southern Canada and U.S., Bermuda Head rarely longer than high, never markedly so (fig. 102); pterostigma usually with central round hyaline spot (fig. 98, a); basiphallus with setae near tip before distiphallus.Paroxyna Hendel 20 spp., Alaska, widespread Canada and U.S.; many Palearctic and Neotropical species 51. Apex of cell r4+5 dark (fig. 100, a).52 Apex of cell r4+5 with at least a small hyaline spot (fig. 104, a), often more broadly hyaline.53 52. Hyaline spots in anterior half of wing disk bordered with a brown color darker than the brown background of disk; some small yellowish spots also present in disk (fig. 99).Acinia Robineau-Desvoidy 1 sp., widespread U.S.; many species southward and in Palearctic Region Hyaline spots in wing disk not bordered with darker brown; all spots hyaline or nearly so (fig. 100).Neotephritis Hendel 2 spp., widespread southern Canada and U.S.; other species southward 53. Wing disk almost completely bordered with brown, leaving a large, uninter¬ rupted hyaline area occupying cells br, r4+5, and dm (fig. 103) . .Metatephritis Foote 1 sp., western U.S. Wing pattern otherwise.54 54. Hyaline spot in apex of cell r4+5 usually as long as wide, bordered by relatively narrow dark arms terminating at apices of veins R4 + 5 and M (fig. 104, a); face and parafacial without dark spots.55 H\ aline spot in apex of cell r4+3 either part of a more extensive hyaline area

Figures 101-109. Characters of Tephritidae. Figures 101, 102. Lateral view, head. 101, Dioxyna sororcula (Wied.); 102, Paroxyna genalis (Thoms.); Figures 103-109. Right wings. 103, Metatephritis fenestrata Foote; 104, Tetreuaresta sp.; 105, Euarestoides acutangulus (Thoms.); 106, Acrotaenia testudinea (Lw.); 107, Tetreuaresta sp., showing haired vein R4+5 (pattern not shown); 108, Eutreta diana (O.S.); 109, Cryptotreta cislimitensis Steysk.

63

Key to the U.S. and Canadian Genera

or consisting of a very small spot; face or parafacial or both often with dark spots .56 55. Vein R4+5 strongly setulose dorsally (fig. 107, a); dark brown color of wing disk extensive, contrasting markedly with the relatively small, welldefined hyaline markings (fig. 104) .Tetreuaresta Hendel Many Neotropical species, at least 4 in Mexico; may eventually be found in the U.S. Vein R4+5 with only a few small setae near its base; contrast in wing disk colors not as marked as above (fig. 105) .Euarestoides Benjamin 4 spp., widespread southern Canada and U.S.; several species south¬ ward 56. Wing with several large dark bulla-like markings anteriorly and at least 2 transverse hyaline bands in apical third (fig. 106); postocular setae light, concolorous .Acrotaenia Loew 1 sp., Florida; many species southward Wing pattern otherwise; a narrow hyaline area present at wing apex or wing almost entirely dark except for reticulations; postocular setae mixed small black and large white .57 57. Wing dark to extreme base, disk usually with many very small white to yellowish spots; hyaline area at wing apex usually uninterrupted (fig. 108, a), absent or with spots at apices of veins R4 + 5 and M in males of several species.Eutreta Loew 17 spp., widespread Canada and U.S.; many species southward Wing hyaline in basal !A to Vs; disk with relatively large hyaline spots; hyaline area at wing apex usually containing dark spots at apices of veins R4+5 and M (fig. 109, a) .Cryptotreta Blanc and Foote 1 sp., California; at least 1 Mexican species

SYSTEMATIC TREATMENT OF THE GENERA

Genus Acidogona Loew Acidogona Loew 1873: 285 (type species, Trypeta melanura Loew 1873: 283, by original designation); 328 (in key); 331 (in list). — Williston 1896: 122 (in key). — Coquillett 1899c: 268 (in key). - Aldrich 1905: 609 (in catalog). - Williston 1908: 287 (in key). - Hendel 1914b: 94 (in key, world). - V. T. Phillips 1923: 122 (in key; review, e. U.S.). - Johnson 1925b: 264 (N. Engl.). - F. H. Benjamin 1934: 31 (review, Fla.). - Curran 1934: 289 (in key). - Foote 1965a: 663 (in catalog). Wasbauer 1972: 98 (hosts). Acidigona [error]: Loew 1873: 330 (taxonomy). — Coquillett 1910: 502 (type data).

Trypeta (Acidogona): Osten Sacken 1878: 192 (in catalog).

recognition. The wing pattern (fig. 110) of melanura, the sole species, is reticu¬ late, remotely resembling those of Neotephritis spp., but the first flagellomere is long, relatively broad, and distinctly pointed (as in fig. 34), and the abdomen is yellow or light brown with strong dark brown to black markings. Some other species of Teph ritidae, notably those in Rhagoletis and Zonosemata, have a sharply pointed first flagellomere but without exception have banded wings.

discussion. The genus Acidogona is not discussed in the literature to any great extent. It is most closely related to Xenochaeta, several Palearctic genera, and proba¬ bly also Jamesomyia.

65

66

Systematic Treatment of the Genera

Figure 110. Right wing, Acidogona melanura (Lw.).

•

Acidogona melanura

o

Acinia picturata

Map 1. Distribution of Acidogona melanura and Acinia picturata.

67

Acinia Acidogona melanura (Loew) (Figs. 68, 70, 110; Map 1)

Trypeta melanura Loew 1873: 283 (female syntypes, MCZ; D.C.). — Coquillett 1910: 502 (type data).

Acidogona melanura: Loew 1873: 285, 330 (taxonomy). — Aldrich 1905: 609 (in catalog). - Hendel 1914b: 94 (type data). - V. T. Phillips 1923: 142 (review, e. U.S.). - Johnson 1925b: 264 (N. Engl.). - F. H.Benjamin 1934: 32 (review, Fla.). Brimley 1938: 384 (N. Car.). - V. T. Phillips 1946: 105 (hosts). - Hennig 1952: 220 (in catalog, immature stages). — Foote 1965a: 663 (in catalog). — Borror et al. 1974: 516 (illustration, adult). — Wasbauer 1972: 98 (hosts).

Trypeta (Acidogona) melanura: Osten Sacken 1878: 192 (in catalog). Acidogona melaneura [error]: Curran 1934: 292 (illustration, wing). recognition. As for the genus.

DISTRIBUTION. Map 1.

host.

Hieracium

discussion.

is the only known host.

A. melanura, the only known member of the genus, was discussed in

some detail by Benjamin (1934). He described and illustrated the egg, larva, puparium, and adult. The larva feeds singly in an individual flower, which remains unopened until the adult emerges. This species is of no known economic importance.

Genus Acinia Robineau-Desvoidy Acinia Robineau-Desvoidy 1830: 775

(type species, Acinia jaceae Robineau-

Desvoidy 1830: 775, by designation of Rondani 1871a: 4; 1871b: 4) (— Acinia

corniculata (Zetterstedt)). - F. H. Benjamin 1934: 46 (review, Fla.). - Hendel 1935: 53 (taxonomy). - Aczel 1958: 75 (revision, world). - Foote and Blanc 1963: 6 (review, Calif.). - Foote 1964b: 84 (taxonomy). - Foote 1965a: 664 (in catalog).

- Foote 1967b: 17 (in catalog, Neotropical). - Cole and Schlinger 1969: 352 (review, w. U.S.). - Wasbauer 1972: 98 (hosts). - Foote and Freidberg 1980: 29 (type data). Tephritis Latreille: Loew (part) 1873: 302 (taxonomy); 328 (in key); 330 (in list). Aldrich (part) 1905: 193 (in catalog).

Trypeta (Tephritis): Osten Sacken (part) 1878: 193 (in catalog). Euribia: V. T. Phillips (part) 1923: 149 (in key; review, e. U.S.). RECOGNITION. The genus

Acinia is rather unusual among the Tephritinae, having

an entirely yellow head, body, and legs combined with light brown or yellowish bristles and setulae. All species of Acinia have three pairs of frontal and two pairs of

Systematic Treatment of the Genera

scutellar bristles. The single U.S. species, picturata (Snow), is distinctive in the nature of the wing markings (see species discussion).

Acinia contains 15 known species, 6 of which occur in Europe. Except for picturata, which is widespread in North America and the Antilles, the distribution.

remaining species are restricted to South America. discussion. The nomenclatural history of

Acinia and its species is marked by a

great deal of confusion; a rather complete discussion was presented by Benjamin (1934). See Aczel (1958) for a revision of the New World species. The larvae of all the known species develop in the flower heads of composites, but little detail concerning their biology is available.

Acinia picturata (Snow) Sourbush seed fly (Figs. 99, 111; Map 1)

Trypeta fucata Fabricius: Osten Sacken 1858: 79 (in catalog). Tephritis fucata: Loew 1873: 302, 330, 336 (taxonomy). — Johnson 1895a: 280 (distribution). - Johnson 1895b: 337 (distribution). - F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 611 (in catalog). - Johnson 1910: 803 (N.J.). - Johnson 1913b: 84 (Fla.). - Johnson 1930: 151 (Nantucket).

Trypeta (Tephritis) fucata: Osten Sacken 1878: 193 (in catalog). - Johnson 1895a: 280 (Jamaica). - Johnson 1904b: 175 (N.J.).

Tephritis picturata W. A. Snow 1894: 173 (male and female syntypes, UKL; Fla.). — Johnson 1910: 803 (Fla.). - Johnson 1913b: 84 (Fla.). - V. T. Phillips (part) 1923: 150 (synonymy).

Euaresta fucata: Johnson 1903a: 100 (biology). Trypeta picturata: Aldrich 1905: 612 (in catalog). Euribia fucata: Hendel 1914c: 67 (revision). - V. T. Phillips (part) 1923: 150 (re¬ view).

Acinia fucata: F. H. Benjamin 1934: 48 (review, Fla.). - Aczel (part) 1949: 271 (in catalog). - Foote 1962: 177 (taxonomy). - Foote and Blanc 1963: 78 (review, Calif.).

Tephritis picturara (error): Quisenberry 1951: 59 (taxonomy). Acinia picturata: Aczel 1958: 101 (revision). — Foote 1964b: 84 (nomenclature). — Foote 1965a: 664 (in catalog). - Foote 1967b: 57.5 (in catalog). - Stegmaier 1967a: 54 (biology, hosts). - Goeden 1971: 47 (host). - Wasbauer 1972: 98 (hosts). - Sutherland 1978: 35, 44, 98 (common name). - Hardy and Delfinado 1980: 40 (Hawaii).

Trypeta germanna Walker (nomen nudum): Foote 1964d: 325 (?N. A., synonymy). recognition. In addition to the characters discussed for the genus, the nature of

the wing markings (fig. Ill) makes the recognition of picturata quite easy. At least the large hyaline spots in cells br and r4+5 are bordered with a brown color notably

69

Aciurina

Figure 111. Right wing, Acinia picturata (Snow).

darker than the yellowish wing background, and all radial cells contain some yellow¬ ish spots in the same darker brown background. The subapical row of contiguous hyaline spots crossing cells r2+3, r4+5, and m (fig. Ill, a) leave the extreme apices of all these cells dark. Aczel (1958) presents characters which separate picturata from all other American species. hosts.

As far as known, picturata infests only species in the plant genera Plucbea

and Tessaria. distribution.

Map 1. In addition, picturata is known from various localities in

northern Mexico, Jamaica, Puerto Rico, and Antigua. In the early literature, picturata was incorrectly identified by North American authors as the South American fucata (Fabricius), which is a distinct and discussion.

easily recognized species. This situation was explained in some detail by Foote (1964b). Although Benjamin (1934) presented additional data concerning picturata, the biology and economic significance of this species is poorly understood. The puparium of picturata is unusual in being concave and whitish ventrally. This species has been introduced into Hawaii as a biological control agent for Plucbea species.

Genus Aciurina Curran Aciura Robineau-Desvoidy: Coquillett (part) 1899c: 263 (taxonomy). - Aldrich (part) 1905: 607 (in catalog). Aciurina Curran 1932b: 9 (type species, Aciurina trixa Curran 1932b: 11, by origi¬ nal designation). - Curran 1934: 293 (in key). - Blanc and Foote 1961: 76 (partial key to species). - Foote and Blanc 1963: 7 (review, Calif.). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 98 (hosts). - Furniss and Barr 1975: 20 (galls, para¬ sites). - Wangberg 1978b: 39 (biology). - Wangberg 1981: 711 (biology). Freidberg 1984: 129 (gall biology). - Steyskal 1984: 582 (revision). - Dodson 1987a: 607 (biology, hosts). Tepbrella Bezzi: Bates (part) 1935: 103 (revision). - Aczel 1953: 193 (revision, New World). Aciurina belongs to a group of tephritids having the occipital bristles stubby and whitish, a relatively wide frons, the dorsocentral bristles situated between the supra-alars and the transverse suture, only one pair of scutellar bristles, recognition.

Systematic Treatment of the Genera a polished abdomen, and a visible bulla in cell r4+5. The genus closely resembles Valentibulla, from which it can be distinguished by the absence of a dark V-shaped mark embracing a marginal hyaline spot at the apex of cell r4+5 (fig. 113), although in some species of Aciurina, the latter cell may be marginally hyaline in specimens with an extremely reduced brown wing pattern (see figs. 121, 124). Aciurina comprises 11 species in the United States and Mexico; there may be additional species in Mexico and farther south. distribution.

discussion.

Except for a questionable record of Rosaceae for maculata, all spe¬

cies with known hosts are gall formers on plants of the family Asteraceae. Wasbauer (1972) and Steyskal (1984) summarized the published host information, but McArthur et al. (1979), Wangberg (1981), and Dodson (1987a) suggested that some records may be erroneous and that the species are more host-specific than indicated by Wasbauer’s and SteyskaPs lists. Biological observations of various species have been made as noted in the ensuing discussions. Various species of Aciurina were discussed by Curran (1932b) under the generic name Tephrella. Steyskal (1984) synoptically revised Aciurina and briefly but effectively summarized most of the available information; we are indebted to him for much of the information upon which our treatment of this genus is based.

Key to U.S. Species of Aciurina 1. A dark crossband, sometimes interrupted in cell dm, passing through vein r-m, bifurcate anteriorly, 1 branch running through pterostigma, the oth¬ er to costa apicad of vein RT (fig. 112, b); a complete brown crossband running through vein dm-cu without an accessory arm (fig. 112, c); cell c with a median dark area (fig. 112, a) .idahoensis Steyskal Wing largely without such crossbands, but if almost so (as in some speci¬ mens of bigeloviae with reduced pattern), then dark mark in cell c narrow or lacking (figs. 121, a; 124) .2 2. Cell r1 with 3 hyaline spots along costa, the apical one just before the end of vein R2+3 (fig* 113); cell r4+5 uniformly dark brown except for small rounded spot adjacent to vein M close to apical one of 3 separate hyaline spots in cell m (fig. 113, c); bulla large; last section of vein R4+5 strongly bowed anteriorly (fig. 113, b); abdomen wholly shining black .. .trilitura Blanc and Foote Cell rj with only 1 or 2 hyaline spots along costa, or with markings in cells r4 + 5 and m otherwise .3 3. Cell be dark brown, similar to dark areas of other parts of wing (fig. 114, a) ..4 Cell be with at least a central hyaline area, sometimes wholly hyaline, sometimes yellowish and decidedly paler than dark areas of other parts of wing (fig. 119, a) .8 4. Cell r4 + 5 crossed by obliquely transverse preapical hyaline band (fig. 114, b); cell m with a small subapical and large subbasal hyaline indentation,

71

Aciurina

Figures 112-117. Right wings, Aciurina spp. 112, idahoensis Steysk.; 113, trilitura Blanc and Foote; 114, maculata (Cole); 115, lutea (Coq.); 116, aplopappi (Coq.); 117, thoracica Curr.

the latter sometimes V-shaped caused by presence of marginal brown spot (fig. 114, c) .maculata (Cole) Cell r4 + 5 wholly brown (fig. 115, c) or with 1 or 2 small hyaline spots; otherwise differing .5 5. Alula hyaline or yellowish (fig. 115, a); cells m and cuaj each with a very large subbasal hyaline mark and a small subapical hyaline spot (fig. 115, b), the latter sometimes absent in cell cua: .lutea (Coquillett) Alula brown (fig. 116, a), sometimes with central hyaline spot.6

6. Cells r2+ ^ and r4+5 apicad of vein dm-cu with at most 1 paler brown spot in dark brown area; cell cua, with at least 3 hyaline spots in addition to the basal hyaline spot (fig. 116, c); cell m with 2 hyaline indentations (fig. 116, d); anal lobe with 3-5 hyaline spots (fig. 116, b) . .aplopappi (Coquillett) Cells r7 + , and r4 + 5 apicad of vein dm-cu with several round paler brown spots; cell cuaj with 1 or 2 (rarely 3) hyaline spots in addition to the basal hyaline spot (fig. 117, b); cell m with 1 or 2 hyaline indentations or spots (fig. 117, c); anal lobe with 1-3 minute marginal hyaline spots (fig. 117, a) and sometimes 1 similar central spot.7

7. Cell m with a broad, roughly parallel-sided median stripe and subbasal hyaline indentation extending halfway from wing margin to vein M (fig. \\~j, c) .thoracica Curran Cell M with median hyaline stripe either narrowly oval, wedge-shaped, or

72

Systematic Treatment of the Genera

a

Figures 118-124. Right wings, Aciurina spp. 118, mexicana (Aczel); 119, ferruginea (Doane); 120, opaca (Coq.); 121, notata (Coq.); 122—124, bigeloviae (Ckll.).

broken, broadest anteriorly and strongly tapering toward wing margin (fig. 118, a), the subbasal indentation or spot usually lacking, at most short-oval.mexicana (Aczel)

8. Cell r4 + 5 wholly brown or with small rounded hyaline spot near wing tip adjacent to vein M; cell c broadly hyaline distally, with transverse median brown bar (fig. 119, b); cell dm with preapical hyaline spot not extending to veins at sides of cell (fig. 119, c) .ferruginea (Doane) Cell r4+5 with preapical hyaline mark occupying at least half of width of cell (fig. 120, c), sometimes with an additional subbasal hyaline mark (fig. 120, b); cell c with distal dark area (fig. 120, a) or its apex sometimes largely hyaline (figs. 121, b; 122, a).9 9. Cell c brown in distal half (fig. 120, a), a little pale apically in very pale specimens; a uniform parallel-sided hyaline band extending from costa at

73

Aciurina midlength of cell r1 to posterior third of cell dm (fig. 120, b). .opaca (Coquillett) Cell c hyaline at apex, with median transverse brown bar (fig. 122, a), or wholly hyaline (fig. 121, a); if complete transverse brown band is present through vein dm-cu (fig. 121, e), additional brown markings are also present apicad thereof along costa (fig. 121, d) .10 10. Pterostigma along costa no more than 1.5 times as long as its greatest width (fig. 121, c); vein dm-cu nearly straight (fig. 121, e), the lower apical angle of cell dm about 65° (fig. 121, f); wing predominantly hyaline .notata (Coquillett) Pterostigma along costa at least 2.0 times as long as its greatest width (fig. 124, a); vein dm-cu usually bowed apicad (fig. 124, b), the lower apical angle of cell dm seldom less than 90° (fig. 124, c); wing patterns as in figs. 122-124, the dark markings highly variable .... bigeloviae (Cockerell)

Aciurina aplopappi (Coquillett) (Fig. 116; Map 2)

Trypeta (Aciura) aplopappi Coquillett 1894: 72 (male lectotype, NMNH; Los An¬ geles County, Calif.; designated by Steyskal [1984]).

Aciura aplopappi: Aldrich 1905: 607 (in catalog). - Thompson 1907: 71 (biology). - V. T. Phillips 1946: 105 (host).

Aciurina aplopappi: Curran 1932b: 10 (in key). - Foote and Blanc 1963: 7 (review, Calif.). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 98 (hosts). - Steyskal 1984: 588 (revision).

Tepbrella aplopappi: Bates 1935: 107 (review). - Aczel 1953: 194 (in key).

Map 2. Distribution of Aciurina aplopappi, A. bigeloviae, A. idahoensis, A. maculata, A. mexicana, and A. notata.

74

Systematic Treatment of the Genera

recognition.

A. aplopappi, mexicana, and tboracica have been shown by

Steyskal (1984) to resemble each other so closely in details of the aculeus that he placed these three species together in an “aplopappi group.” All three may be recog¬ nized by the largely brown alula (fig. 116, a). In contrast to the other two species in this group, cells r2+3 and r4+5 in aplopappi contain at most only one brown spot (fig. 116), and cell cuat has at least three hyaline spots, one of them basal. distribution.

Map 2. A male has been collected at 9,624 ft. at Sonora Pass,

Calif. Steyskal (1984) recorded aplopappi from Gutierrezia sarothrae and Haplopappus pinifolius. hosts.

A. aplopappi, a little-known species, is restricted to California and Arizona and has been collected to date mainly in March and April. Very little biological information is available (see Thompson 1907). discussion.

Aciurina bigeloviae (Cockerell) (Figs. 82, 122-124; Map 2) Trypetidae sp. of Cockerell 1889c: 324 (gall). Trypeta sp. of Cockerell 1889d: 363 (gall). Trypeta bigeloviae Cockerell 1890a: 75 (March; description of gall only, Westcliff, Colo., no type selected). - Cockerell 1890b: 324 (December; adult). - Ashmead 1890: 427 (sep. p. 25) (biology). - Cockerell 1893a: 112 (description gall, adult). — Cockerell 1893b: 369 (Custer County, Colo). Trypeta bigeloviae var. disrupta Cockerell 1890b: 324 (male holotype, WSUP; West Cliff, Custer County, Colo.). - Cockerell 1893a: 112 (description gall). - Cockerell 1893b: 369 (Custer County, Colo.). - Steyskal 1984: 592 (synonymy). Eurosta (Trypeta) bigeloviae Townsend 1893b: 49 (female holotype, UKL; Colo.), 52 (synonymy). - Foote 1962: 173 (taxonomy). - Byers et al. 1962: 180 (type data). Eurosta bigeloviae: Ashmead 1894: 342 (parasite). - Cockerell 1899: 91 (descrip¬ tion). - Aldrich 1905: 609 (in catalog). — Thompson 1907: 71 (biology). — Felt 1918: 198 (description gall). - Essig 1938: 605 (review, w. N.A.). - Felt 1940: 323 (description gall). - Essig 1958: 605 (review, w. N.A.). Trypeta bigeloviae: C. F. Baker 1895: 174 (description gall). Aciura bigeloviae: Caudell 1902: 37 (gall). - Janes and Thomas 1932: 103 (Utah). Tepbritus (error) bigeloviae: Ballou 1926: 28 (gall). Aciurina trixa Curran 1932b: 11 (female holotype, AMNH; Stansbury I., Utah). Curran 1934: 295 (figure wing). - Knowlton and Harmston 1937: 145 (Utah). Foote and Blanc 1963: 11 (review, Calif.). - Foote 1965a: 670 (in catalog). Wasbauer 1972: 99 (hosts). - Arnaud and Owen 1981: 147 (type data). - Steck 1981: 33 (biology). - Wangberg 1981: 721 (gall, biology). - Steyskal 1984: 592 (synonymy). - Dodson and George 1986: 63 (biology). - Dodson 1987b: 494 (biology).

75

Aciurina Tephrella Tepbrella Tephrella Tephrella

bigeloviae: Bates 1935: 110 (review). bigeloviae var. disrupta: Bates 1935: 110 (taxonomy). trixa: Bates 1935: 110 (review). semilucida Bates 1935: 111 (female holotype, NMNH; Riparia, Wash.). —

Steyskal 1984: 592 (synonymy). Euribia bigeloviae: Phillips 1946: 114 (hosts). Aciurina bigeloviae: Foote and Blanc 1963: 8 (review, Calif.). — Foote 1965a: 670 (in catalog). — Sabrosky 1971: 85 (correction, N.A. catalog). — Wasbauer 1972: 98 (hosts). — Wangberg 1977: 237 (parasite). — McArthur et al. 1979: 81 (galls). Wangberg 1981: 730 (taxonomy). — Steyskal 1984: 592 (revision). — Dodson and George 1986: 63 (biology). Aciurina bigeloviae var. disrupta: Foote 1965a: 670 (in catalog). Aciurina semilucida: Foote 1965a: 670 (in catalog). — Novak et al. 1967: 148 (host, gall). - Wasbauer 1972: 99 (hosts). - Wangberg 1981: 719 (biology). Urophora sabroskyi Steyskal 1979: 55 (male holotype, NMNFf; Wawawai, Garfield County, Wash.). - Norrbom 1989: 64 (taxonomy, synonymy). As recognized by Steyskal (1984), bigeloviae has a more variable wing pattern than any other species of Aciurina. The wing pattern ranges from predominantly dark brown to largely hyaline with many intermediate forms (figs. recognition.

122-124). However, very little variation exists in the structure of the male and female postabdomen. In bigeloviae, regardless of the type of wing pattern, a subapical hyaline spot is always present in cells r4+5 and m, almost always broadly con¬ nected across vein M but sometimes obliterated by the coalescence of hyaline areas. In this respect, it is somewhat similar to opaca, maculata, and trilitura, but it may be distinguished from those species by the characters given in the key.

DISTRIBUTION.

Map 2.

A. bigeloviae was orginally described from white woolly galls on Bigelovia (now Chrysothamnus) in Colorado, and it is commonly found on C. nauseosus within its range. Steyskal (1984) recorded it from Artemisia, Bebbia, and two other species of Chrysothamnus, and Wasbauer (1972) listed it from Haplopappus. However, Dodson (1987b) questioned the accuracy of non-Chrysothamnus records for this species (as trixa). The galls are so distinctive that little confusion hosts.

exists concerning the insects to which they give rise. A. bigeloviae is the most widely distributed and commonly encoun¬ tered of the Chrysothamnus-dwelling Aciurina species. Dodson and George (1986) considered trixa and bigeloviae distinct species, or at least host races, based on ecological and biochemical differences. If they are separate species, bigeloviae as recognized by Steyskal may be a complex of several sibling species. The names synonymized by Steyskal and Norrbom remain available, and some may be resur¬ discussion.

rected based on future biological investigations.

76

Systematic Treatment of the Genera

Map 3. Distribution of Aciurina ferruginea, A. lutea, A. opaca, A. thoracica, and A. trilitura.

Aciurina ferruginea (Doane) (Fig. 119; Map 3) Aciura ferruginea Doane 1899: 182 (holotype, sex not known, WSUP; Pullman, Wash.). - C. F. Baker 1904: 30 (Nev.). - Aldrich 1905: 607 (in catalog). — Janes and Thomas 1932: 103 (Utah). — Foote 1966b: 123 (type data). — Zack 1984: 31 (type data). Aciurina ferruginea: Curran 1932b: 10 (in key). - Foote and Blanc 1963: 8 (review, Calif.). — Foote 1965a: 670 (in catalog). - Tauber and Tauber 1967: 907 (re¬ productive behavior, biology). - Tauber and Tauber 1968: 553 (description gall). Wasbauer 1972: 99 (hosts). - Wangberg 1977: 238 (parasite). - Steck 1981: 19 (host). - Wangberg 1981: 712 (biology). - Steyskal 1984: 593 (revision). Tepbrella ferruginea: Bates 1935: 109 (review). recognition.

The wing pattern of ferruginea is not as distinctive as some other

species of Aciurina, and care must be taken to prevent confusion. The species is somewhat similar to opaca, from which it is distinguished by the lack of a connection between the hyaline spots in cells rl and dm (fig. 119). Use of the remaining key characters will make precise identification possible. DISTRIBUTION.

Map 3.

Steyskal (1984) recorded ferruginea from Chrysothamnus, Haplopappus, and Solidago. hosts.

77

Aciurina discussion.

Steyskal (1984) described the aculeus of ferruginea in detail. This

species apparently occurs in the adult stage from July through September in many parts of its range, later in the season than is usual for species in this genus. Tauber and Tauber (1967, 1968) and Wangberg (1981) discussed its biology in some detail.

Aciurina idahoensis Steyskal (Fig. 112; Map 2) Aciurina sp. A of Wangberg 1981: 724 (biology). — Steyskal 1984: 595 (revision). Aciurina sp. B of Wangberg 1981: 111 (biology). — Steyskal 1984: 595 (revision). Aciurina idahoensis Steyskal 1984: 594 (male holotype, NMNH; Murphy, Idaho). The unique wing pattern (fig. 112) enables one to recognize ida¬ hoensis immediately. It exhibits a central brown transverse wedge-shaped bar ante¬ riorly surrounding the hyaline spot immediately apicad of cell sc, thus forming a distinct Y. This bar extends posteriorly to at least the middle of cell dm, rarely to the hind wing margin where it may coalesce with the brown apical bar adjacent to it. The latter completely crosses the wing and is always connected anteriorly to the dark mark along the costa apical to it. With the exception of opaca and some specimens of bigeloviae with scanty brown wing markings, it is the only Aciurina species in which the second hyaline costal spot apicad of the pterostigma extends uninterrupted into recognition.

cell dm or beyond (see also figs. 120, 123). DISTRIBUTION.

host.

Map 2.

Chrysothamnus viscidiflorus is the only host recorded by Steyskal (1984).

discussion.

Found to date only in Idaho, idahoensis has been captured from late

April through July. Steyskal (1984) indicated a probability that the “species” named “A” and “B” by Wangberg (1981) are conspecific with idahoensis. Wangberg de¬ scribes the hosts, galls, and biology of those two forms in considerable detail.

Aciurina lutea (Coquillett) (Fig. 115; Map 3) Aciura lutea Coquillett 1899c: 264 (female holotype, NMNFT. Pareah, Utah). - C. F. Baker 1904: 30 (Nev.). - Aldrich 1905: 607 (in catalog). Aciurina lutea: Curran 1932b: 10 (in key). - Foote 1965a: 670 (in catalog). Wangberg 1981: 715 (biology). - Steyskal 1984: 595 (revision). Tephrella lutea: Bates 1935: 107 (review). recognition.

A. lutea is another quite distinctive Aciurina species in having two

hyaline areas in each of cells cuaj and m, the basal one very large and broad and the apical one very small and inconspicuous (fig. 115, b). In the former cell, the small

Systematic Treatment of the Genera subapical spot is sometimes lacking. The alula is entirely hyaline or yellowish in strong contrast to the brown color of the wing.

DISTRIBUTION.

Map 3.

Cbrysothamnus viscidiflorus, the only known host, was recorded by Steyskal (1984). host.

Wangberg (1981) presented an extensive discussion of many aspects of the biology of lutea as it occurs in Idaho. discussion.

Aciurina maculata (Cole) (Figs. 85, 114; Map 2) Aciura maculata Cole in Cole and Lovett 1919: 252 (male holotype, CAS; Jackson County, Oreg.). - Cole and Lovett 1921: 325 (Oreg.). - V. T. Phillips 1946: 105 (host). - Arnaud 1979: 329 (type data). Aciurina maculata: Curran 1932b: 10 (in key). - Foote and Blanc 1963: 9 (review, Calif.). - Foote 1965a: 670 (in catalog). - Novak et al. 1967: 148 (host, gall). Wasbauer 1972: 99 (hosts). - Wangberg 1977: 238 (parasite). - McArthur et al. 1979: 81 (galls). - Steck 1981: 33 (biology). — Wangberg 1981: 716 (biology). Steyskal 1984: 595 (revision). Aciurina pacifica Curran 1932b: 10 (female holotype, AMNH; Yakima, Wash.). Bates 1935: 108 (synonymy). - Arnaud and Owen 1981: 149 (type data). Tephrella maculata: Bates 1935: 108 (review).

As in most specimens of bigeloviae, which maculata most closely resembles, the apical spot in cell m continues anteriorly across vein M to end in a rather extensive hyaline area in the outer third of cell r4+5 (fig. 114, b). From recognition.

bigeloviae, this species is separated most readily by the nearly uninterrupted dark brown color of cell be (compare figs. 114, a and 122, 124). A. maculata also closely resembles lutea (fig. 115), from which it is separated by the different arrangement of hyaline areas in cell cuaj. Cell r4+5 of lutea is entirely brown (fig. 115, c). DISTRIBUTION.

A. maculata has been recorded an Amelancbier species.

HOSTS. tionably

Map 2.

from

Cbrysothamnus, Senecio,

and ques¬

Various aspects of the biology of maculata have been discussed by Novak et al. (1967), Steck (1981), and Wangberg (1977, 1981). Adults have been collected throughout its range from March to early July. discussion.

79

Aciurina

Aciurina mexicana (Aczel) (Fig. 118; Map 2) Tephrella mexicana Aczel 1953: 194 (male holotype, NMNH; Nogales, Mex.). Aciurina mexicana: Foote and Blanc 1979: 161 (taxonomy). — Steyskal 1984: 590 (revision). — Jenkins 1990: 66 (mating behavior). Structurally, mexicana resembles aplopappi and thoracica more closely than any other Aciurina. Steyskal (1984) demonstrated similarities in the aculeus of all three species. A. mexicana is unique in the genus in that most of the wing is dark. The hyaline spot in cell m (fig. 118, a), which is relatively narrow or divided near its center into two separate spots, is the principal character by which it can be distinguished from thoracica (see also the recognition characters for aplo¬ recognition.

pappi). DISTRIBUTION.

host.

Map 2.

The only known host is Baccharis sarothroides (Steyskal 1984).

discussion.

A. mexicana, an essentially Mexican species, has rarely been found

in large numbers, nor has it been collected often. Adults have been found in Arizona and California during the latter half of April through the first half of June. Jenkins (1990) described its mating behavior.

Aciurina notata (Coquillett) (Fig. 121; Map 2) Trypeta notata Coquillett 1899c: 262 (female syntypes, NMNH; Albuquerque, N. Mex.). - Cockerell 1900a: 198 (description gall). - Aldrich 1905: 605 (in catalog). - Thompson 1907: 71 (biology). - V. T. Phillips 1946: 127 (host). - Essig 1958: 604 (description gall). Tephrella notata: Bates 1935: 112 (review). Aciurina notata: Foote 1960b: 254 (nomenclature). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 99 (hosts). - Steyskal 1984: 596 (revision). RECOGNITION.

A. notata is quite distinctive in wing pattern in that a large share

of the wing is hyaline rather than brown. It is similar to some specimens of bigeloviae in having a reduced wing pattern, but the pterostigma is much shorter in comparison with its width than in bigeloviae (compare figs. 121, c and 122, b).

DISTRIBUTION.

host.

Map 2.

Chrysothamnus nauseosus has been recorded by Steyskal (1984).

80

Systematic Treatment of the Genera

A rare species apparently restricted to New Mexico where it has been found during April, May, and June. No biological studies of notata have been reported. discussion.

Aciurina opaca (Coquillett) (Fig. 120; Map 3) Aciura opaca Coquillett 1899c: 263 (female holotype, NMNH; Elko, Nev.). — Aldrich 1905: 607 (in catalog). Acidia (?) johnsoni Thomas 1914: 426 (female holotype, MCZ; Colo.). - Foote 1965a: 670 (synonymy). Acidia johnsoni: Bates 1934a: 295 (taxonomy). Aciurina opaca: Curran 1932b: 10 (in key). - Foote 1965a: 670 (in catalog). Wangberg 1981: 718 (biology). - Steyskal 1984: 596 (revision). Tephrella johnsoni: Bates 1935: 106 (review). Tephrella opaca: Bates 1935: 106 (review). Aciurina johnsoni: Foote 1960b: 254 (generic transfer). A. opaca is distinguished from all other Aciurina by its unique wing pattern, in which the apical end of cell c is broadly brown (fig. 120, a), the usual recognition.

hyaline spot in cell

at the apex of the pterostigma is usually absent, and a uniform

parallel-sided band extends from the costa at the midlength of cell r1 to the lower third of cell dm (fig. 120, b). Among the species of Aciurina having darkly colored wings, opaca also is distinctive in having the anal lobe and the cells basad and anteriad to it almost completely hyaline. Steyskal (1984) described its unique rounded aculeus tip, which indicates that its oviposition habits may be somewhat different from those of most other American fruit flies. DISTRIBUTION.

host.

Map 3.

Steyskal (1984) recorded only Chrysothamnus nauseosus as a host.

A. opaca, a relatively rare species, is spread over much of the western United States, where it has been found from March through June. discussion.

Aciurina thoracica Curran (Fig. 117; Map 3) Aciurina thoracica Curran 1932b: 11 (female holotype, AMNH; San Diego County, Calif.). — Tilden 1951: 160, 167 (host). — Foote and Blanc 1963: 9 (review, Calif.). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 99 (hosts). - Arnaud and Owen 1981: 147 (type data). - Steyskal 1984: 592 (revision). - Goeden 1986: 326 (host). Tephrella thoracica: Bates 1935: 108 (review). - Aczel 1953: 194 (in key).

81

Acrotaenia A. tboracica is distinguished from aplopappi and mexicana, the

recognition.

two species of Aciurina most closely resembling it, by the presence of more than one pale brown spot in cells r2+3 and r4+5 in combination with the presence in cell m of a broad roughly parallel-sided median stripe and a subbasal hyaline indentation ex¬ tending halfway from the wing margin to vein M (fig. 117, c). Steyskal (1984) described details of the aculeus. See also the recognition section of aplopappi. DISTRIBUTION.

HOSTS.

Map 3.

Three species of Baccharis were recorded by Steyskal (1984).

discussion.

A. thoracica is fairly widespread in the western United States, and

adults have been collected from late February into early August. Almost nothing is known about its biology.

Aciurina trilitura Blanc and Foote (Fig. 113; Map 3) Aciurina trilitura Blanc and Foote 1961: 75 (female holotype, CDFA; Camp Baldy Road, San Bernardino County, Calif.). - Foote and Blanc 1963: 10 (review, Calif.). -Foote 1965a: 670 (in catalog). - Wasbauer 1970: 50 (type data). - Arnaud 1979: 329 (type data). - Steyskal 1984: 596 (revision). A. trilitura may be recognized immediately by the presence of three hyaline spots along the costa distad of the pterostigma in cell rl5 the central one crossing vein R?+3 and continuing into cell r2+3 (fig* 113). A small subapical spot is always present in cell r4+3 contiguous with, or only narrowly separated from, the apical-most of the three hyaline spots in cell m (fig. 113, c). The bulla is prominent RECOGNITION.

and is bordered anteriorly by the strongly bowed vein R4+5DISTRIBUTION.

host.

Map 3.

Cbrysotbamnus nauseosus is the only recorded host, according to Steyskal

(1984). discussion.

Originally thought to be restricted to California, trilitura is now

known to occupy a much wider range. Adults have been collected in California from late April until early June.

Genus Acrotaenia Loew Acrotaenia Loew 1873: 274 (type species, Trypeta latipennis Wiedemann, by desig¬ nation of Wulp 1899: 914); 328 (in key). - Williston 1896: 122 (in key). - Aldrich

82

Systematic Treatment of the Genera

1905: 608 (in catalog). - Williston 1908: 287 (in key). - Coquillett 1910: 503 (type data). - Hendel 1914b: 94 (in key). - V. T. Phillips 1923: 122 (in key). Curran 1932b: 4 (in key). - Bates 1934c: 1 (revision). - Curran 1934: 291 (in key). — Foote 1960h: 107 (Fla., review). — Foote 1965a: 664 (in catalog). Trypeta (Acrotaenia): Osten Sacken 1878: 191 (in catalog). The wing in all Acrotaenia species is exceptionally broad in rela¬ tion to its width, and the wing pattern (fig. 125) is unlike that of any other genus in recognition.

the United States or Canada. The combination of the distinctive basal reticulation and apical bands, and usually very extensive dark marks anterobasally, distinguish this genus from all others known in the New World. Acrotaenia is a tropical and subtropical genus, occurring from northern Mexico (Durango) south to Brazil and in most of the Antilles. distribution.

Acrotaenia testudinea (Loew) (Figs. 106, 125) Trypeta testudinea Loew 1873: 111 (female lectotype here designated, ZMHU; Cuba). - Coquillett 1910: 503 (type data). Acrotaenia testudinea: Loew 1873: 274, 329 (taxonomy). - Bates 1933b: 164 (tax¬ onomy). - Foote 1960h: 107 (review). - Foote 1965a: 664 (in catalog). The wing pattern of testudinea (fig. 125) is unique among all fruit fly species occurring in the United States and Canada. No other wing pattern consists recognition.

of a combination of dark apical stripes on a light field, basal hyaline spots in a dark field, and a series of black, bullalike spots in the vicinity of the pterostigma. distribution.

Not mapped. A female of testudinea was trapped in 1936 on Big

Pine Key, Monroe County, Fla. This is the first and only known record of the species in the United States despite the extensive use of fruit fly traps in Florida and other southern states since 1936. It is recorded principally from the Greater Antilles, but the total extent of its distribution is not known. Not known. Other Acrotaenia species have been reared from flowers of Otopappus and Wedelia (Asteraceae). host.

83

Anastrepha A. testudinea is encountered rarely in fruit fly collections. The lectotype has a blue label stating only “Cuba, Otto” and an accession label stating discussion.

“2563.

55

Genus Anastrepha Schiner Anastrepha Schiner 1868: 263 (type species, Dacus serpentinus Wiedemann 1830: 521, by original designation). — Aldrich 1905: 601 (in catalog). — Williston 1908: 283 (in key). - Bezzi 1909: 280 (revision). - Coquillett 1910: 506 (type data). Hendel 1914b: 84 (in key). - V. T. Phillips 1923: 120 (in key). - Curran 1932b: 3 (in key). - Curran 1934: 287 (in key). - C. T. Greene 1934: 127 (revision). - Lima 1934: 487 (hosts). - Stone 1942a: 11 (revision). - Foote and Blanc 1963: 11 (review, Calif.). - Foote 1965a: 673 (in catalog). - Sabrosky 1967: 121 (correc¬ tion, N.A. catalog). — Wasbauer 1972: 100 (hosts). — Steyskal 1977a: 75 (taxon¬ omy, synonymy). - Steyskal 1977b: 1 (key to known species). - Norrbom 1985: 1 (phylogeny, taxonomy, hosts, key to species). - Norrbom and Kim 1988b: 1 (species groups, hosts). - Maddison and Bartlett 1989: 30 (zoogeography). Norrbom and Foote 1989: 15 (review, taxonomy, zoogeography). - Steck et al. 1990: 333 (larval identification, key to larvae). - Hernandez 1992: 1 (revision of Mexican species). Acrotoxa Loew 1873: 227 (type species, Dacus fraterculus Wiedemann 1830: 524, designated by Bezzi 1909: 280). - W. A. Snow 1895: 117 (taxonomy). - Williston 1896: 121 (in key). - Coquillett 1899c: 259 (synonymy). - Coquillett 1910: 503 (type data). Trypeta (Acrotoxa): Osten Sacken 1878: 189 (in catalog). Anastrepha (Pseudodacus) Hendel 1914b: 97 (type species, Anastrepha daciformis Bezzi 1909: 283, by original designation). Pseudodacus: Stone 1939a: 282 (revision). — Foote 1965a: 673 (in catalog). — Steyskal 1977a: 77 (synonymy). Phobema Aldrich 1925: 7 (type species, Phobema atrox Aldrich 1925: 7, by original designation). - Steyskal 1977a: 77 (synonymy). Lucumaphila Stone 1939b: 340 (type species, Lucumaphila sagittata Stone 1939b: 347, by original designation). - Aczel 1949: 233 (in catalog). - Foote 1965a: 673 (in catalog). - Steyskal 1977a: 77 (synonymy). RECOGNITION.

Anastrepha is a genus of relatively large yellowish to orange-

brown flies with pale white and limited brown to black body markings. The dorsocentral bristles (fig. 22, b) are situated slightly anterior to the level of the acrostichal bristles (fig. 22, c) but not farther forward than halfway between the latter and the supra-alar bristles (fig. 22, a). The aculeus and oviscape are relatively long, the tip of the aculeus having value for the recognition of many species. In all species, the apical fourth to fifth of vein M is curved anteriorly, and in most species (all those treated herein), joins the apical end of the costa in a smooth, sweeping curve (fig. 24, a).

Systematic Treatment of the Genera Steyskal (1977a) synonymized the genera Lucumapbila Stone, Phobema Aldrich, and Pseudodacus Hendel with Anastrepba. Because these and some species of Toxotrypana are the only other New World taxa with vein M strongly curved forward apically, any species with that character and not having the distinctive features of Toxotrypana may invariably be recognized as an Anastrepba species. discussion. Because many species of Anastrepba were described from trapped

specimens, the majority from Panama and Brazil, very limited host and distributional data are available for the genus. Several of the most economically important species have extensive host lists, but no hosts are known for more than half of the species. The immature stages cannot be identified reliably except under special circumstances because they have been described for relatively few species. Although morphological characters presumably have been worked out for the larvae of several economically important species, there is no assurance that larvae of others are not similar in appearance, host preference, and distribution. Moreover, continuing studies demon¬ strate that a greater degree of morphological variation exists in the larvae of known species than was originally anticipated. At present there are few reliable morphologi¬ cal characters for the recognition of larvae of most species of Anastrepba. Steck et al. (1990) discussed methods for the identification of larvae in this genus, including the use of discriminant analysis, and present a key for the identification of the larvae of 13 species based mainly on Neotropical material. Males of most species in the genus cannot be identified unless they possess distinc¬ tive nongenitalic characters because, historically, mainly female genitalic characters have been used for interspecies comparisons. Scattered evidence indicates that geni¬ talic characters are present that hold promise for species recognition of males in the future (Korytkowski and Ojeda 1968, Norrbom 1985, Norrbom and Kim 1988a). The genus is confined to the New World, occurring from Florida and Texas south to Uruguay and Argentina and in most of the islands of the Caribbean. Occasional introductions have been reported in southern California. Only 20 species have been found in the United States, some of them only rarely. The genus attains its greatest development in the tropical areas of its range; no fewer than 60 species have been discovered in a small area of Panama alone. Evidence indicates that similar numbers of species will be found in other tropical areas such as Chiapas, Mexico, where a very wide variety of host species is present. Very probably, several previously undiscovered species also will be found in re¬ stricted island habitats in the Caribbean. One of these, stonei Steyskal (1977a), which was described from the Bahamas, has been collected in southern Florida (this record was made known to us too late for inclusion in the formal part of this handbook). Anastrepba is probably the most economically important fruit fly genus in the New World, principally because of the host relationships of a handful of the approxi¬ mately 190 known species. A few of the most significant are distincta Greene, ludens (Loew), fraterculus (Wiedemann), obliqua (Macquart), serpentina (Wiedemann), striata Schiner, and suspensa (Loew). Additional species attack other economically important fruit in localized situations (Norrbom and Kim 1988b), and outbreaks of

Anastrepba species not known to be important to agriculture are being discovered regularly in various localities throughout the New World. In the United States, ludens and sus-

pensa are the two species of primary significance. See the discussions of individual species for additional details.

Key to Females of U.S. Species of Anastrepha The males of many species in this genus cannot be identified at the species level. To identify the females of many of the species with confidence, one must examine microscopic features of the aculeus and its apex (see figs. 147—156). Directions for freeing the aculeus from the oviscape are given in the introductory section on tech¬ niques. 1. Wing pattern with middle part of S band absent or separated from basal and anteroapical parts (figs. 127, 128); posterior orbital bristle absent; sides of scutellum with basal third dark; aculeus tip simple, no more than minutely serrate (fig. 147, a), shaft no more than 0.04 mm wide .... 2 Wing pattern with S band uninterrupted (fig. 129, a) and completely cross¬ ing cell dm; posterior orbital bristle usually present; sides of scutellum with at most the extreme base dark .3 2. Dark markings of thorax confined to posterior quarter of scutum (fig. 157, a); r-m and middle of cell dm covered by spot; C band and apical part of S band at least partially interrupted by hyaline spot on costal margin imme¬ diately beyond pterostigma (fig. 127, a) .pallens Coquillett Scutum and pleura marked heavily with dark brown (fig. 158); area around r-m and middle of cell dm hyaline; C band and apical part of S band completely fused along costal margin (fig. 128, a).bicolor (Stone) 3. Wing pattern predominantly dark brown; distal arm of V band (fig. 126) reduced, not joining proximal arm at anterior end of latter (fig. 129, b), or absent (fig. 130, a); mediotergite (fig. 164, a) mostly dark brown or black; abdomen bicolored, reddish brown and yellow.4 Wing pattern not with this combination of characters; mediotergite usually dark only laterally or entirely yellow; abdomen entirely yellowish. .5 4. Distal arm of V band present, separated from proximal arm (fig. 129, c) or narrowly joined to latter at vein M; scutum without dark markings except along scutoscutellar suture; abdominal syntergite1+2 and tergites 3 and 4 with sometimes medially interrupted dark bands apically . .ocresia (Walker) Distal arm of V band absent (fig. 130, a); scutum usually largely dark brown; abdomen brown except for yellowish T-shaped medial mark .. .serpentina (Wiedemann) 5. Scutum with broad, dark brown dorsocentral stripes sometimes interrupted at transverse suture, these stripes with 1 pre- and 1 postsutural area without microtrichia but with black pile strongly contrasting with the dense microtrichia and hoary white pile bordering them medially (best viewed from front) (fig. 159, a); aculeus tip at least 0.19 mm wide; wing pattern as in fig. 131.striata Schiner

costal band

s band

a

Figures 126—135. Right wings, Anastrepha spp. 126, schematic showing terminology of wing bands; 127, pallens Coq.; 128, bicolor (Stone); 129, ocresia (Walker); 130, serpentina (Wied.); 131, striata Schiner; 132, dentata (Stone); 133, sagittata (Stone); 134, edentata Stone; 135, ludens (Lw.).

86

Anastrepha Scutum evenly microtrichose and without dark markings except sometimes on posterior V4; aculeus tip rarely this wide .6 6. Aculeus extremely long and slender, needlelike; tip (fig. 147, a) less than 0.06 mm long; eversible membrane with all dorsobasal scales small and weakly sclerotized; alula without microchaetae.7 Aculeus usually shorter in relation to its width; tip more than 0.09 mm long; eversible membrane with large, stongly sclerotized, hooklike dor¬ sobasal scales; microchaetae usually present on alula .8 7. Posterior margin of scutum with 2 or 3 distinct black spots (fig. 160, a); wing pattern as in fig. 132.dentata (Stone) Posterior margin of scutum usually with brown transverse band, without discrete spots (fig. 161, a); wing pattern as in fig. 133. .sagittata (Stone) 8. Costal and S bands broadly separated along vein R4 + 5 by hyaline area (fig. 134, a); tip of aculeus not serrate (fig. 149, a) .edentata Stone Costal and S bands joined along vein R4 + 5, or tip of aculeus serrate .... .9 9. Aculeus tip with lateral serrations extending at least to level of apex of oviduct and usually somewhat more basad (fig. 153, a, b) .10 Aculeus tip without lateral serrations, or, if serrations present, not extend¬ ing to level of end of oviduct (fig. 154, a, b).12 10. Scutum with at most a diffuse irregular brownish spot; aculeus tip elongate (fig. 153) .limae Stone Scutum yellow-brown except for a strong, sharply defined, oval or circular black median spot on scutoscutellar suture (fig. 162, a); aculeus tip broadly spatulate with very fine serrations (fig. 152, a) .11 11. Hyaline area on costa beyond pterostigma always crossing vein R2+3 and nearly always touching vein R4+5 (fig. 140, b) .spatulata Stone This hyaline area never crossing vein R2+3 and usually not reaching it (fig. 141, b); aculeus tip as in fig. 152 .interrupta Stone 12. Posterior margin of scutum with dark brown band (as in fig. 161, a); aculeus not more than 2.0 mm long, the tip not serrate; wing as in fig. 145 .nigrifascia Stone Posterior margin of scutum without dark brown band; aculeus length vari¬ able; tip at least weakly serrate; sides of mediotergite or subscutellum often dark .13 13. Aculeus tip with at least 22 serrations per side (fig. 151, a); wing pattern as in fig. 137.chiclayae Greene Aculeus tip with at most 20 serrations per side (fig. 155, a) .14 14. Aculeus no more than 2.0 mm long, the tip with distinct serrations (visible at 60x magnification) on apical half or more.15 Aculeus more than 2.0 mm. long, the tip with fine serrations (not visible at 60x magnification), or with serrations confined to apical 2/s.18 15. Medial whitish vitta on scutum broad anteriorly; scutoscutellar suture with¬ out dark spot; mediotergite usually with lateral dark stripe but sub¬ scutellum entirely yellow; aculeus 1.30-1.60 mm long, tip 2A to 3A serrate (fig. 154, b); wing as in fig. 143 .obliqua (Macquart) Medial whitish vitta on scutum narrow anteriorly; scutoscutellar suture often with medial dark spot; mediotergite and subscutellum usually both

Figures 136—145. Right wings, Anastrepha spp. 136, ludens (Lw.), variant; 137, chiclayae Greene; 138, suspensa (Lw.); 139, fraterculus (Wied.); 140, spatulata Stone; 141, interrupta Stone; 142, limae Stone; 143, obliqua (Macq.); 144, zuelaniae Stone; 145, nigrifascia Stone.

88

Figure 146. Right wing, Anastrepha distincta Greene. Figures 147-156. Aculeus tips, Anastrepha spp. (redrawn from Stone 1942a). 147, bicolov (Stone); 148, dentcitu (Stone); 149, edentata Stone; 150, ludens (Lw.); 151, chiclayae Greene; 152, interrupta Stone; 153, limae Stone; 154, obliqua (Macq.); 155, suspensa (Lw.); 156, fraterculus (Wied.).

89

90

Systematic Treatment of the Genera

Figures 157-163. Dorsal view, patterns of scutum and scutellum of Anastrepha spp. 157, pallens Coq.; 158, bicolor (Stone); 159, striata Schin.; 160, dentata (Stone); 161, sagittata (Stone); 162, suspensa (Lw.); 163, ludens (Lw.). Figures 164, 165. Posterior view, subscutellum and mediotergite, Anastrepha spp. 164, serpentina (Wied.); 165, ludens (Lw.).

with lateral dark stripe or both entirely yellow; aculeus length variable, but if less than 1.60 mm long, tip no more than 3A serrate.16 16. Aculeus 1.65-1.85 mm long, tip nearly as broad at base of serrations as at apex of genital opening; subscutellum and mediotergite entirely yellow; scutoscutellar suture without medial dark spot; wing as in fig. 144_ .zuelaniae Stone

Anastrepha Aculeus 1.45-1.95 mm long, tip narrower at base of serrations than at apex of genital opening; subscutellum and mediotergite usually with lateral dark stripe; scutoscutellar suture usually with at least a weak medial dark spot..17 17. Apical part of S band broad, ending near or posterior to apex of vein M (fig. 138, c); aculeus 1.45-1.60 mm long; scutoscutellar suture usually with a large, distinct medial dark spot (fig. 162, a); southern Florida, West Indies .suspense (Loew) Apical part of S band narrow, usually ending anterior to apex of vein M (fig. 139); aculeus 1.50-1.95 mm long; scutoscutellar suture usually with a small, often weak spot; southern Texas to Argentina . .fraterculus (Wiedemann) 18. Aculeus shorter than mesonotum (at least in U.S. and Mexican popula¬ tions); color of mediotergite variable, but dark area on subscutellum, if present, usually no wider than area on mediotergite; wing as in fig. 146 .distincta Greene Aculeus longer than mesonotum; sides of subscutellum with dark spot usu¬ ally broader than dark area extending down onto mediotergite (fig. 165, b).ludens (Loew)

Anastrepha bicolor (Stone) (Figs. 128, 147, 158; Map 4)

Pseudodacus bicolor Stone 1939a: 288 (female holotype, NMNH; Edinburg, Tex.). Aczel 1949: 236 (in catalog). - Foote 1965a: 674 (in catalog).

Anastrepha bicolor: Steyskal 1977b: 3 (in key). - Norrbom 1985: 119, 279 (taxon¬ omy, in key). recognition. As in

pallens, the wing pattern of bicolor differs from that of

“typical” Anastrepha in having an incomplete S band (compare figs. 126, 128). The middle part of this band is completely absent in bicolor, and the anteroapical part along the costa from vein Kx to the wing apex is completely fused with the costal band without a hyaline spot apicad of the pterostigma. Cell dm (except along vein dm-cu) and cell br are hyaline, and the distal arm of the V band is completely absent. In contrast to pallens, the scutal and pleural patterns are heavily dark brown rather than yellow or orange-brown (fig. 158). The shaft of the aculeus is extremely narrow in proportion to its length and terminates in a simple unserrated tip (fig. 147) (see recognition characters for dentata). Of the species occurring in the United States,

bicolor is one of four in which the distiphallus is absent in males, the others being dentata, pallens, and sagittata. DISTRIBUTION. Map 4. In areas south of the United States,

from Mexico.

host. Not known.

bicolor is known only

92

Systematic Treatment of the Genera

• Anastrepha bicolor o Anastrepha nigrifascia

• Anastrepha dentata o Anastrepha interrupta ▲ Anastrepha ocresia

Map 4. Distribution of Anastrepha bicolor, A. dentata, A. edentata, A. interrupta, A. limae, A. nigrifascia, A. ocresia, and A. sagittata.

discussion.

A. bicolor is one of two U.S. species previously assigned to the genus

Pseudodacus Hendel because of its aberrant wing pattern, needlelike aculeus, and other characters.

Anastrepha chiclayae Greene (Figs. 137, 151; Map 5)

Anastrepha chiclayae Greene 1934: 167 (female holotype, NMNH; Hacienda Ouefe, Chiclayo, Peru). - Stone 1942a: 41 (revision). - A. C. Baker et al. 1944: 140 (review, biology). - Aczel 1949: 203 (in catalog). - Foote 1965a: 673 (in catalog).

Anastrepha

Anastrepha fraterculus

Anastrepha distincta

Anastrepha pallens

Map 5. Distribution of Anastrepha chiclayae, A. distincta, A. fraterculus, A. pallens, A. spatulata, and A. zuelaniae.

- Wasbauer 1972: 100 (hosts). - Steyskal 1977b: 11, 20 (in key). - Norrbom 1985: 239, 287, 296 (hosts, in key). - Norrbom and Kim 1988b: 15 (hosts). recognition. A small, entirely yellow and white species,

chiclayae is rather

difficult to distinguish externally from other Texas species with a “typical” Ana¬

strepha wing pattern. The costal and S bands to some extent are confluent in many specimens; specimens in which these bands are completely separated somewhat resemble those of edentata, a Florida species, from which it may be distinguished by the aculeus (fig. 151), the tip of which features numerous fine lateral serrations. In

chiclayae, these lateral serrations occupy most of the lateral margins of the tip, similar to those in limae (fig. 153), but in the former, the serrations do not extend quite as far as the terminus of the oviduct. distribution. Map 5. In addition to the Texas localities shown,

chiclayae has

been collected in several localities in Mexico and Panama and in at least one location in Peru. It probably occurs throughout Central America and northwestern South America wherever its hosts are present. hosts. The only known field infestation by

chiclayae is of Passiflora sp. in Mex¬

ico. Nine additional plant hosts were listed by Norrbom and Kim (1988b) as ques¬ tionable. discussion.

A. chiclayae is of no known economic importance.

94

Systematic Treatment of the Genera

Anastrepha dentata (Stone) (Figs. 24, 132, 148, 160; Map 4) Lucumaphila dentata Stone 1939b: 343 (female holotype, NMNH; Tequila, Jalisco, Mex.). — Aczel 1949: 234 (in catalog). - Foote 1965a: 673 (in catalog). Anastrepha dentata: Steyskal 1977b: 31 (in key). - Norrbom 1985: 123, 289 (taxon¬ omy, hosts, in key). recognition.

A. dentata and sagittata are similar to bicolor and pallens in

having an extremely slender, needlelike aculeus, the tip of which is not expanded but has extremely minute serrations (figs. 147, a; 148, a), unlike the aculeus of most Anastrepha, which are generally transversely flattened and wider in proportion to their length. These are the only four U.S. species of Anastrepha with this character. In all of them, the distiphallus is also absent in the male. The latter two species can be distinguished from the former two by their atypical wing pattern. A. dentata is distinctive in showing two or three dark unconnected spots of varying extent on the scutum immediately anterior to the scutoscutellar suture (fig. 160) (compare with scutal markings of A. pallens, fig. 157); it contrasts with those markings in sagittata, which are uniform and less distinct along the posterior margin of the scutum (fig. 161, a). Map 4. Essentially a Mexican species, dentata rarely occurs in the United States; it was last collected in two localities in Texas in 1940. distribution.

host.

Not

known.

discussion.

A. dentata is of no known economic importance.

Anastrepha distincta Greene (Fig. 146; Map 5) Anastrepha distincta Greene 1934: 149 (female holotype, NMNH; Chiclayo, Peru). - Stone 1942a: 106 (revision). - A. C. Baker et al. 1944: 132 (review, biology). Aczel 1949: 207 (in catalog). — Bush 1962: 93 (cytotaxonomy). - Foote 1965a: 673 (in catalog). - Wasbauer 1972: 100 (hosts). - Steyskal 1977b: 29, 30 (in key). - Norrbom 1985: 240, 292 (hosts, in key). - Norrbom and Kim 1988b: 16 (hosts). — Steck et al. 1990: 344 (in key to larvae). Populations of distincta from Peru have a slightly longer aculeus and might be confused with ludens, which also has a “typical” Anastrepha wing pattern, a yellow and white scutum, and a relatively long, slender aculeus. Females recognition.

from Mexico and Texas always can be distinguished by the relative length of the aculeus and usually by the markings on the subscutellum and mediotergite. The aculeus tip is similar to that shown in fig. 150, but the lateral serrations are some¬ times weaker and may be confined to the apical third of the tip. The V band is

Anastrepha separated from the S band and is often narrowly broken anteriorly. A. edentata is the only other U.S. species with a relatively long, slender oviscape, but it has a nonserrated aculeus tip, and the S and costal bands, which are usually confluent along vein R4+5 in distincta, are well separated. distribution.

Map 5. A. distincta is found throughout Mexico and Central and

South America south to Brazil. A. distincta is the only Anastrepha species known to prefer Inga species as hosts throughout its range; Norrbom and Kim (1988b) listed 18 species of that genus as hosts. Those authors also listed plant species in the genera Chrysophyllum, Eu¬ genia, and Mangifera as being field-infested, and they list eight additional plant hosts.

genera containing doubtful or laboratory hosts.

Anastrepha edentata Stone (Figs. 134, 149; Map 4) Anastrepha sp. F: A. C. Brown 1937: 19 (Fla.). Anastrepha edentata Stone 1942a: 48 (female holotype, NMNH; Key Largo, Fla.). Foote 1965a: 673 (in catalog). - Weems 1968b: 1 (review). - Steyskal 1977b: 14 (in key). - Norrbom 1985: 273, 288 (phylogeny, in key). RECOGNITION.

A. edentata is easily recognized among species occurring in Flori¬

da by the hyaline area that separates the costal and S bands. This area runs obliquely from the apex of the pterostigma posteriorly to the posterior wing margin (fig. 134, a) and is nearly parallel-sided until it crosses vein R4+5. In this respect, it resembles some Texas specimens of chiclayae, from which it is easily distinguished by the completely unserrated aculeus tip (fig. 149, a) and a much longer oviscape. distribution.

host.

Not

Map 4. A. edentata also is known from Puerto Rico.

known.

DISCUSSION.

Although formerly occurring in the Florida Keys throughout the

year, edentata has not been collected there in recent years.

Anastrepha fraterculus (Wiedemann) South American fruit fly (Figs. 139, 156; Map 5) Dacus fraterculus Wiedemann 1830: 524 (male syntype, NMW; Brazil). - Coquillett 1910: 503 (type data). - Hardy 1968: 145 (type data). Trypeta fraterculus: Loew 1873: 222 (review). Acrotoxa fraterculus: Loew 1873: 227, 329, 336 (taxonomy). Trypeta (Acrotoxa) fraterculus: Osten Sacken 1878: 189 (in catalog).

96

Systematic Treatment of the Genera Anastrepba fraterculus: Aldrich 1905: 602 (in catalog). — Hendel 1914c: 15 (review). - Rust 1918: 457 (review, biology). - C. T. Greene 1929: 496 (description larva, pupa). - Emmart 1933: 184 (description egg). - Bates 1934b: 1 (morphology larva). - C. T. Greene (part) 1934: 164 (review). - E. Cameron 1941: 13 (parasite). - Stone 1942a: 78 (revision). - A. C. Baker et al. 1944: 138 (review, biology). V. T. Phillips 1946: 106 (hosts). - Aczel 1949: 210 (in catalog). — Hennig 1952: 216 (in catalog immature stages). - Commonw. Inst. Entomol. 1958a: Map No. 88 (distribution). - Ebeling 1959: 246 (review). - Christenson and Foote 1960: 171 (biology). - Bush 1962: 93 (cytotaxonomy). - Wasbauer 1964: 6 (review). Foote 1965a: 673 (in catalog). - Hardy 1968: 145 (type data). - Wasbauer 1972: 100 (hosts). - Steyskal 1977b: 12, 20, 22 (in key). - Wharton and Marsh 1978: 148 (parasites). - Berg 1979: 21 (in key to larvae). - Nation 1981: 121 (sex pheromone glands). - Weems 1980: 1 (review). - Morgante et al. 1983: 234 (mating behavior). - Norrbom 1985: 1, 242, 298 (phylogeny, hosts, in key). Norrbom and Kim 1988b: 19 (hosts). - Steck et al. 1990: 344 (in key to larvae). Anastrepba fratercula [error]: Cole 1927: 445 (male genitalia). recognition.

Among all U.S. Anastrepba species, fraterculus, obliqua, and sus-

pensa present the most difficult identification problems in the genus; these three species are likely to be confused because of the similarity of their external features. Stone (1942a) lists the following critical differences between fraterculus and obliqua: A. fraterculus: (a) Aculeus usually longer than the distance on vein M from the junction of MP and M to vein r-m. (b) Subscutellum darkened laterally. A. obliqua: (a) Aculeus always shorter than the distance on vein M from the junction of vein MP and M to vein r-m. (b) Subscutellum not darkened laterally. The apical arm of the S band of fraterculus is narrow compared with that of suspensa (see recognition section for that species). There is frequently a distinct scutoscutellar black spot, but it is usually smaller than in suspensa. One of the most important distinguishing features is the nature of the aculeus tip, which has serrations only on its apical third (fig. 156, a) in contrast to that of obliqua (fig. 154, b). Map 5. A. fraterculus occurs from Texas to Argentina; in some areas it is extremely abundant. See Weems (1980) and below for further information about the status of various populations of this species. distribution.

A. fraterculus is the most economically important Anastrepba in Brazil and in some other South American countries because of its wide host range that includes many economically important fruits. Norrbom and Kim (1988b) listed hosts.

plants in the following genera as being field-infested throughout the range of frater¬ culus: Alcbornea, Anacardium, Annona, Averrboa, Campomanesia, Citrus, Coffea, Crataegus, Cydonia, Diospyros, Dovyalis, Eriobotrya, Eugenia, Eeijoa, Ficus, For-

Anastrepha tunella, Fragaria, Inga, Juglans, Malpighia, Malus, Mangifera, Manilkara, Masticbodendron, Myrcia, Persea, Pouteria, Prunus, Psidium, Punica, Pyrus, Rubus, Solanum, Spondias, Syzygium, Terminalia, Theobroma, Turpinia, Vitis, and Ximenia. Those authors also included questionable records and laboratory rearings. discussion.

A. fraterculus is widespread and variable morphologically and in its

pest status in different regions (Weems 1980). Isozyme and karyotype studies (Solferini and Morgante 1987, Steck 1991) suggest that what has been considered frater¬ culus consists of several closely similar “sibling species.” Closely related species in Brazil have been described (Zucchi 1979), but other species in this complex may not yield to the usual morphological taxonomic techniques. We expect that various biological studies will be required to reveal the true status of fraterculus in the wider sense.

Anastrepha interrupta Stone (Figs. 141, 152; Map 4) Anastrepha Sp. E: A. C. Brown 1937: 20 (Fla.). Anastrepha interrupta Stone 1942a: 62 (female holotype, NMNH; Jensen, Fla.). McClanahan 1951: 44 (host). - Shaw 1962: 409 (review). - Foote 1965a: 673 (in catalog). - Weems 1967a: 1 (review). - Marsh 1970: 31 (parasite). - Wasbauer 1972: 102 (host). - Steyskal 1977b: 16 (in key). - Wharton and Marsh 1978: 148 (parasite). - Norrbom 1985: 246, 293 (hosts, in key). - Norrbom and Kim 1988b: 30 (hosts). - Steck and Wharton 1989: 995 (description larva). - Steck et al. 1990: 343 (in key to larvae).

A. interrupta and spatulata are unique among U.S. Anastrepha in having a dark scutoscutellar spot in combination with a very short oviscape and a recognition.

distinctively shaped aculeus tip (fig. 152). This species closely resembles spatulata in wing pattern; in both species the costal and S bands are touching to broadly con¬ fluent, and the V band is complete but usually separated from the S band. A. interrupta is distinguished from spatulata principally by the nature of the hyaline spot distad of the pterostigma (fig. 141, b); in interrupta, it is small with indistinct borders and rarely touches vein R2+3, whereas in spatulata it is much more promi¬ nent and almost always terminates broadly on vein R4+5. distribution.

Map 4. A. interrupta was originally known only from Florida,

but it also occurs in the Bahamas. In past years, it has occurred commonly in some of the areas shown on the map. hosts.

A. interrupta was reared from the fruits of Schoepfia chrysophylloides and

5. schreberi (Norrbom and Kim 1988b). discussion.

A. interrupta is of no known economic importance. Steck and

Wharton (1989) described the third instar in detail.

98

Systematic Treatment of the Genera

Anastrepha limae Stone (Figs. 142, 153; Map 4) Anastrepha limae Stone 1942a: 67 (female holotype, NMNH; Panama City, Pan¬ ama). - Aczel 1949: 216 (in catalog). - Foote 1965a: 673 (in catalog). - Wasbauer 1972: 102 (host). - Steyskal 1977b: 18 (in key). - Norrbom 1985: 246, 294 (host, in key). - Norrbom and Kim 1988b: 31 (host). - Steck and Wharton 1989: 997 (description larva, puparium). - Steck et al. 1990: 342 (in key to larvae). Among the Texas species of Anastrepha without dark scutal or scutellar markings, limae is distinguished principally by the nature of the aculeus tip recognition.

(fig. 153), which displays fine lateral serrations from the apex to a point basad of the opening of the oviduct. In this respect it somewhat resembles chiclayae, the lateral serrations of which (fig. 151) do not extend as far basad. It differs from spatulata and interrupta by having an elongate ovipositor, the tip of which is not spatulate. distribution.

Map 4. A. limae was described from Panama, where it has been

reared from its host plant at several localities. It probably occurs throughout Mexico and Central America wherever its host occurs. host.

Passiflora quadrangular is is the only known host (Norrbom and Kim

1988b). discussion.

Only two specimens of limae have ever been found in the United

States, and it is rarely encountered elsewhere. Steck and Wharton (1989) described the third instar and puparium in detail.

Anastrepha ludens (Loew) Mexican fruit fly, orange worm, orange fruit worm, Morelos orange worm, gusano de la naranja (Figs. 1-6, 135, 150, 163, 165; Map 7) Trypeta ludens Loew 1873: 223 (male lectotype, here designated, NMW; Mexico). L. O. Floward 1888: 45 (review). - Riley 1889: 45 (biology). - Johnson 1898: 53 (review). - Isaac 1905: 1 (hosts). - Crawford 1910: 321 (early history, biology). Crawford 1913: 2 (hosts). Acrotoxa ludens: Loew 1873: 327, 329 (taxonomy). Trypeta (Acrotoxa) ludens: Osten Sacken 1878: 189 (in catalog). Anastrepha ludens: Aldrich 1905: 602 (in catalog). - Herrera 1908: 169 (review, Mexico). - Banks 1912: 33 (description larva). - Cole 1927: 444 (male genitalia). - Crawford 1927: 421 (review, hosts). - Mackie 1928: 295 (review, biology, hosts). - C. T. Greene 1929: 495 (description larva, pupa). - Fleury 1932: 316 (review, Tex.). - Emmart 1933: 184 (description egg). - Bates 1934b: 1 (morphol¬ ogy larva). - Darby and Kapp 1934: 1 (biology). - C. T. Green 1934: 151 (review).

Anastrepha - Emmart 1935: 119 (chromosomes). - Plummer and Stone 1935: 1 (hosts). Quayle 1938: 240 (review). - Plummer and McPhail 1941: 1 (hosts). - C. L. Metcalf 1942: 507 (Ill.). - Stone 1942a: 97 (revision). - A. C. Baker et al. 1944: 1 (review, biology). — V. T. Phillips 1946: 30, 106 (description, biology larva; hosts). - Aczel 1949: 216 (in catalog). - Hennig 1952: 216 (catalog immature stages). Hoidale 1953: 559 (review). - Peterson 1953: 331, 333, 335, 337 (figures of larvae). - Berry 1955: 414 (review, hosts). - Flitters et al. 1956: 1 (bioclimatic cabinet studies). - Messenger and Flitters 1957: 119 (bioclimatic cabinet studies). - Christenson 1958: 11 (review of research). - Commonw. Inst. Entomol. 1958b: Map No. 89 (distribution). - Flitters and Messenger 1958: 7 (bioclimatic cabinet studies). - Ebeling 1959: 244 (review). - Christenson and Foote 1960: 171 (biolo¬ gy). - Bush 1962: 91 (cytotaxonomy). - Christenson 1963: 441 (review, current investigations). - Foote and Blanc 1963: 12 (review, Calif.). - Weems 1963: 1 (review). - Wasbauer 1964: 7 (review). - Flitters and Messenger 1965: 1 (biology). -Foote 1965a: 673 (in catalog). - Mena 1965: 13 (hosts). - Rubio and McFadden 1966: 1015 (associates, host). - Bateman 1972: 495 (biology). - Wasbauer 1972: 102 (hosts). - Wright and Burgess 1975: 1248 (olfactory physiology). - Steyskal 1977b: 10, 12, 30 (in key). - Steyskal 1977c: 12 (McPhail trap). - Wharton and Marsh 1978: 148 (parasites). - Berg 1979: 20, 22 (in key to larvae). - Nation 1981: 121 (sex pheromone glands). - Weems 1981b: 4 (review). - Dickens et al. 1982: 9 (behavior, development). - Rhode and Sanchez 1982: 98 (trapping). Heppner 1984: 1 (description larva). - Norrbom 1985: 247, 286, 292 (hosts, in key). _ Norrbom and Kim 1988b: 31 (hosts). - Carroll and Wharton 1989: 201 (morphology immature stages). - E. J. Harris 1989: 76 (pest status). - Steck et al. 1990: 343 (in key to larvae). Anastrepha lathana Stone 1942a: 105 (female holotype, NMNH; Cuernavaca, Mex¬ ico). - Stone 1942b: 304 (distribution, Tex.). - Aczel 1949: 215 (in catalog). Foote 1965a: 673 (in catalog). - Wasbauer 1972: 102 (host). - Steyskal 1977b: 10, 30 (in key). - Norrbom 1985: 246,293 (host, in key). - Norrbom and Kim 1988b: 30 (host). - Hernandez 1990: 106 (synonymy). - Hernandez 1991: 233 (syn¬ onymy). A. ludens is characterized by a relatively long aculeus and oviscape, the former 3.4-4.7 mm long and the latter correspondingly long and tapering in its apical third. This external character alone will alert the identifier to the possi¬ bility of ludens in areas where ludens is already known to exist, especially in large numbers (e.g. the Rio Grande Valley of Texas). The apical third of the aculeus tip (fig. 150) is slightly expanded in the area of the lateral serrations, which are relatively few and not prominent. A. suspensa (fig. 155) and fraterculus (fig. 156) differ in having a much shorter aculeus and aculeus tip with more prominent lateral serrations and by recognition.

other characters as well. A. ludens also usually can be recognized by the color of the postnotum. A pair of lateral dark spots is usually present on the subscutellum; they typically extend ventrally onto the mediotergite (fig. 165, b). The V band is usually not connected to the S band and is faint anteriorly in most specimens.

100

Systematic Treatment of the Genera

distribution.

Map. 7. A. ludens occurs from Texas south to Costa Rica. It is

indigenous to Mexico, from where it migrates annually across the Rio Grande Valley into the United States. It has been the subject of a sterile male release control program in southern California and has been eradicated there. In much of its range, ludens is one of the most commonly encountered species in Citrus; of these crops, ludens normally prefers grapefruit over orange. hosts.

Norrbom and Kim (1988b) listed plant species in the following genera as hosts: Anacardium, Annona, Carica, Casimiroa, Citrus, Coffea, Cydonia, Diospyros, Inga, Malus, Mammea, Mangifera, Mastichodendron, Passiflora, Persea, Prunus, Psidium, Punica, Pyrus, Sargentia, Spondias, and Syzygium. Several other plant genera were listed by those authors as doubtful hosts. discussion.

A. ludens is a distinctly subtropical to temperate fly. For that reason

and because of its wide host range, including fruits of economic importance such as grapefruit and orange, ludens is economically the most significant Anastrepba spe¬ cies in the United States. The lectotype has the following labels: white square with “89,” “ludens det. Low,” and “Mexico, coll. Winthem.” The holotype of lathana is only an abnormally small specimen of ludens.

Anastrepha nigrifascia Stone (Fig. 145; Map 4) Anastrepba sp. W: A. C. Brown 1937: 21 (Fla.). Anastrepba nigrifascia Stone 1942a: 91 (female holotype, NMNH; Big Pine Key, Fla.). - Foote 1965a: 673 (in catalog). - Weems 1967b: 1 (review). - Wasbauer 1972: 107 (hosts). — Steyskal 1977b: 25 (in key). — Norrbom 1985: 147, 284, 290 (taxonomy, hosts, in key). — Norrbom and Kim 1988b: 40 (hosts). recognition.

A rather small, primarily yellow species, nigrifascia is distinguish¬

able from all other Anastrepba species occurring in Florida by the presence of a distinct lateral brown band at the posterior margin of the scutum somewhat similar to that shown in fig. 161 for sagittata. Differences from the latter species are given in the key. In nigrifascia, the aculeus is not more than 2.0 mm long, and the tip is not serrate. Map 4. A. nigrifascia is one of the very few Anastrepba not known to occur outside the United States and the Bahamas. In 1935-1936, it was distribution.

commonly encountered in the Florida Keys but has not been collected there since then. hosts.

According to Norrbom and Kim (1988b), nigrifascia has been reared from

Manilkara emarginata (recorded by Stone [1942a] as Mimusops) and from M. zapota (recorded by Stone [1942a] as Acbras). Mangifera indica is a questionable host.

101

Anastrepha

discussion. A. nigrifascia is primarily a seed feeder. Larvae are found in the flesh of the fruit only on their way to or from the outside surface.

Anastrepha obliqua (Macquart) West Indian fruit fly (Figs. 143, 154; Map 6) Tepbritis obliqua Macquart 1835: 464; 1843a: 225 (female holotype, MNHN; Cuba). - Steyskal 1975a: 357 (type data). Anastrepba acidusa [misidentification]: Loew 1862c: 57 (taxonomy). - Aldrich (part) 1905: 602 (in catalog). — Bates 1934b: 1 (morphology of larva). — C. T. Greene 1934: 162 (review). - Newell 1935: 22 (biology). - McAlister 1936: 440 (biology, hosts, Key West). - Newell 1936: 116 (biology). - A. C. Brown 1937: 20 (distribution). - Quayle 1938: 243 (review). - Aczel (part) 1949: 233 (in catalog). Acrotoxa acidusa: Loew 1873: 335 (taxonomy). — Johnson 1895b: 337 (Fla.). — Johnson 1913b: 83 (Fla.). Trypeta (Acrotoxa?) acidusa: Osten Sacken 1878: 194 (in catalog). Anastrepba fratercuius [misidentification]: Newell 1933: 17 (discovery, Fla.). Anastrepba frater cuius var. mombinpraeoptans Sein 1933: 187 (female holotype, NMNH; Rio Piedras, Puerto Rico). - Steyskal 1975: 357 (synonymy). Anastrepba mombinpraeoptans: Waterston 1941: 38 (introduction, Bermuda). Stone 1942a: 45, 68 (revision). - A. C. Baker et al. 1944: 134 (review, biology). Aczel 1949: 218 (in catalog). — Hennig 1952: 216 (in catalog immature stages). — Commonw. Inst. Entomol. 1958c: Map No. 90 (distribution). - Christenson and

Map 6. Distribution of Anastrepha obliqua and A. suspensa.

Systematic Treatment of the Genera Foote 1960: 171, 185 (biology). — Bush 1962: 93 (cytotaxonomy). — Wasbauer 1964: 7 (review). — Foote 1965a: 673 (in catalog). — Angeles 1966: 299 (hosts). — Weems 1970: 1 (review). — Wasbauer 1972: 104 (hosts). — Steyskal 1977b: 19 (synonymy). — Berg 1979: 21, pi. 4 (in key to larvae). Anastrepba obliqua: Stone 1942a: 75 (revision). — Steyskal 1975a: 357 (synonymy). — Steyskal 1977b: 19 (in key). — Wharton and Marsh 1978: 148 (parasites). — Nation 1981: 121 (sex pheromone glands). — Norrbom 1985: 1, 251, 297 (hosts, in key). — Norrbom and Kim 1988b: 40 (hosts). — Steck et al. 1990: 344 (in key to larvae). Anastrepba fratercuius var. mombinpraeoptera [error]: V. T. Phillips 1946: 106 (hosts). Anastrepba mombimpreoptans [error]: Mena 1965: 13 (hosts).

Externally, obliqua quite closely resembles fraterculus and suspensa, thereby presenting problems in their separation. Stone (1942a) compares a recognition.

number of characters that appear to be critical in separating obliqua from fraterculus (see the listing in the discussion of the latter species). The aculeus is subtly different from those of fraterculus and suspensa, having lateral serrations on more than twothirds of the tip (fig. 154) in contrast to those of the other two species, where they are limited to the apical two-fifths to three-fifths of the tip (figs. 155, 156). In obliqua, the tip also is relatively wider at the base of the serrations compared with the width at the genital opening. The white medial vitta on the scutum is wider in obliqua than in suspensa and fraterculus, and no scutoscutellar black spot or lateral dark marks on the subscutellum are present, although the mediotergite usually has a lateral dark stripe.

Map 6. Of greatest importance to U.S. fruit growers is the fact that obliqua occurs throughout the Greater and Lesser Antilles. It is one of the most distribution.

widespread Anastrepba species, having been recorded from Trinidad, Mexico to Panama, Venezuela, Ecuador, and in the vicinity of Rio de Janeiro, Brazil. It has been trapped in the Florida keys, in the Rio Grande Valley of Texas, and in San Diego, Orange, and Los Angeles counties in California.

hosts.

For A. obliqua, Norrbom and Kim (1988b) listed more than 60 plant

species belonging to 24 plant families. Those plant genera recorded as being fieldinfested are: Alcbornea, Anacardium, Annona, Averrboa, Brosimum, Citrus, Coffea, Crataegus, Diospyros, Dovyalis, Eriobotrya, Eugenia, Geoffraea, Godmania, )ambosa, Malpigbia, Malus, Mangifera, Manilkara, Passiflora, Pouteria, Prunus, Psidium, Pyrus, Spondias, and Syzygium. A. obliqua is one of the most important fruit fly pests of mango, but it also favors fruits of other species of Anacardiaceae such as Spondias and will attack many other fruits such as guava and rose apple. It occurs rarely in Citrus. Its economic significance is increasing because markets for mangoes in the United States and Europe are expanding.

103

Anastrepha

Anastrepha ocresia (Walker) (Fig. 129; Map 4) Trypeta ocresia Walker 1849: 1016 (female syntype, BMNH; Jamaica). — Osten Sacken 1858: 79 (in catalog). — Loew 1862c: 60 (taxonomy). — Loew 1873: 231 (taxonomy). Acrotoxa ocresia: Loew 1873: 231, 337 (taxonomy). Trypeta (Acrotoxa) ocresia: Osten Sacken 1878: 189, 195 (in catalog). Anastrepha ocresia: Aldrich 1905: 602 (in catalog). — Hendel 1914c: 15 (review). — C. T. Greene 1934: 158 (review). — A. C. Brown 1937: 21 (distribution, Fla.). — Stone 1942a: 24 (revision). - A. C. Baker et al. 1944: 125 (review, biology). Aczel 1949: 221 (in catalog). - Foote 1964d: 322 (type data). - Foote 1965a: 673 (in catalog). — Weems 1968a: 1 (review). — Wasbauer 1972: 107 (hosts). — Steyskal 1977b: 8 (in key). - Norrbom 1985: 257, 282 (hosts, in key). - Norrbom and Kim 1988b: 51 (hosts). A. ocresia and serpentina belong to a small group of Anastrepha species having dark wing markings that contrast with the light hyaline areas of the wing. Unlike most species in the genus, the lighter yellowish area in cell c extends well into cell r, immediately posterior to the pterostigma. Of all Anastrepha species with these characters, ocresia is recognizable by the reduced apical arm of the V band (fig. 129, c), although there is some variation in the shapes of both arms of that band from those shown in fig. 129. In contrast to those of serpentina, the dark markings of the scutum are restricted to the scutoscutellar area. The abdominal markings are also recognition.

distinctive. Map 4. A. ocresia supposedly occurs in the Florida Keys but to our knowledge has not been collected there since 1936. It also occurs in Cuba, distribution.

Jamaica, Puerto Rico, and Hispaniola. hosts.

Psidium guajava and Manilkara zapota are the only hosts known for

ocresia (Norrbom and Kim 1988b). discussion.

Little is known about this species, which is rarely collected within

its limited range.

Anastrepha pallens Coquillett Bumelia fruit fly (Figs. 127, 157; Map 5) Anastrepha pallens Coquillett 1904: 35 (male holotype, NMNH; Brownsville, Tex.). - Hendel 1914a: 69 (in key). - C. T. Greene 1934: 154 (review). - V. T. Phillips 1946: 31,106 (description, biology larva; hosts). - Steyskal 1977b: 3 (in key). Norrbom 1985: 112, 257, 278 (taxonomy, hosts, in key). Anastrepha (Pseudodacus) pallens: McPhail and Berry 1936: 405 (biology).

104

Systematic Treatment of the Genera

Pseudodacus pallens: Stone 1939a: 283 (revision). - Stone 1942a: 10 taxonomy). A. C. Baker et al. 1944: 115 (review, biology). - Aczel 1949: 238 (in catalog). Hennig 1952: 216 (catalog immature stages). — Peterson 1953: 333 (figures of larva). - Foote 1965a: 674 (in catalog). - Wasbauer 1972: 125 (hosts). - Norrbom and Kim 1988b: 51 (hosts). recognition.

A. pallens somewhat resembles A. bicolor in having an atypical S

band configuration in the wing pattern; additionally, in both species, the distal arm of the V band is completely absent (fig. 127, b) and the aculeus is very slender and needlelike (see recognition characters for A. dentata). In contrast to bicolor, pallens has a hyaline spot apicad of the pterostigma that at least partially separates the costal band and apical part of the S band. The middle part of the S band is present, although it is separated from the basal and apical parts by hyaline areas in cell r2+3 and along the basal part of vein CuAj. The bands are yellowish or light brown, are lighter in color than in most Anastrepha species, and contrast less strongly with the hyaline areas of the wing. The pattern is sometimes difficult to see clearly. In contrast to bicolor, most of the scutal pattern in pallens is yellowish or orange with dark brown markings (fig. 157, a) only near the posterior margin that are typical of this species. In the male, the distiphallus is absent (see recognition characters for den¬ tata). distribution.

Map 5. A. pallens has been somewhat more commonly encoun¬

tered than bicolor in Texas. It has been recorded south from Mexico to Honduras (Norrbom 1985). host.

Norrbom and Kim (1988b) indicated that Bumelia celastrina is the only

recorded host; previous records of B. spiniflora probably represent misidentifications of celastrina. discussion.

A. bicolor and pallens previously had been assigned to the genus

Pseudodacus Hendel because of the lack of a typical Anastrepha wing pattern, their needlelike aculei, and other characters.

Anastrepha sagittata (Stone) (Figs. 133, 161; Map 4) Lucumapbila sagittata Stone 1939b: 347 (female holotype, NMNH; Cuernavaca, Mexico). — Aczel 1949: 235 (in catalog). - Foote 1965a: 673 (in catalog). Wasbauer 1972: 118 (hosts). Anastrepha sagittata: Steyskal 1977a: 77 (type data). — Steyskal 1977b: 32 (in key). Norrbom 1985: 130, 289 (taxonomy, hosts, in key). - Norrbom and Kim 1988b: 55 (host). One of four Anastrepha species occurring in the United States having an extremely narrow, needlelike aculeus and lacking a distiphallus in the male recognition.

105

Anastrepha

(see additional comments in the recognition section of A. dentata), sagittata is distin¬ guished by the “typical” Anastrepha wing pattern in combination with the presence of a dark, more or less uniform transverse marking at the posterior border of the scutum adjacent to the scutoscutellar suture (fig. 161, a); the latter character sepa¬ rates it from dentata (compare fig. 160). Map 4. Originally described from Mexico, sagittata is rare in the United States, having been collected only at Mission, Texas, in 1938. It is part of the hamata (Loew) species complex, which includes several poorly differentiated forms distribution.

occurring south to Bolivia and Brazil (Norrbom 1985). The larvae of sagittata feed on the seeds of Pouteria campechiana, the only known host (Norrbom and Kim 1988b). This plant has previously been reported in the fruit fly literature as belonging to the genus Lucuma. host.

discussion.

This species is of no known economic importance.

Anastrepha serpentina (Wiedemann) Black fruit fly (Figs. 130, 164; Map 7) Dacus serpentinus Wiedemann 1830: 521 (female syntypes, NMW; Brazil). - Loew 1873: 227 (taxonomy). — Stone 1942b: 298 (type data). Acrotoxa serpentinus: Loew 1873: 227, 329 (taxonomy). Anastrepha serpentina: Coquillett 1910: 506 (type data). - Hendel 1914b: 84 (type data). - V. T. Phillips 1923: 127 (type data). - C. T. Greene 1929: 497 (description larva, pupa). - Emmart 1933: 184 (description egg). - Bates 1934b: 1 (morphol¬ ogy larva). - C. T. Greene 1934: 142 (revision). - Stone 1942a: 27 (revision). A. C. Baker et al. 1944: 115, 125 (biology). - V. T. Phillips 1946: 33, 107 (description, biology larva; hosts). - Shaw 1947: 34 (hosts, distribution). - Aczel 1949: 227 (in catalog). - Hennig 1952: 216 (in catalog immature stages). - Bush 1962: 95 (cytotaxonomy). - Wasbauer 1964: 7 (review). - Foote 1965a: 673 (in catalog). - Spishakoff 1966: 1010 (rearing). - Weems 1969b: 1 (review). Bateman 1972: 498 (ecology). - Wasbauer 1972: 107 (hosts). - Steyskal 1977b: 8 (in key). - Wharton and Marsh 1978: 148 (parasites). - Berg 1979: 23, pis. 2, 5 (in key to larvae). - Norrbom 1985: 259, 282 (hosts, in key). -Sarma et al. 1987: 271 (biochemistry). - Norrbom and Kim 1988b: 55 (hosts). - Steck et al. 1990: 344 (in key to larvae). As in ocresia and a few non-U.S. Anastrepha species, the very dark wing markings of serpentina contrast strongly with the light hyaline areas of the wing. The S band is quite slender and is not connected to the proximal area of the V band, the apical arm of which is absent in all specimens. A. serpentina and ocresia are the only species of Anastrepha occurring in the United States that have a distinct pale yellow to hyaline area in cell r1 immediately posterior to the pterostigma, but RECOGNITION.

• Anastrepha ludens

• Anastrepha serpentina

• Anastrepha striata

Map 7. Distribution of Anastrepha ludens, A. serpentina, and A. striata.

106

107

Anastrepha

the former may be distinguished from the latter by the complete absence of the distal arm of the V band and the difference in abdominal markings (see key). The scutum of this species (not figured) is characterized by contrasting light and dark markings; the subscutellum and mediotergite (fig. 164, a) are very dark, with a lighter brownish or yellowish spot or stripe dorsally. Map 7. A. serpentina seldom has been found in Texas since about 1959. It was trapped in southern California in 1989. In addition to the records shown here, it occurs abundantly in Mexico and most countries of Central and South distribution.

America south to Brazil. Norrbom and Kim (1988b) listed about 40 plant species in 13 plant families as hosts. Those authors listed field infestations of species in the following genera: Alchornea, Annona, Bumelia, Byrsonima, Cbrysophyllum, Citrofortunella hosts.

x mitis, Citrus, Cydonia, Diospyros, Dovyalis, Ficus, Lacmellea, Malus, Mammea, Mangifera, Manilkara, Masticbodendron, Micropbolis, Mimusops, Persea, Pouteria, Prunus, Psidium, Pyrus, and Spondias. Sapotaceae appears to be the favored host family. A. serpentina is commonly encountered in collections, probably because of its relatively wide host range. It is not known as an economically impor¬ tant species except in areas where one or more of its hosts are grown commercially. discussion.

Anastrepha spatulata Stone (Fig. 140; Map 5) Anastrepha spatulata Stone 1942a: 61 (female holotype, NMNH; Hacienda Sta. Engracia, Tamaulipas, Mexico). - Aczel 1949: 228 (in catalog). - Bush 1962: 94 (cytotaxonomy). - Shaw 1962: 413 (review). - Foote 1965a: 673 (in catalog). Steyskal 1977b: 16 (in key). - Norrbom 1985: 293 (in key). recognition.

A. spatulata

and

interrupta

are the only

Anastrepha

species that

have a very short aculeus in combination with a dark scutoscutellar spot (similar to that in fig.

162)

has a “typical”

and a distinctively shaped aculeus tip (as in fig.

Anastrepha

broadly confluent, and the band by a hyaline area. (fig.

140,

As

wing pattern; the costal and

V

bands are touching to

band is complete but is usually separated from the

S

stated in the key, the hyaline spot distad of the pterostigma

b) is much larger and more distinct than in

across cell r2+3 and terminating rather broadly on vein

distribution.

S

152). A. spatulata

interrupta,

usually passing

R4+5.

Map 5. In the past, spatulata has been trapped extensively in the

Rio Grande Valley in Texas; otherwise it is known to occur only in parts of Mexico and Panama. host.

Not

known.

108

Systematic Treatment of the Genera

discussion.

Almost nothing is known about the habits and biology of spatulata.

Anastrepha striata Schiner (Figs. 131, 159; Map 7) Anastrepha striata Schiner 1868: 264 (female holotype, NMW; Venezuela). - C. T. Greene 1929: 497 (description larva, pupa). - Emmart 1933: 184 (description egg). — Bates 1934b: 1 (morphology larva). - C. T. Greene 1934: 145 (review). — Stone 1942a: 29 (revision). - A. C. Baker et al. 1944: 130 (review, biology). - V. T. Phillips 1946: 107 (hosts). - Aczel 1949: 229 (in catalog). - Hennig 1952: 216 (in catalog immature stages). - Christenson and Foote 1960: 171 (biology). — Bush 1962: 94 (cytotaxonomy). — Harper 1964: 86 (Calif.). — Wasbauer 1964: 7 (re¬ view). -Foote 1965a: 673 (in catalog). — Hardy 1968: 136 (type data). - Wasbauer 1972: 108 (hosts). — Steyskal 1977b: 5 (in key). — Wharton and Marsh 1978: 148 (parasites). — Berg 1979: 23, pi. 6 (in key, larvae). — Zucchi 1979b: 265 (type data, taxonomy). — Nation 1981: 121 (sex pheromone glands). — Weems 1982: 1 (re¬ view). - Norrbom 1985: 263, 282 (hosts, in key). — Norrbom and Kim 1988b: 62 (hosts). — Steck et al. 1990: 343 (in key to larvae). A small to medium-sized species with a “normal” Anastrepha wing pattern, striata is one of the few species occurring in the United States that has recognition.

distinct dark scutal markings (fig. 159) in addition to darkening along the scutoscutellar suture. On the sublateral dark scutal areas (Fig. 159, a), a pair of dense patches of short, brownish black setae is present, as well as some hoary pile visible only when viewed from in front, but the lateral half of the scutal brown stripe is denuded. A. striata is the only U.S. species having such scutal characters. The aculeus tip is distinctively broad and wedge-shaped with a very blunt apex and extremely fine lateral serrations. The size of the hyaline mark at the apex of vein Rl varies consider¬ ably in this apecies. distribution.

Map 7. A. striata is included in this handbook by virtue of only

one U.S. collection in McAllen, Texas (it has not been seen there since 1959), one collection in San Ysidro, California, in 1963, and one in Los Angeles in 1989. These occurrences probably originated by infested fruit having been brought into the United States. It occurs in Mexico, Central America, and throughout much of South America. hosts.

The preferred host fruit of striata is guava. Norrbom and Kim (1988b)

listed about 20 additional host plants belonging to eight plant families; striata has been found in field infestations in the following genera: Annona, Bumelia, Chrysophyllum, Citrus, Diospyros, Mangifera, Manihot, Myrcia, Pouteria, Prunus, Psidium, Solanum, Spondias, Syzygium, and Terminalia. A. striata is one of the most commonly encountered species in the genus south of the United States. discussion.

109

Anastrepha Anastrepha suspensa (Loew)

Caribbean fruit fly, Carib fly (Figs. 137, 155, 162; Map 6) Trypeta suspensa Loew 1862c: 69 (male syntype, MCZ; Cuba). - Loew 1873: 222

(review). Acrotoxa suspensa: Loew 1873: 227, 239 (taxonomy). Trypeta (Acrotoxa) suspensa: Osten Sacken 1878: 189 (in catalog). Anastrepha suspensa: Aldrich 1905: 602 (in catalog). - C. T. Greene 1934: 147

(review). — Newell 1935: 24 (biology, Fla.). — McAlister 1936: 440 (biology, hosts, Key West). — Newell 1936: 116 (biology, Fla.). — Stone 1942a: 74 (revision). — V. T. Phillips 1946: 107 (hosts). - Aczel 1949: 230 (in catalog). - Christenson and Foote 1960: 171 (biology). — Foote 1965a: 673 (in catalog). — Weems 1965b: 1 (review). - Weems 1966: 401 (review, Fla.). - Baranowski 1969: 10 (review, para¬ site). — Baranowski and Swanson 1970: 161 (parasite). — Eden 1970: 150 (review, Fla.). - Kamasaki et al. 1970: 639 (laboratory culture). - Marsh 1970: 31 (para¬ site). - Lopez et al. 1971: 1541 (lure). - Prescott and Baranowski 1971: 297 (biology). — Baranowski and Swanson 1972: 249 (parasite). — Boush et al. 1972: 30 (bacterial associates). — Chambers 1972: 279 (lures). — Nation 1972: 1364 (behavior). - Swanson and Baranowski 1972: 271 (hosts). - von Windeguth et al. 1972: 127 (hosts, Fla.). - Wasbauer 1972: 109 (hosts). - Baranowski 1974: 17 (parasite). - Burditt et al. 1974: 387 (hosts). - Nation 1974: 731 (morphology). Taschenbert et al. 1974: 731 (biology). - Agee and Park 1975: 171 (eyes, vision). Baranowski 1975: 279 (biology, rearing). - Chambers 1975: 19 (mass rearing). Chow and Nation 1975: 515 (biology). - Conley 1975: 12 (biology). - Nation 1975: 27 (pheromone). - Perdomo et al. 1975: 291 (traps). - Sharp et al. 1975: 241 (biology). - Ashley et al. 1976: 391 (biology of parasite). - Chambers 1976: 25 (review). - Greany et al. 1976: 207 (parasite). - Lawrence et al. 1976: 33 (biology of parasite). - Leppla and Turner 1976: 61 (behavior). - Perdomo et al. 1976: 1208 (traps). - Sharp 1976: 255 (behavior, flight). - von Windeguth et al. 1976: 248 (host). - Webb et al. 1976a: 761 (biology). - Webb et al. 1976b: 415 (sound production). - Wolfenbarger et al. 1976: 118 (dispersal). - Agee et al. 1977: 359 (eye structure). - Greany et al. 1977a: 63 (trapping). - Greany et al. 1977b: 153 (parasite). - Greany et al. 1977c: 189 (parasite). - Greany et al. 1977d: 319 (biology of parasite). - Mazomenos et al. 1977: 139 (mating be¬ havior). - Prokopy et al. 1977: 463 (behavior). - Sharp and Webb 1977: 525 (behavior). - Steyskal 1977a: 75 (taxonomy). - Steyskal 1977b: 12, 20 (in key). Steyskal 1977c: 14 (trap). - Dodson 1978: 231 (morphology). - Greany et al. 1978: 20 (trapping). - Lawrence et al. 1978a: 93 (biology of parasite). - Lawrence et al. 1978b: 253 (biology of parasite). - Wharton and Marsh 1978: 148 (para¬ sites). - Ashley and Chambers 1979: 363 (biology of parasite). - Berg 1979: 22 (in key to larvae). - Burditt and McGovern 1979: 1 (attractants). - Greany and Szentesi 1979: 239 (behavior). - Lawrence 1979: 214 (description egg, larva). Sharp 1979: 70 (wing dimensions). - Szentesi et al. 1979: 227 (behavior). - Agee and Chambers 1980: 7 (mass rearing). - Morgante et al. 1980: 622 (systematics).

110

Systematic Treatment of the Genera — Sharp et al. 1980: 239 (morphology). — Bennett and Baranowski 1981: 197 (parasite). - Kitto et al. 1981: 1761 (immunotaxonomy). — Lawrence 1981: 249 (biology of parasite). — Nation 1981: 121 (sex pheromone glands). — Webb et al. 1981: 476 (laboratory studies, biology). — Burditt 1982: 367 (traps). — Burditt and Baranowski 1982: 85 (brief review). — Dodson 1982: 189 (mating, territoriality).— Goodin 1982: 14 (biology). - Greany et al. 1982: 374 (traps). — Sharp and Little 1982: 146 (flight muscle). — Witherell 1982: 580 (traps). — Burk 1983: 678 (mating). — Burk and Webb 1983: 678 (mating). — Sharp and Chambers 1983: 923 (aggregation). - Sharp et al. 1983: 302 (flight studies). - Webb et al. 1983: 996 (mating behavior). — Heppner 1984: 1 (description larva). - Sharp and Chambers 1984: 768 (physiology). - Sivinski 1984: 126 (mating). - Norrbom 1985: 264, 298 (hosts, in key). - Elzinga and Broce 1986: 208 (labellar morphology). Norrbom and Kim 1988b: 65 (hosts). - E. J. Harris 1989: 77 (pest status). - Steck et al. 1990: 344 (in key to larvae). Anastrepba unipuncta Sein 1933: 190 (female holotype, NMNH; Rio Piedras, Puer¬ to Rico). - Lima 1934: 506 (synonymy). Anastrepba longimacula C. T. Greene 1934: 146 (holotype female, NMNH; Hope, Jamaica). — Stone 1942a: 74 (synonymy). recognition. A. suspensa possesses external characters that closely resemble those of fraterculus and obliqua; therefore, the separation of these three species is sometimes difficult. One of the most obvious distinguishing marks in suspensa is the presence (except in some specimens from Jamaica) of a dark spot at the junction of the scutum and scutellum (fig. 162, a). This spot is sometimes present in fraterculus but is usually smaller, and it is absent in obliqua. The apical part of the S band in suspensa is relatively wide compared with that in the other two species, and its inner margin is less concave. It covers the apex of vein M or ends immediately anterior to it (fig. 138, c), whereas in the other two species it normally ends well anterior to the apex of vein M, or its inner margin is strongly concave (figs. 139, b; 143). As in the identification of other species of Anastrepba, the shape of the aculeus tip is impor¬ tant. In suspensa, as in fraterculus, the serrations occupy no more than three-fifths of the tip (fig. 155, a), whereas those in obliqua occupy at least two-thirds (fig. 154, b); this character is variable and should be used with care. distribution. Map 6. A. suspensa is limited to the Greater Antilles and the Bahamas and has occurred in Florida since about 1930. hosts. A. suspensa has been recorded from about 100 different host species occurring in many plant families. Norrbom and Kim (1988b) listed species in the following genera as having been infested in the field: Annona, Atalantia, Averrboa, Biscbofia, Bligbia, Britoa, Canella, Capsicum, Carica, Carissa, Casimiroa, Cbrysobalanus, Cbrysophyllum, Citrofortunella x mitis. Citrus, Clausena, Coccoloba, Diospyros, Dovyalis, Drypetes, Eriobotrya, Eugenia, Ficus, Flacourtia, Fortunella, Garcinia, Malpigbia, Malus, Mangifera, Manilkara, Mimusops, Momor-

Anastrepha dica, Muntingia, Murraya, Myrcianthes, Myrciaria, Phoenix, Pimenta, Pouteria, Prunus, Pseudanamomis, Psidium, Punica, Pyrus, Rheedia, Rubus, Severinia, Spondias, Synsepalum, Syzygium, Terminalia, Trevesia, and Triphasia. To date, it has not become a serious pest of commercial citrus in Florida. It is known to infest a larger number of plant species than any other Anastrepha that occurs in the United States, possibly because of the extensive field research that has been conducted in Florida. Although specimens have been trapped occasionally in Florida since the 1930s, suspensa was not permanently established until 1965, when what was apparently a new “strain,” presumably introduced from the Antilles, developed into an infestation in the southern third of the state. For excellent reviews of this situation, see Weems (1965, 1966). A great deal of research has been and is being undertaken by personnel of the USDA-ARS Insect Attractants, Behavior, and Basic Biology Re¬ search Laboratory in Gainesville, Fla., on all aspects of suspensa in relation to its environment. Much of the literature resulting from this research is cited above. discussion.

Anastrepha zuelaniae Stone (Fig. 144; Map 5) Anastrepha zuelaniae Stone 1942a: 82 (female holotype, NMNH; Barro Colorado I., Canal Zone, Panama). - Aczel 1949: 233 (in catalog). - Bush 1962: 91 (cytotaxonomy). — Foote 1965a: 673 (in catalog). — Wasbauer 1972: 111 (hosts).— Steyskal 1977b: 22 (in key). - Norrbom 1985: 271, 296, 297 (hosts, in key). Norrbom and Kim 1988b: 76 (hosts). A. zuelaniae is a rather small, yellowish brown species with a short aculeus and confluent costal and S bands. The V band is usually distinctly separated from the S band and is faint anteriorly. The most reliable character for its recognition is the shape of the aculeus tip, which has rather coarse lateral teeth along its posterior two-thirds and a rather distinct constriction immediately basad of the basal serration or tooth. It differs from the aculeus tip of obliqua (fig. 154) in that the lateral serrations extend somewhat farther basad, and the constriction is rather more pronounced in zuelaniae. The absence of dark lateral markings on the mediotergite RECOGNITION.

distinguish it from fraterculus, obliqua, and suspensa. Map 5. A. zuelaniae has been collected in Mexico and Panama and probably occurs throughout Mexico and Central America concurrently with its distribution.

host plant. host.

Zuelania guidonia is the only host known to be infested in the field (Norr¬

bom and Kim 1988b). discussion.

A. zuelaniae is not commonly encountered in Texas, it has not been

seen there since 1942.

112

Systematic Treatment of the Genera

Genus Bactrocera Macquart Bactrocera Macquart 1835: 452 (type species, Bactrocera longicornis Macquart 1835: 452, by monotypy). — Drew 1989a: 10 (subgeneric classification). — Drew 1989b: 1 (revision, Australasian and Oceanian Region). - Evenhuis 1989b: 2 (review of Hardy and Foote [1989]). — Hardy and Foote 1989: 503 (catalog, Australasian Region). Strumeta Walker 1856: 33 (type species, Strumeta conformis Walker 1856: 33, by monotypy). Dacus Fabricius: Hendel 1914b: 74 (in key). — Foote and Blanc 1963: 14 (review, Calif.). — Hardy 1973: 1 (taxonomy, Thailand). — Hardy 1974: 1 (taxonomy, Philippines). — Foote and Freidberg 1980: 30 (type data). — Munro 1984: 1 (revi¬ sion, Afrotropical Region [as family Dacidae]). recognition.

Species of the distinctive genus Bactrocera are quite different from

any of the native New World fruit flies. The genus may be identified with confidence by the size of cell bm, which in all species is at least two times as wide as cell cup, and by the complete lack of ocellar, postocellar, dorsocentral, presutural, and katepisternal bristles (fig. 14). The bristle of the postpronotal lobe (fig. 14, b) is usually absent. The radial veins and cells of the wing tend to be grouped together in the anterior half of the disk, so that vein R4+5 terminates well short of the wing apex (figs. 166—171). Cell cup is drawn out into an extremely long and narrow elongation (fig. 13, c). These characters also apply to the genus Dacus, which differs from Bactrocera in that the abdominal terga (see fig. 7) are completely fused. Based on an extensive study of Dacus species in the Oriental and Australasian Regions, especially those of New Guinea, Drew (1989a, 1989b) ele¬ discussion.

vated Bactrocera, formerly representing a subgenus of Dacus, to full generic rank. Bactrocera now includes almost all of the Oriental and Australasian Dacinae with free abdominal terga that were formerly placed in the genus Dacus. The latter, primarily Afrotropical, and Monacrosticbus Bezzi, a monotypic genus from the Phil¬ ippines, are the only other genera in the tribe Dacini. Because of the wide variety of hosts and extensive distribution of several of its species, Bactrocera is the most economically important fruit fly genus in the Oriental Region, and it competes with the medfly in importance where it occurs in the rest of the world. It now comprises about 30 subgenera and includes more than 800 species worldwide. See Evenhuis (1989) for additional names in synonymy, and our discus¬ sion of the genus Dacus for additional information concerning both genera. Bactrocera is included in this handbook only because of the past accidental intro¬ duction, or the possibility of introduction, of several of its species, as follows: B. correcta: Field-collected three times in Orange County, California, in 1986. Eradicated. B. cucurbitae: Accidentally introduced to California 1984-1986. Not established.

113

Bactrocera B. dorsalis: Repeatedly field-collected in California since 1966. Eradi¬ cated. B. latifrons: Larvae intercepted by USDA-APHIS in peppers from Hawaii. B. tryoni: A single female field-collected in California in 1985. B. zonata: Field-collected in California in 1984, 1987, and 1990. Erad¬ icated. The key and discussions presented herein provide a means of separating only these six species from each other. We have not attempted to accommodate any other species of this large genus that may be discovered in North America in the future.

Key to Species of Bactrocera Introduced or Suspected of Being Introduced into North America 1. Vein dm-cu with a prominent dark mark covering vein dm-cu (fig. 166, b) in addition to costal and cubital markings; costal band expanded in apical third to half of cell r4+5 into a large, prominent dark mark (fig. 166, a) .cucurbitae (Coquillett)

Figures 166-171 Right wings, Bactrocera spp. 166, cucurbitae (Coq.); 167, correcta (Bigot); 168, zonata (Saund.); *169, dorsalis (Hend.); 170, latifrons (Hend.); 171, tryoni (Frogg.).

Systematic Treatment of the Genera Vein dm-cu without a dark mark (fig. 167, d), or if a dark mark is present, very light-colored, narrow, and inconspicuous; costal band otherwise (fig. 167, c) .2 2. Costal band yellow in costal cells and pterostigma (fig. 167, a), very light brown throughout its length, nearly or entirely evanescent in cell r2+3 distad of pterostigma (figs. 167, b; 168, a) .3 Costal band of even darkness and thickness through pterostigma and cells rj and r2 + 3 (fig. 169, a), although it may be expanded at apex of vein (fl8- 170> a) .4 3. Scutum predominantly black; a complete transverse band present across lower face; wing pattern as in fig. 167 .correcta (Bezzi) Scutum rufous; face with 2 black spots; wing pattern as in fig. 168 . .zonata (Saunders) R4+5

4. Abdominal tergum yellow to orange with transverse black bands on tergites 2 and 3 and a narrow median black stripe extending from tergite 3 to end of tergite 5; wing pattern as in fig. 169 .dorsalis (Hendel) Abdominal tergum rufous and dark markings otherwise .5 5. Tergite 5 with a pair of shining oval yellow-brown spots; femora each with a preapical ventral brown mark; frons wide, 0.85-0.9 times its height (measured from antennal base to vertex); wing pattern as in fig. 170 .latifrons (Hendel) Tergite 5 entirely dark; femora without preapical ventral brown mark; frons narrower, 0.55 to 0.6 times its height; wing pattern as in fig. 171. .tryoni (Froggatt)

Bactrocera correcta (Bezzi) Guava fruit fly (Fig. 167; Map 8) Bactrocera zonata: Bezzi 1913: 94 (misidentification, not zonata Saunders 1841). Chaetodacus correctus Bezzi 1916: 107 (new name for Bactrocera zonata Bezzi 1913) (syntypes, depository not known; Bihar and Madras, India). Dacus correctus: Hardy 1973: 38 (description). - Weems 1987: 1 (review). recognition.

A species of moderate size with a predominantly black scutum

and a pair of rather broad, yellow, lateral postsutural yellow stripes. The scutellum is broadly yellow. The abdomen is primarily yellow with a dark transverse band across the base of tergite 3, from which a narrow dark central streak extends well onto tergite 5; in this respect it resembles the abdominal pattern of dorsalis. In wing pattern, correcta is similar to zonata in that the costal band is yellowish and evanes¬ cent or completely interrupted in cell r3 beyond the apex of vein R2+3 (fig- 167, b), but it is easily distinguished from zonata, and from most other species of Bactrocera including dorsalis, in having a complete transverse black band across the lower third of the face rather than a pair of black spots. The posterior fifth of the hind tibia of the male is abruptly narrowed to its apex. Map 8. B. correcta is known only from Burma, Thailand, Sri Lanka, and four states of India (Bihar, Madras, Mysore, and Punjab). distribution.

115

Bactrocera

Bactrocera correcta ▲ Bactrocera tryoni

o

Map 8. Distribution of Bactrocera correcta, B. cucurbitae, B. dorsalis, B. tryoni, B. zonata, and Dacus bivittatus.

Weems (1987) indicated that B. correcta has been recorded from a wide range of hosts, including the genera Acbras, Citrus, Coffea, Eugenia, Mangifera, Prunus, Psidium, Ricinus, Santalum, Syzygium, and Zizyphus. Weems (1987) also stated that some plants in the genera Aegle, Carey a, Ficus, Lagenaria, and Manilkara hosts.

must be considered as potential hosts. Almost nothing is known about the biology of this distinctive spe¬ cies of Bactrocera. B. correcta has been field-collected eight times in North America, all in Orange County, California, in the period from early August 1986 to early November 1989. The three adult male flies were caught in traps baited with methyl eugenol. An eradication treatment was applied. These adults most likely were intro¬ duced into California as larvae in host fruits by air passengers or mail from countries discussion.

where this species is known to occur.

Bactrocera cucurbitae (Coquillett) Melon fly (Fig. 166; Map 8) Dacus cucurbitae Coquillett 1899b: 129 (male and female syntypes, NMNH; Honolulu, Haw.). - Banks 1912: 32 (description of larva). - Back and Pemberton 1914: 269 (biology). - Cole 1927: 444 (male genitalia). - V. T. Phillips 1946: 35, 100 (description biology larva; hosts). - Hardy 1949: 181 (description of adults and immature stages). - Hagen and Finney 1950: 735 (biology). - Hennig 1952: 208 (in catalog of immature stages). - Finney 1953: 11 (review). - Peterson 1953: 331, 333, 337 (figures of larva). - Messenger and Flitters 1954: 756 (biology). Commonwealth Inst. Entomol. 1956: Map No. 64 (distribution). - Harper 1957:

116

Systematic Treatment of the Genera

134, 141 (history, Calif.). - Christenson 1958: 11 (review of research). - Flitters and Messenger 1958: 7 (biology). - Christenson and Foote 1960: 171 (biology). — Foote and Blanc 1963: 14 (review, Calif.). - Wasbauer 1964: 4 (review). — Weems 1964b: 1 (review). - Cunningham 1966: 865 (sex of pupae). - Kamasaki 1970: 1353 (associates). - Blanc 1970: 72 (status in Calif.). — Bateman 1972: 493 (biology). - Hardy 1973: 38 (taxonomy). - Hardy 1974: 27 (taxonomy). Wharton and Marsh 1978: 148 (parasite). - Berg 1979: 16, pi. 10 (in key, larvae). - Sharp 1979: 70 (wing dimensions). - Weems 1981b: 7 (review). Bactrocera cucurbitae: Greene 1929: 492 (description of larva, pupa). P. cucurbitat [error]: Kamasaki 1979: 1353 (associates). recognition.

The wing of cucurbitae is distinctive in having a somewhat wider

costal band than dorsalis, terminating apically in a large, rounded dark spot covering the apical third of vein R4+5 and filling the apex of cell r4+5 (fig. 166, a). The wing also bears a rather extensive dark spot covering vein dm-cu, which extends to the posterior wing margin in the apex of cell cual (fig. 166, b). The dark mark covering cell cup does not attain the posterior margin of the wing. The body color is predomi¬ nantly rufous with three yellow postsutural vittae on the scutum, and the legs are almost entirely yellow. distribution.

Map 8. B. cucurbitae is widespread and common throughout

the Oriental Region and much of the Pacific, including Hawaii. It is also found in Mauritius, East Africa, Kenya, and Tanzania. hosts.

Weems (1964b) indicated that more than 125 species of plants are af¬

fected. This species is one of the most important pests of vegetables wherever it occurs, especially cucurbits, tomatoes, and peppers, and at least three genera of wild plants: Mormordica, Passiflora, and Sicyos. discussion.

B. cucurbitae was first collected in North America as a trapped

female at Westwood, Los Angeles County, Calif., in 1956. Since then, single speci¬ mens were trapped in 1984 at Pasadena, in 1985 in San Diego and Pasadena, in 1986 at Los Angeles and Hollywood, in 1989 at West Covina, and in 1990 near Los Angeles International Airport. None of these finds represented an established infesta¬ tion. Because of the common occurrence of this fly in Hawaii, these specimens probably emerged from host fruits or vegetables which originated there and were introduced to California by mail or in passenger baggage.

Bactrocera dorsalis (Hendel) Oriental fruit fly (Figs. 13, 14, 169; Map 8) Dacus dorsalis Hendel 1912: 18 (male and female syntypes, DEI; Koshun, Taiwan). - E. Cameron 1941: 14 (parasite). - V. T. Phillips 1946: 110 (hosts). - Hardy 1949: 181 (description of larva, pupa, adult). - Carter 1952: 274 (recent re-

117

Bactrocera

search). — Hennig 1952: 208 (in catalog of immature stages). — Carter 1953: 551 (review, world). — Finney 1953: 11 (review). — Fullaway 1953: 148 (review, Hawaii). — Peterson 1953: 331, 333, 337 (figures of larva). - Messenger and Flitters 1954: 756 (biology). — Christenson 1958: 11 (review of research). — Flitters and Messenger 1958: 7 (biology). — Ebeling 1959: 245 (review, world). — Christenson and Foote 1960: 171 (biology). — Commonw. Inst. Entomol. 1960: Map No. 109 (distribution). — Harper 1961: 126, 130 (history, Calif.). — Christenson 1963: 441 (review, current investigations). — Foote and Blanc 1963: 15 (review, Calif.). — Wasbauer 1964: 3 (review). - Weems 1964a: 1 (review). — Steiner 1965: 374 (methodology). - Cunningham 1966: 865 (sex of pupae). — Blanc 1970: 72 (status in Calif.). - Kamasaki 1970: 1353 (associates). - Bateman 1972: 493 (biology). - Hardy 1973: 41 (taxonomy). — Hardy 1974: 29 (taxon¬ omy). — Wharton and Marsh 1978: 148 (parasites). — Berg 1979: 17, pi. 9 (in key to larvae). - Sharp 1979: 70 (wing dimensions). - Weems 1981b: 5 (review). Heppner 1988: 1 (review).

B. dorsalis is one of many species of Bactrocera with a very nar¬ row costal band (fig. 169, a) not crossing vein R4+5 along most of the length of cell or filling the apex of cell r4+5. The wing bears a dark area along cell cup which does not attain the hind margin of the wing, and vein dm-cu has no dark marking whatever. The face bears a pair of large black spots; the scutum has two postsutural yellow stripes; the abdomen is yellow to orange with transverse black bands on tergites 2 and 3 and a central longitudinal black stripe extending down tergites 3-5; recognition.

the legs are entirely yellow; and the aculeus is 4.5-5.0 mm long.

distribution.

Map 8. B. dorsalis is widespread in the Oriental Region and also

occurs in the Hawaiian Islands, Micronesia, and the Moluccas.

hosts.

Weems (1964a) indicated that more than 150 plant species have been

infested by dorsalis.

Economically, dorsalis is one of the most important fruit flies in Southeast Asia. Since its establishment in Hawaii in 1946, it has been field-collected repeatedly in California. The first trappings were in Orange and Santa Barbara counties in 1960. Subsequent field collections were made in the Los Angeles basin (east to San Bernardino County and north to Ventura County) and in the San Diego and Santa Clara Valley areas in the years 1966, 1972, and 1974, and in every year since then. One male was trapped in Sacramento County in May 1989, and one female was trapped in Contra Costa County in 1990. Successful programs have eradicated every infestation thus far. Most introductions likely have been from Hawaii, where this fly heavily infests a great variety of fruits and some vegetables. Quarantine interceptions are numerous in passenger baggage and mail shipments. DISCUSSION.

Systematic Treatment of the Genera

Bactrocera latifrons (Hendel) Malaysian fruit fly, pepper fruit fly, solanaceous fruit fly (Fig. 170) Chaetodacus latifrons Hendel 1915: 425 (syntypes male and female, DEI; Taiwan). — Hardy and Adachi 1956: 17 (taxonomy, hosts, distribution). — Hardy 1973: 44 (taxonomy). — E. J. Harris 1989: 75 (pest status, Hawaii). recognition.

In latifrons, the narrow costal band is similar in width to that of

dorsalis, but it expands over the apical third of vein R4+5, where it is considerably wider than near the pterostigma. The dark area covering cell cup is somewhat larger than in dorsalis and extends to the posterior margin of the wing. Vein dm-cu is not covered by a darkened area. The scutum bears two dark yellow lateral postsutural stripes on a very dark brown background. There are no other distinguishing marks on the scutum or on the rufous abdominal tergum, and the legs are yellow except for a preapical ventral brown mark on each femur. The apex of the aculeus is distinctly trilobed. distribution.

Not mapped. B. latifrons is found in Taiwan, India, Pakistan,

Malay Peninsula, Thailand, Laos, and Sri Lanka and has recently been introduced to Hawaii. hosts.

B. latifrons has been reared from Solanum incanum, S. indicum, S. ver-

bascifolium, S. sarmentosum, Baccaurea motleyana, Ponderosa tomato, and chili peppers in various parts of its range. It is also known to infest eggplant. discussion.

Adults of B. latifrons have not yet been encountered or field-

collected in the United States. It was found in Hawaii for the first time early in 1983 infesting chili peppers grown mainly by Indochinese residents in Honolulu. B. lati¬ frons is potentially able to infest commercially grown tomatoes, eggplant, and other vegetables grown in large quantities in California. Larvae in peppers have been intercepted in mail entering California from Hawaii.

Bactrocera tryoni (Froggatt) Queensland fruit fly (Fig. 171; Map 8) Tepbritis tryoni Froggatt 1897: 410 (female holotype, depository not known; Queensland, Australia). Dacus tryoni: Drew 1974: 85 (description, figure male). — Drew et al. 1978: 46 (description, biology, economic importance). - Weems 1965a: 1 (review). B. tryoni is medium-sized with a yellowish head and a rather narrow frons. The scutum is reddish brown with fuscous markings, yellow postrecognition.

119

Bactrocera

pronotal lobes, and a row of narrow yellow postsutural stripes; the scutellum is almost entirely yellow. The abdominal tergum is reddish brown with fuscous mark¬ ings, and a pair of shining yellow-brown spots occurs on the fifth tergite of the male. The apical half of the second tergite is amber to whitish. The legs are yellowish, the femora lack distinct brown markings, and the hind tibiae are darker than the general leg color. B. tryoni differs in several respects from all others treated here, especially in the presence of the paired abdominal spots and the darkened hind tibiae. It has many characters in common with other Australian Dacinae and is not easily distinguished from them. Map 8. B. tryoni is endemic in the western third of Queensland and New South Wales and the western tip of Victoria in Australia. In the past 15 distribution.

years, it has spread to Papua New Guinea, New Caledonia, Austral Islands, and many of the Society Islands. In 1971, this fly was detected on Easter Island, where it was promptly eradicated. In Queensland, tryoni has been recorded from more than 100 host plants. All commercial fruit crops (except pineapple and strawberries) and numerous wild fruits, including species of Eugenia and Passiflora (Weems 1965a), are attacked. hosts.

B. tryoni is the most destructive fruit and vegetable pest in Aus¬ tralia. According to Drew et al. (1978), it appears to be an “ecological homologue” of dorsalis in that the two species are of equal pest status, infest the same range of host plants, and possess similar ecological traits. Only one specimen of the Queens¬ land fruit fly, an adult female, has been field-collected in America north of Mexico. It was captured in a fruit fly trap at La Mesa, San Diego County, Calif., in October, 1985, but no established infestation was found there. It probably emerged from an infested fruit carried into the area from Australia, New Guinea, or New Caledonia discussion.

by an air passenger or in a mail shipment.

Bactrocera zonata (Saunders) Peach fruit fly (Fig. 168; Map 8) Dasyneura zonata Saunders 1841: 61 (holotype, sex not known, BMNH; Bengal, India). Dacus zonatus: Hardy 1973: 54 (taxonomy). The costal band of the wing of zonata is comparatively lightcolored, definitely yellowish at the base of the wing, evanescent along the outer third of cell r2+3, and expanding at the apex of vein R4+5 to a small but noticeable darker spot which appears to be more or less isolated from the costal band. The dark marking in cell cup attains the hind margin of the wing. The face bears two rounded black spots, the scutum has two presutural yellow stripes and a broad rufous post¬ sutural area, the abdominal tergum is almost all yellow to rufous with a narrow recognition.

120

Systematic Treatment of the Genera

black transverse band at the base of the third tergite and a narrow central longitudi¬ nal one from the base of the fourth almost to the apex of the fifth tergite, and the legs are entirely yellow. distribution.

Map 8. B. zonata is widespread over India, Pakistan, and gener¬

ally over Southeast Asia. To date it has not been found in the Hawaiian Islands. hosts.

B. zonata

infests primarily peach and mango but attacks other fruits as

well.

discussion.

The trapping of two males in March 1984 at El Segundo, three

males in Westchester and one male in Lennex in 1987, and one male in August 1990 in Hollywood, Los Angeles County, Calif., represent the only field collections of this fruit fly in the Western Hemisphere. The traps were baited with methyl eugenol. These adult flies no doubt emerged from host fruits such as peach, mango, or orange carried by air passengers from countries where this species occurs. An eradication program using the male annihilation technique was conducted.

Genus Callachna Aldrich Oedaspis (part): Loew 1873: 263 (taxonomy); 328 (in key). - Aldrich 1905: 606 (in catalog). Trypeta (Oedaspis) (part): Osten Sacken 1878: 191 (in catalog). Procecidochares (part): V. T. Phillips 1923: 122 (in key); 136 (review). Callachna Aldrich 1929: 11 (type species, Trypeta gibba Loew 1873: 260, by origi¬ nal designation). - Curran 1932b: 4 (in key). - Curran 1934: 291 (in key). - Foote 1960c: 671 (taxonomy). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 111 (hosts). — Freidberg 1984: 129 (gall biology). Callachna quite closely resembles Cecidocharella and Stenopa in possessing a shining, swollen black scutellum, black body bristles, and a wing pat¬ recognition.

tern of dark stripes strongly contrasting with a clear wing. In contrast to those two genera, Callachna possesses a paired patch of white bristles on the lower outer corners of the face (fig. 33, b), a character not possessed by any other North Ameri¬ can genus of fruit flies. The wing pattern is reminiscent of Procecidochares, but a

Figure 172. Right wing, Callachna gibba (Lw.).

120

Systematic Treatment of the Genera

black transverse band at the base of the third tergite and a narrow central longitudi¬ nal one from the base of the fourth almost to the apex of the fifth tergite, and the legs are entirely yellow. distribution.

Map 8. B. zonata is widespread over India, Pakistan, and gener¬

ally over Southeast Asia. To date it has not been found in the Hawaiian Islands. hosts.

B. zonata

infests primarily peach and mango but attacks other fruits as

well.

discussion.

The trapping of two males in March 1984 at El Segundo, three

males in Westchester and one male in Lennex in 1987, and one male in August 1990 in Hollywood, Los Angeles County, Calif., represent the only field collections of this fruit fly in the Western Hemisphere. The traps were baited with methyl eugenol. These adult flies no doubt emerged from host fruits such as peach, mango, or orange carried by air passengers from countries where this species occurs. An eradication program using the male annihilation technique was conducted.

Genus Callachna Aldrich Oedaspis (part): Loew 1873: 263 (taxonomy); 328 (in key). - Aldrich 1905: 606 (in catalog). Trypeta (Oedaspis) (part): Osten Sacken 1878: 191 (in catalog). Procecidochares (part): V. T. Phillips 1923: 122 (in key); 136 (review). Callachna Aldrich 1929: 11 (type species, Trypeta gibba Loew 1873: 260, by origi¬ nal designation). - Curran 1932b: 4 (in key). - Curran 1934: 291 (in key). - Foote 1960c: 671 (taxonomy). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 111 (hosts). — Freidberg 1984: 129 (gall biology). Callachna quite closely resembles Cecidocharella and Stenopa in possessing a shining, swollen black scutellum, black body bristles, and a wing pat¬ recognition.

tern of dark stripes strongly contrasting with a clear wing. In contrast to those two genera, Callachna possesses a paired patch of white bristles on the lower outer corners of the face (fig. 33, b), a character not possessed by any other North Ameri¬ can genus of fruit flies. The wing pattern is reminiscent of Procecidochares, but a

Figure 172. Right wing, Callachna gibba (Lw.).

121

Callachna

supernumerary vein runs between the costa and vein R2+3 (fig. 172, a), a character shared by very few other fruit fly genera. The lateral margins of the scutum appear to be swollen and have shiny black areas that contrast markedly with the tomentose central portion. The frons is more than two times as wide as either compound eye. In at least some specimens, a second pair of dorsocentral bristles lies anterior to the suture as in Procecidochares atra. The monotypic genus Callachna is apparently wholly North Ameri¬ can in distribution because no other species properly belonging in the genus have discussion.

been found elsewhere.

Map 9. Distribution of Callachna gibba, Cecidocharella borrichia, and Ceratitis capitata.

122

Systematic Treatment of the Genera

Callachna gibba (Loew) (Figs. 29, 33, 36, 172; Map 9) Trypeta gibba Loew 1873: 260 (female holotype, MCZ; Texas). - Osten Sacken 1880: 53 (host). - Riley and Howard 1890: 349 (parasite). - Webster 1894: 38 (parasite). - L. O. Howard 1903: 139 (parasite). - V. T. Phillips 1923: 137 (taxon¬ omy). Oedaspis gibba: Loew 1873: 263, 329 (taxonomy). - F. H. Snow 1903: 219 (Kans.). - Aldrich 1905: 606 (in catalog). - M. T. Cook 1907: 10 (biology). Trypeta (Oedaspis) gibba: Osten Sacken 1878: 191 (in catalog). Cecidochares gibba: Bezzi 1910: 25 (review). Procecidocbares gibba: Bezzi and Tavares 1916: 162 (in key). Trypeta gibbosa [error]: Viereck 1917: 453 (parasite). Procecidocbares polita [misidentification]: V. T. Phillips (part) 1923: 136 (in key). Procecidocbares atra [misidentification]: V. T. Phillips (part) 1923: 137 (review). Callacbna gibba: Aldrich 1929: 2, 11 (review). — Brimley 1938: 384 (N. Car.). — V. T. Phillips 1946: 108 (host). — Foote 1960b: 254 (nomenclature). — Foote 1965a: 660 (in catalog). — Harris and Piper 1970: 134 (host). — Wasbauer 1972: 111 (hosts). recognition.

DISTRIBUTION.

hosts.

As for the genus. Map 9.

At least three species of Ambrosia have been recorded as hosts (Wasbauer

1972). This species probably forms galls on its host plants; species of related genera are known to be gall-makers. Almost nothing is known about the biology of gibba, which is rather remarkable in view of its wide distribution and distinctive appearance. discussion.

Genus Cecidocharella Hendel Cecidocbarella Hendel 1936: 74 (type species, Cecidocharella elegans Hendel 1936: 75, by monotypy). - Aczel 1949: 189 (in catalog). - Aczel 1953: 115 (in key). Foote 1967b: 57.18 (in catalog). - Bush and Huettel 1970: 89 (revision, key to known species). — Wasbauer 1972: 111 (hosts). recognition.

The wing pattern of both sexes (fig. 173) is similar to that of

Stenopa, with bold dark bands on a nearly transparent wing. Stenopa may be distinguished easily from Cecidocbarella by the characters given in the key. A dis¬ tinctly swollen, shining black scutellum is characteristic of both genera as it is of Callacbna, Procecidocbares, and Procecidocbaroides, but the wing pattern distin¬ guishes Cecidocbarella from the latter three genera. The dorsocentral bristles are

123

Ceratitis

Figure 173. Right wing, Cecidocharella borricbia Bush and Huettel.

situated well behind the suture, there are three pairs of frontal bristles and two pairs of scutellars, and almost all the longer body bristles are black. Cecidocharella, known to occur in Mexico, Argentina, Paraguay, and Brazil, also may occupy intervening areas in Central America and continental South America. Other similar genera occur in the Neotropical Region. discussion.

Cecidocharella borrichia Bush and Huettel (Figs. 38, 173: Map 9) Cecidocharella borrichia Bush and Huettel 1970: 89 (male holotype, NMNH; Port Isabel, Texas). recognition.

As

for

the genus.

Map 9. C. borrichia is thought to occur well into Mexico but probably not farther south. Its host plant occurs in sandy marshes, dunes, and prairies on the coastal plain from southern Virginia to Florida and around the Gulf; the species probably is restricted to that geographical area. distribution.

host.

Borrichia frutescens

is the only known host.

Bush and Huettel (1970) briefly discussed the biology of borrichia and presented some cytological information. The immature stages have never been described. Only two other species of Cecidocharella are known; both occur only in discussion.

South America.

Genus Ceratitis Macleay Ceratitis Macleay 1829: 482 (type species, Ceratitis citriperda Macleay 1829: 482, by monotypy) (= capitata (Wiedemann)). - Aldrich 1905: 600 (in catalog). Williston 1908: 283 (in key). - Coquillett 1910: 519 (type data). - Hendel 1914b: 76 (in key). - Hendel 1927: 58 (review, Palearctic Region). - Curran 1932b: 3 (in key). - Foote 1965a: 672 (in catalog). - Foote and Freidberg 1980: 30 (type data). - Maddison and Bartlett 1989: 31 (zoogeography). Ceratites [error]: Williston 1896: 120 (in key).

124

Systematic Treatment of the Genera

recognition.

Flies of the genus Ceratitis are medium-sized and have a swollen,

shiny, blackish scutellum with yellow markings, capitata-Yike wing markings with yellowish areas between the brown bands, and various formations of black spots in the cells of the basal half of the wing. This genus contains so many diverse characters that it is not in the purview of this handbook to discuss them in detail. The distin¬ guishing characters of capitata and remarks on its relationships within the genus are presented in the following account of that species. Worldwide, the genus Ceratitis comprises several subgenera, the taxonomic status and relationships of which are still under discussion by world discussion.

authorities on the group. Ceratitis appears to be closely related to a number of other genera, mainly African, but the exact status of these relationships also is unsettled at present. Ceratitis contains several species with comparatively restricted geographic and host ranges; some of these are of considerable economic importance where they occur. Except for C. malgassa Munro, which was reported from Puerto Rico by Steyskal (1982) and has never been found again, capitata is the only Ceratitis species ever encountered in the New World, probably because of its wide-ranging habitats and the fact that its hosts are involved in world-wide commerce.

Ceratitis capitata (Wiedemann) Mediterranean fruit fly, medfly (Figs. 30, 32, 35, 174; Map 9) Trypeta capitata Wiedemann 1824: 55 (male syntype(s), ZMUC; “India orient.” [prob. error]). — Coquillett 1910: 519 (type data). Ceratitis capitata: McCallan 1891: 120 (biology). - Riley 1891: 255 (Bermuda). Riley and Howard 1891a: 5 (host, biology, Bermuda). - Johnson 1904a: 79 (Ber¬ muda). - Aldrich 1905: 601 (in catalog). - Coquillett 1910: 519 (type data). Banks 1912: 31 (description of larva). - Johnson 1913a: 450 (Bermuda). - Severin 1913a: 68 (introduction). - Severin 1913b: 399 (life history). - Back 1914: 1 (biology). — Hendel 1914b: 76 (type data). - Hendel 1914c: 10 ( review, Neotropi¬ cal Region). - Ogilvie 1927: 38 (parasite). - Ogilvie 1928a: 44 (history, Bermuda). - Ogilvie 1928b: 289 (parasite). - Ogilvie 1928c: 30 (parasite). - C. T. Greene 1929: 492 (description of larva, pupa). - Hume 1929: 29 (review, Fla.). - Kellogg et al. 1929: 146 (review, Fla.). — M. H. Walker 1929: 10 (review). — Newell 1930: 512 (review, Fla.). - Strong 1930: 509 (history, Fla.) - Gjullin 1931: 248 (survival in U.S.). — Keck 1931: 86 (cage tests). - Newell 1931a: 10 (history, Fla.). - Newell 1931b: 49 (history, Fla.). - Quayle 1931: 1064 (biology). - Bates 1934b: 1 (mor¬ phology of larva). - Curran 1934: 288, 294 (figures of wing, head). - Quayle 1938: 228 (review, world). - Waterston 1941: 3 (hosts). - Waterston 1944: 56 (reviews, Fla., Bermuda). - V. T. Phillips 1946: 36, 108 (description, biology larva; hosts). - Aczel 1949: 191 (in catalog). - Hardy 1949: 181 (description of all stages). - Hagen and Finney 1950: 735 (biology). - Commonw. Inst. Entomol.

125

Ceratitis

1951: Map No. 1 (distribution). - Hennig 1952: 215 (in catalog of immature stages). — Carter 1953: 555 (biology). — Finney 1953: 11 (biology). — Peterson 1953: 333, 337 (figures of larvae). - Messenger and Flitters 1954: 756 (biology). Brogdon 1956: 23 (host). - Christenson et al. 1956: 1 (status of research). Denmark 1956: 85 (review, Fla.). - Ayers 1957: 67 (history, Fla.). - Ayers and Rohwer 1957: 12 (history, Fla.). - Oberbacher and Denmark 1957: 6 (history, Fla.). — Shepherd 1957: 101 (history, Fla.). — Steiner 1957: 508 (trap). — Steiner et al. 1957: 505 (lures). — Wolfenbarger 1957: 29 (history, Fla.). — Christenson 1958: 11 (review of research). — Flitters and Messenger 1958: 7 (biology). — Rohwer 1958: 23 (history, Fla.). - Simanton 1958: 679 (lures). - Ebeling 1959: 244 (review, world). — Christenson and Foote 1960: 171 (biology). — Steiner et al. 1961: 30 (lures, summary of research). — Cowperthwaite 1962: 19 (review, Fla.). Christenson 1963: 441 (review of current research). - Wasbauer 1964: 5 (review). — Foote 1965a: 672 (in catalog). — Cunningham 1966: 864 (sex of pupa). — Stephenson and McClung 1966: 374 (review, Tex.). - Kamasaki 1970: 1353 (asso¬ ciates). - Bateman 1972: 499 (ecology). - Rhode 1976: 11 (review, N.A.). Wharton and Marsh 1978: 148 (parasites). - Berg 1979: 18, pis. 7, 8 (in key, illustrations of larva). - Prokopy and Hendrichs 1979: 642 (mating behavior). Sharp 1979: 70 (wing dimensions). - Hagen et al. 1981: 5 (review). - Poinar and Hyslop 1981: 641 (nematode parasites). - Prokopy 1981b: 5 (distribution). - J. Walsh 1981: 1221 (review, Calif.). - Weems 1981a: 1 (review). - Weems 1981b: 2 (review). - Burditt and Baranowski 1982: 85 (review). - Jordan 1982: 70 (review, Calif.). - Steyskal 1982: 165 key to species). - Am. Soc. Hortic. Sci. 1983: 38 (review, Calif.). - APHIS 1983: 1 (review, Calif.). - J. C. Davis et al. 1983: 322 (ultrastructure of eye). - Rohwer and Williamson 1983: 41 (pest risk evaluation). Sharp et al. 1983: 302 (flight studies). - Spider et al. 1984: 1441 (host data). Heppner 1985: 1 (description of larva). - Elzinga and Broce 1986: 208 (labellar morphology). — Liquido et al. 1991: 1 (hosts). RECOGNITION.

The following characters together distinguish capitata from all

other species of Tephritidae occurring in North America: small to medium-sized, brightly colored flies; scutellum swollen, rounded above, shiny black with a thin sinuate yellow steak near base dorsally; scutum yellowish with numerous black areas in a characteristic pattern; abdomen yellowish with two narrow transverse lightcolored bands; wing relatively broad in comparison with its length (fig. 174), cloudy yellow, with three brown bands on apical two-thirds, all separated from each other, and smaller dark irregular-shaped streaks within the cells in the proximal half; cell

Figure 174. Right wing, Ceratitis capitata (Wied.).

126

Systematic Treatment of the Genera cup with its apical extension expanded as shown in fig. 32, a; and males with a black, diamond-shaped expansion of the apex of the anterior orbital bristle (fig. 35, a). These characters also distinguish capitata from all other species in the genus wherev¬ er they may occur worldwide. Several species of the subgenus Ceratitis closely resem¬ ble capitata in the thoracic pattern, the apical expansion of the elongation of cell cup, the presence of dark markings in the basal half of the wing, and in having the anterior orbital bristle of the male modified in some way. In capitata, it is black and resembles a diamond apically as shown in fig. 35 rather than some other shape. Species in other genera closely related to Ceratitis also generally resemble capitata in wing pattern and habitus but differ in not having the characteristic capitata-like scutal or scutellar pattern or the modified male orbital bristle. Details of the wing pattern also differ in these species, and some of them may possess prominent patches of long, expanded setae (“feathering”) on some or all of the male femora. Map 9. The medfly has occurred in Hawaii since its accidental introduction in 1910. It was introduced but eradicated in Florida in 1929-1930, distribution.

1956-1957, 1962—1963, 1981, and 1990; in Texas in 1966; and several times in California, with a major infestation there in 1980-1982. hosts.

C. capitata has been reported to infest more than 260 different fruits,

flowers, vegetables, and nuts. Weems (1981a) listed these in the following categories: heavily or generally infested, 42; occasionally infested, 15; rarely infested, 25; labo¬ ratory infestations, 21; unknown importance, the remainder. Liquido et al. (1991) prepared an annotated world review of the hosts of the medfly, which includes plant species belonging to almost 180 genera. C. capitata is probably the world’s most economically important fruit fly, partly because of its ability to infest a wide variety of host plants, some of discussion.

which figure largely in world trade, and partly because of its unusual ability to expand its range wherever it succeeds in establishing itself. C. capitata has been the object of intense eradication efforts because of its potential for markedly reducing the value of commercially valuable fruit. We have not attempted to record all of the literature references referring to capitata in the United States because numerous excellent reviews are available, many of which have been referenced in the synonymy presented above. The bibliographies contained in these reviews significantly expand the material we have presented.

Genus Chaetostomella Hendel Chaetostomella Hendel 1927: 124 (type species, Trypeta onotrophes Loew 1846: 498, by original designation) ( = lurida (Loew)). - Steyskal and Foote 1977: 146 (taxonomy). - White 1986: 147 (nomenclature). Sitarea Robineau-Desvoidy: Foote 1984: 123 (in catalog).

Chaetostomella Medium-sized yellowish flies with dark body markings and strongly banded wings, distinctly convergent posterior orbital bristles, a flat scutellum, and a habitus similar to that of Orellia species. The single pair of dorsocentral bristles are situated in or slightly behind a transverse line through the supra-alar bristles. Chaetostomella is separable from Neaspilota and Orellia, both of which have convergent posterior orbital bristles, by the presence of well-developed setae on the anterior oral margin in addition to the usual genal bristle and a black spot on the recognition.

scutellum. discussion.

Steyskal and Foote (1977) compared C. undosa with the type spe¬

cies, onotrophes. The genus is principally Palearctic in distribution with about 10

o •

Chaetostomella undosa Chetostoma californicum

▲ Chetostoma ruhidum

Map 10. Distribution of Chaetostomella undosa, Chetostoma californica, and C. rubidum.

128

Systematic Treatment of the Genera

Figure 175. Right wing, Cbaetostomella undosa (Coq.).

species in Europe, North Africa, the Near East, and the former USSR. Foote (1984) treated Cbaetostomella as a junior synonym of Sitarea Robineau-Desvoidy, 1830, but White (1986) considered the latter a synonym of Orellia.

Chaetostomella undosa (Coquillett) (Figs. 45, 175; Map 10) Trypeta undosa Coquillett 1899c: 262 (female holotype, NMNH; Colo.). - Aldrich 1905: 605 (in catalog). Orellia undosa: McFadden and Foote 1960: 256 (revision). - Foote and Blanc 1963: 39 (review, Calif.). - Foote 1965a: 671 (in catalog). - Wasbauer 1972: 121 (hosts). Cbaetostomella undosa: Steyskal and Foote 1977: 147 (description, taxonomy). Steck 1981: 92 (biology). — Steck 1984: 669 (biology, hosts, description of larva). - Pemberton et al. 1985: 791 (hosts). - Goeden and Ricker 1986c: 957 (host).

C. undosa was originally included by McFadden and Foote (1960) in Orellia because of its similarity to those species. However, in addition to the recognition.

characters given for the genus, undosa may be separated from Orellia occidentalis, which it strongly resembles, by the wing bands, which are much bolder and darker and have darkened borders with a noticeably lighter interior and by the postocular setae, all of which are whitish in Orellia but are mixed black and white in undosa.

DISTRIBUTION.

hosts.

Map 10.

C. undosa

has been reared from various species of

Cirsium,

but its host

range is relatively unknown.

Genus Chetostoma Rondani Cbetostoma Rondani 1856: 112 (type species, Cbetostoma curvinerve Rondani 1856: 112, by original designation). - Foote and Blanc 1963: 12 (review, Calif.).Foote 1965a: 677 (in catalog). - Foote 1984: 79 (in catalog). Cbaetostoma [emendation]: Loew 1873: 332 (taxonomy). - Hendel 1914b: 85 (in key). - Quisenberry 1949c: 81 (taxonomy).

Chetostoma Medium-sized yellowish to brown flies without black body or leg markings except sometimes on abdominal tergites; wings with four narrow bands on a hyaline field; all bristles well developed, black; three frontal bristles; two orbital bristles, the posterior pair reclinate; a single pair of dorsocentral bristles situated slightly ahead of a transverse line through the supra-alar bristles; scutellum flat; and the gena with 6-12 well-developed black setae at the anterior oral margin, the longest one-half or more as long as the genal bristle behind them. In wing pattern, Chetostoma resembles Epochra, but in that genus the antenna is longer, the wing bands are wider, and the black genal setae anterior to the genal bristle are absent. The aculeus tip in Chetostoma is transversely flattened; this character will distin¬ guish our two species from all other Nearctic tephritids except several species of recognition.

Myoleja. Very little is known about the species discussed herein. Chetostoma is of no known economic importance. The genus is represented by nine additional species in Europe and Asia; one of the North American species has been found in discussion.

Mexico.

Key to U.S. and Canadian Species of Chetostoma Preapical dark band lying upon vein dm-cu joining apical band at costa (fig. 176, a) .rubidum (Coquillett) Preapical dark band lying upon vein dm-cu extending anteriorly only to vein M or slightly beyond (fig. 177, b) .californicum Blanc

Chetostoma californicum Blanc (Fig. 177; Map 10) Chaetostoma californica Blanc 1959: 202 (male holotype, CAS: La Mesa, San Diego County, Calif.) Chetostoma California [error]: Foote and Blanc 1963: 13 (review, Calif.). Chetostoma californica: Wasbauer 1964: 8 (review). Chetostoma californicum: Foote 1965a: 677 (in catalog). - Arnaud 1979: 329 (type data). — Goeden 1986: 327 (host).

Figures 176, 177. Right wings, Chetostoma spp. 176, rubidum (Coq.); 177, californicum Blanc.

130

Systematic Treatment of the Genera recognition.

The two North American species of Chetostoma are easily recog¬

nized by their wing patterns. In californicum, the subapical band (fig. 177, b) con¬ sists only of a dark streak covering vein dm-cu; it extends only from about vein M to the posterior margin of the wing, leaving an extensive break between it and the proximal end of the apical band. In addition, the discal band tends to become evanescent within cell dm (fig. 177, a), whereas in rubidum this band is uniformly brown throughout its length. DISTRIBUTION.

host.

Map 10.

C. californicum has been reared from Lonicera albiflora in Arizona.

Chetostoma rubidum (Coquillett) (Figs. 52, 176; Map 10) Epocbra rubida Coquillett 1899c: 260 (male and female syntypes, NMNH, MCZ; Colo.). — Essig 1938: 604 (note, taxonomy). Cbaetostoma elizabetbae Quisenberry 1949c: 81 (male holotype, originally CSUFC, transferred to NMNH; Colo.). — Blanc 1959: 202 (synonymy). Myoleja rubida: Foxlee 1957: 36 (B.C.). Cbaetostoma rubida: Blanc 1959: 202 (taxonomy). Chetostoma rubida: Harper 1962: 85 (Calif.). Chetostoma rubidum: Foote 1965a: 678 (in catalog). The two species discussed herein are easily distinguished from each other by the nature of the apical and subapical wing bands. In rubidum, the recognition.

subapical band (fig. 176, a) extends unbroken completely across the wing and con¬ nects with the proximal end of the apical band, whereas this band is broadly broken in californicum (fig. 177, b). In addition, the discal band is uniformly brown throughout its length, not tending to become evanescent in cell dm as in califor¬ nicum. DISTRIBUTION.

host.

Not

Map 10.

known.

Genus Cryptotreta Blanc and Foote Cryptotreta Blanc and Foote 1961: 82 (type species, Eurosta pallida Cole 1923: 472, by original designation). - Foote and Blanc 1963: 14 (review, Calif.). - Foote 1965a: 662 (in catalog). - Steyskal and Foote 1977: 147 (taxonomy).

Cryptotreta The wing pattern of Cryptotreta (fig. 178) is reminiscent of that of some species of Eutreta in that the dark areas of the wing are punctuated by indis¬ recognition.

tinctly bordered yellowish spots. Except for scattered dark spots, the basal quarter of the wing disk is hyaline, quite unlike most Eurosta species, and the frequent oily appearance of the head and body in Eurosta is not at all evident. Three pairs of frontal bristles and two pairs of scutellars are present, and the dorsocentral bristles are inserted immediately behind the suture. The thorax is densely tomentose, and the abdomen is subshining with yellow and black markings. The oviscape is very dark with a diffuse, rounded, central orange area. As far as known, Cryptotreta includes only two species, pallida (Cole), which occurs only in Mexico, and cislimitensis Steyskal from California. discussion.

Cryptotreta cislimitensis Steyskal (Figs. 109, 178) Cryptotreta pallida Cole [misidentification]: Blanc and Foote 1961: 83 (type data). — Foote and Blanc 1963: 14 (review, Calif.). - Foote 1965a: 662 (in catalog). — Arnaud 1979: 330 (type data). Cryptotreta cislimitensis Steyskal in Steyskal and Foote 1977: 147 (male holotype, NMNH; Chula Vista, Calif.). As for the genus. Steyskal (in Steyskal and Foote 1977) detailed the characters by which cislimitensis differs from pallida (Cole), a species apparently recognition.

restricted to Baja California. distribution.

Not mapped. C. cislimitensis has been found in California at

Chula Vista, San Diego, and San Ysidro. host.

Not

known.

discussion.

C. cislimitensis was misidentified as pallida, a Mexican species, by

Blanc and Foote (1961), Foote and Blanc (1963), and Foote (1965a). Steyskal (in Steyskal and Foote 1977) described cislimitensis from the California material origi¬ nally reported by us and from specimens seen subsequent to our studies.

Figure 178. Right wing, Cryptotreta cislimitensis Steysk.

132

Systematic Treatment of the Genera

Genus Dacus Fabricius Dacus Fabricius 1805: 272 (type species, Dacus armatus Fabricius 1805: 273, by subsequent designation of Hendel 1927: 24). — Flardy 1955: 428 (taxonomy of Dacini). — Drew 1972: 11 (taxonomy of Dacini). - Munro 1984: 1 (revision, Afrotropical Region [as family Dacidae]). — Drew 1989a: 10 (subgeneric classifica¬ tion).

recognition.

Species of the distinctive genus Dacus are quite different from any

of the native New World fruit flies. They share with Bactrocera a number of charac¬ ters not found in native New World Tephritidae. These include the size of cell bm, which in all species is at least two times as wide as cell cup, and by the complete lack of ocellar, postocellar, dorsocentral, presutural, and katepisternal bristles (fig. 14). The bristle of the postpronotal lobe (fig. 14, b) is usually absent. The radial veins and cells of the wing tend to be grouped together in the anterior half so that vein R4+5 terminates well short of the wing apex (fig. 179). Cell cup is drawn out into an extremely long and narrow elongation (fig. 13, c). Dacus differs from Bactrocera, the only other similar genus found in North America, in having fused abdominal tergites (see fig. 7).

discussion.

In raising Bactrocera to full generic status, Drew (1989a, 1989b)

included the Oriental and Australasian subgenera whose species have free abdominal terga (see our discussion of Bactrocera). He distinguished between Bactrocera and Dacus, the only two dacine genera resulting from his studies, on the basis of zoo¬ geographic and host characters and on the presence or absence of fusion of the abdominal tergites. For a more comprehensive discussion of this situation, see Drew (1989a).

Dacus bivittatus (Bigot) Pumpkin fruit fly (Fig. 179; Map 8) Leptoxys bivittatus Bigot 1858: 29 (female syntype(s), MNHN; Gabon). Dacus bivittatus: Munro 1964b: 9 (review, biology). — Bot 1965: 166 (biology). Schmidt 1967: 329 (biology, host). - Cogan and Munro 1983: 519 (in catalog, Afrotropical). - Munro 1984: 33 (taxonomy). - Hancock 1989: 3 (brief review).

Figure 179. Right wing, Dacus bivittatus (Bigot).

133

Dioxyna A large species 9—10 mm long, bivittatus is most easily distin¬ guished from all the closely related species of Bactrocera by its fused abdominal tergites and by its wing pattern (fig. 179), which features a prominent costal band gradually expanding toward the wing tip. This band occupies at least the anterior half of cell r4+5 and occasionally attains vein M in the apical fourth of the wing (fig. 179, a). Three narrow yellow postsutural stripes usually are present on the scutum, the scutellum and posterior margin of the first abdominal segment are yellow, a narrow vertical yellow stripe is present on the pleuron, and doubled quadrangular yellow spots are present just below and behind the wing base. The distal half to third recognition.

of all femora are darkened. distribution.

Map 8. D. bivittatus, endemic only in tropical and subtropical

Africa, occurs from Kenya and Ghana south to Cape Province. Munro (1984) listed six plant families known to be attacked by bivittatus. It is principally a pest of cucurbits, specifically Cucumis melo, C. sativus, Cucurbita pepo, and Momordica sp. It has also been recorded infesting coffee berries. It inflicts hosts.

considerable economic damage in various parts of its range. discussion.

D. bivittatus, a distinctive African fly, has been field-collected only

once in the Western Hemisphere; a single male was caught in a methyl eugenol trap in a plum tree at Cerritos, Los Angeles County, Calif., in 1987. Increased trapping revealed no establishment of this species, thus eradicative measures were deemed unnecessary. D. bivittatus is not known to occur outside Africa, and its presence in California is probably the result of entry of infested host material by mail or pas¬ senger baggage.

Genus Dioxyna Frey Ensina Loew [misidentification]: Loew 1873: 291, 328 (review). - Aldrich (part) 1905: 610 (in catalog). - V. T. Phillips 1923: 123, 144 (in key, review). - Curran 1932b: 2 (in key). Trypeta (Ensina): Osten Sacken 1878: 193 (in catalog). Paroxyna Hendel [misidentification]: Hendel (part) 1927: 146 (revision). - F. H. Benjamin (part) 1934: 40 (review, Fla.). Dioxyna Frey 1945: 62 (type species, Trypeta sororcula Wiedemann 1830: 509, by original designation). - Munro 1957: 936 (revision). - Foote and Blanc 1963: 15 (review, Calif.). - Foote 1965a: 666 (in catalog). - Wasbauer 1972: 111 (hosts). Novak 1974b: 4 (revision). - Hardy 1988a: 77 (taxonomy). - Maddison and Bartlett 1989: 33 (zoogeography). In America north of Mexico, Dioxyna is represented by only two species, sororcula (Wiedemann) and tbomae (Curran). Both are relatively small flies RECOGNITION.

Systematic Treatment of the Genera about 3 mm long. This genus is very similar to Paroxyna but can be distinguished from it by the head, which is longer than high; by the pterostigma, which in U.S. and Canadian specimens always lacks a hyaline spot; and by the absence of setulae on the aedeagus just basal to the distiphallus. In Paroxyna the head is higher than long, a hyaline spot often is present in the pterostigma, and aedeagal setulae are present. Like Paroxyna, Dioxyna has a long, geniculate proboscis and wings with numerous round, hyaline spots, giving them a speckled appearance. Other identification characteristics are given in the key to genera. Our two species of Dioxyna can be confused with two small, dark-legged species of Paroxyna (clathrata and farinata) but can be distinguished from them by their elongated heads. discussion. Dioxyna is represented by one European species, three species in the

Oriental and Australasian Regions, and four species in the New World, two of which occur in America north of Mexico. The revision by Novak (1974b) described and illustrated these two species, provided a key, wing photographs, line drawings of male genitalia, and distribution maps. Little is known concerning the immature stages of species in this genus, but a relatively large number of hosts are known.

Key to U.S. and Canadian Species of Dioxyna Hyaline spot in apex of cell r4+5 small, not touching veins R4+5 or M (fig. 180, a); at least basal half of all femora bluish black; scutellum uniformly brownish black.sororcula (Wiedemann) Hyaline spot in apex of cell r4+5 usually extending to touch apices of veins R4 + 5 and M (fig. 181, a); at least the fore femora amber; scutellum with dark ground color but with amber to yellow apex . .thomae (Curran) Dioxyna sororcula (Wiedemann) (Figs. 62, 97, 180; Map 11) Trypeta sororcula Wiedemann 1830: 509 (syntype male(s), NMW?; Teneriffe, Ca¬ nary Is.). Ensina cbilensis Macquart 1843a: 387; Macquart 1843b: 231 (syntype female, MNHN; Chile).

a

Figures 180, 181. Right wings, Dioxyna spp. 180, sororcula (Wied.); 181, thomae (Curr.).

135

Dioxyna

Map 11. Distribution of Dioxyna sororcula and D. thomae.

Acinia picciola Bigot 1857: 347 (male holotype, UMO or MNHN?; Cuba). - Loew

1873: 291,337 (taxonomy, synonymy). - Coquillett 1899c: 264 (taxonomy, synonymy). Trypeta huwiilis Loew 1862c: 81 (male holotype, MCZ; Cuba). — Loew 1873. 291,

330 (taxonomy). - Johnson 1895b: 337 (Fla.). - Coquillett 1899c: 264 (syn¬ onymy). - Doane 1899: 188 (distribution). - F. H. Snow 1903: 219 (Kans.). F. FI. Snow 1904: 345 (Ariz.). - Aldrich 1905: 610 (in catalog). - Cresson 1907: 101 (N. Mex.). — Tucker 1907: 104 (Colo.). — Cole 1912: 158 (Calif.). Trypeta aurifera Thomson 1869: 585 (male and female syntypes, NRS?; Calif.). Loew 1873: 335, 343 (taxonomy). - Coquillett 1899c: 264 (synonymy). Benjamin 1932: 42 (synonymy).

Systematic Treatment of the Genera

Ensina aurifera: Loew 1873: 335 (taxonomy). - Aldrich 1905: 610 (in catalog). Trypeta (Ensina) humilis: Osten Sacken 1877: 345 (distribution). — Osten Sacken 1878: 193 (in catalog). Tepbritis picciola: Coquillett 1899c: 264 (synonymy). — Johnson 1904a: 79 (Ber¬ muda). Tephritis pucciola [error]: Washburn 1905: 80 (Minn.). Ensina picciola: Johnson 1913a: 452 (Bermuda). — Johnson 1913b: 84 (Fla.). — V. T. Phillips 1923: 149 (review). — Sturtevant 1925: 215 (seminal receptacles). — Olgilvie 1928a: 45 (Bermuda). — Knowlton and Harmston 1937: 145 (Utah). — Brimley 1938: 384 (N. Car.). — Strickland 1946: 166 (Alta.). Paroxyna sororcula (Wiedemann): Hendel 1927: 158 (synonymy). Tephritis humilis: Janes and Thomas 1932: 104 (Utah). Paroxyna picciola: F. H. Benjamin 1934: 42 (review, Fla.). — Needham 1946: 344 (description, biology larva; hosts). — Needham 1948: 435, 444 (biology). — Aczel 1949: 289 (in catalog). — Hennig 1952: 212 (in catalog immature stages). Dioxyna picciola: Munro 1957: 957 (revision). — Foote and Blanc 1963: 16 (review, Calif.). — Foote 1965a: 666 (in catalog). — Novak et al. 1967: 147 (hosts). — Goeden 1971: 47 (host). — Steyskal 1972a: 88 (host). — Wasbauer 1972: 111 (hosts). — Novak 1974b: 5 (taxonomy). — Goeden and Blanc 1986: 90 (hosts). — W. A. Palmer 1987: 192 (hosts). — Flardy 1988a: 77 (synonymy). Dioxyna sororcula: Hardy 1988a: 77 (taxonomy, synonymy). Dioxyna piccola [error]: Hardy 1988a: 77 (nomenclature). recognition.

D. sororcula, a brown fly, differs from thomae in that the distal

hyaline spot in cell r4+5 is relatively small and does not touch either vein R4+5 or M; the femora are bluish black; the entire scutellum is brownish black, and, in the male, the epandrium and surstyli are slightly higher than wide. In thomae, the distal hyaline spot in cell r4+5 is large and extends almost or completely across the cell to touch either or both of veins R4+5 or M, the femora are amber (in some specimens the middle and hind femora are grayish black tomentose), the apex of the scutellum is amber to yellow, and the epandrium and surstyli are distinctly wider than high. The elongate head separates sororcula from such dark-legged Paroxyna species as clathrata and farinata. Map 11. D. sororcula is widespread throughout the tropics and subtropics of the world. It also occurs in Japan and as far south as Chile in South America. distribution.

Wasbauer (1972) listed 21 plant species as hosts. About half of these belong to the genus Bidens; the others are in the genera Actinospermum, Aster, Coreopsis, Helenium, and Tagetes. hosts.

In all recent New World literature, the name picciola has been used for this species. However, we have adopted Hardy’s (1988a) synonymy on the discussion.

strength of his statement: “I have compared numerous specimens from the Oriental

137

Dyseuaresta

and Australasian regions with picciola from continental United States and have not found distinct differences in wing markings, sterna of male abdomen, or male geni¬ talia and find no characters that I would consider valid for separating species. The specimens of typical sororcula also fit all the characters of picciola given by Novak (1974b)/’ D. sororcula is a highly variable species and very probably should be accorded further study. It is commonly collected in America north of Mexico. Ben¬ jamin (1934) illustrated the egg, larva, and adult.

Dioxyna thomae (Curran) (Fig. 181; Map 11) Ensina thomae Curran 1928a: 70 (female holotype, AMNH; St. Thomas I., Virgin Is.). — Arnaud and Owen 1981: 149 (type data). Paroxyna thomae: F. H. Benjamin 1934: 41 (review, Fla.). Dioxyna thomae: Munro 1957: 938 (revision). — Foote 1965a: 666 (in catalog). — Wasbauer 1972: 112 (host). — Novak 1974b: 8 (revision). D. thomae is readily distinguished from sororcula by the charac¬ ters discussed in the Recognition section for that species. Like sororcula, thomae can be confused with Paroxyna clathrata and P. farinata but can be recognized as a species of Dioxyna by its elongate head. recognition.

distribution.

hosts.

Map 11. D. thomae also occurs in the West Indies and Guyana.

Bidens bipinnata is the only host listed by Benjamin (1934). We have seen

specimens reared from Wedelia sp. discussion.

Unlike sororcula, thomae is rarely collected in the Nearctic Region.

Genus Dyseuaresta Hendel Euaresta Loew: Loew (part) 1873: 296, 328 (taxonomy, in key). - Aldrich (part) 1905: 612 (in catalog). Trypeta (Euaresta): Osten Sacken 1878: 194 (in catalog). Dyseuaresta Hendel 1928: 368 (type species, Euaresta adelphica Hendel 1914c: 72, by original designation). - Curran 1932b: 3 (in key). - Bates 1934c: 15 (review). Benjamin 1934: 51 (review, Fla.). - Curran 1934: 293 (in key). - Aczel 1940: 235 (in key). - Foote 1960e: 72 (taxonomy). - Foote 1960f: 146 (taxonomy). - Foote 1965a: 668 (in catalog). - Foote 1967b: 22 (in catalog). - Wasbauer 1972: 112 (hosts). - Foote 1980: 26 (taxonomy, in key). Dyseuaresta closely resembles Euaresta and some species of Tephritis in having distinct hyaline spots on an evenly dark wing, a heavily tomentose recognition.

Systematic Treatment of the Genera thorax and abdomen, yellow legs, the dorsocentral bristles inserted close to the suture, and two frontal bristles. Dyseuaresta differs from both genera by the presence of only one pair of scutellar bristles. Males of Dyseuaresta are easily distinguished from those of Euaresta by the lack of swollen fore femora, and they differ from those of both Euaresta and Tephritis by the quite different aspect of the external male genitalia. In Dyseuaresta, the outer surstyli are elongate and posteriorly curved, whereas they are simple in Euaresta and Tephritis. Euaresta males have the epandrium swollen and striate. Dyseuaresta runs with some difficulty in the key (see couplet 39) because the bulla is evanescent and can be seen only in the darkest specimens. A hyaline spot is present in the apex of cell r4+5. The wing pattern is quite different from that of most species of Trupanea (compare figs. 81 and 83). discussion.

Dyseuaresta undoubtedly originated in the Neotropical Region,

where nine species occur in addition to mexicana (Foote 1967b). It is most closely related to the Neotropical genera Lamproxynella Hering and Pseudoedaspis Hendel, which have similar chaetotaxy and male genitalia. Foote (1980) discussed its poorly known relationships and indicated a possible distribution from the United States south to Brazil.

Dyseuaresta mexicana (Wiedemann) (Figs. 83, 182; Map 12) Trypeta mexicana Wiedemann 1830: 511 (male syntype(s), ZMHU; Mexico). — Osten Sacken 1858: 79 (in catalog). - Loew 1862c: 59 (taxonomy). — Osten Sacken 1862: 96 (description of male). — Loew 1873: 317 (review). Euaresta mexicana: Loew 1873: 330, 337 (taxonomy). — Aldrich 1905: 613 (in catalog). — Johnson 1913b: 84 (Fla.). Trypeta (Euaresta) mexicana: Osten Sacken 1878: 194 (in catalog). Dyseuaresta mexicana: Hendel 1928: 368 (taxonomy). — F. H. Benjamin 1934: 51 (review, Fla.). — V. T. Phillips 1946: 113 (host). — Aczel 1949: 279 (in catalog). — Aczel 1952a: 122 (in catalog). - Foote 1965a: 668 (in catalog). - Foote 1967b: 22 (in catalog). - Stegmaier 1968a: 19 (biology, hosts). - Wasbauer 1972: 112 (hosts). recognition.

distribution.

As for the genus. Map 12. D. mexicana also has been recorded from Mexico,

Cuba, Puerto Rico, St. Vincent, the Bahamas, and Paraguay, but at least some of these records may be misidentifications of other Dyseuaresta species.

Figure 182. Right wing, Dyseuaresta mexicana (Wied.).

Dyseuaresta

o

Dyseuaresta mexicana

•

Epochra canadensis

Map 12. Distribution of Dyseuaresta mexicana and Epochra canadensis.

hosts.

At least three species of Melanthera were recorded as hosts

by

Wasbauer

(1972).

D. mexicana closely resembles many poorly known forms that are found south of the United States. Its generic relationships are not clearly understood, primarily because it has not been studied at all extensively. Stegmaier (1968a) dis¬ cussed the biology of this species in Florida. The immature stages have never been DISCUSSION.

described.

140

Systematic Treatment of the Genera

Figure 183. Right wing, Epocbra canadensis (Lw.).

Genus Epochra Loew Epocbra Loew 1873: 238 (type species, Trypeta canadensis Loew 1873; 235, by monotypy). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 603 (in catalog). Coquillett 1910: 539 (type data). - Cresson 1914a: 275 (taxonomy). - V. T. Phillips 1923: 120, 133 (in key, revision). — Curran 1932b: 3 (in key). — Curran 1934: 287 (in key). - Foote and Blanc 1963: 16 (review, Calif.). - Foote 1965a: 677 (in catalog). - Wasbauer 1972: 112 (hosts). Trypeta (Epochra): Osten Sacken 1878: 189 (in catalog). Epochroa [error]: Williston 1896: 121 (in key). — Williston 1908: 286 (in key). recognition.

Small to medium-sized flies with a largely unmarked light yellow

body, banded wings, and all of the body setae and bristles black. In older specimens, the abdomen may be darkened dorsally, but no patterns on the tergites are normally evident. Presutural bristles (fig. 53, a) are lacking; the dorsocentral bristles are closer to a line through the supra-alar than through the acrostichal bristles; the posterior orbital bristles are reclinate; the scutellum is flat; and the apex of the first flagellomere nearly attains the oral margin (fig. 54, b). The wing pattern of Epocbra (fig. 183) somewhat resembles that of several species of Rbagoletis in exhibiting the four Rbagoletis-Yike bands (see the discussion of the latter genus). However, it differs from those species and all other North American genera in that the discal band terminates in the anterior wing margin distinctly distad of the pterostigma, and the subapical band terminates at or just anterior to vein R4+5. The long, fine setae on the anatergite distinguish Epocbra from all other genera in the New World. No species other than canadensis or its synonym has ever been associated with this genus, but A. L. Norrbom (unpublished data) reared a second discussion.

(undescribed) species from a Ribes species in Mexico. Epocbra is the only New World representative of the Euphrantini.

Epocbra canadensis (Loew) Currant fly, currant fruit fly, yellow currant fly, mouche du groseillier (Figs. 54, 183; Map 12) Trypeta canadensis Loew 1873: 235 (female holotype, MCZ; Canada). — Gillette 1892: 18 (biology). — C. F. Baker 1895: 174 (biology). — Felt 1898: 160 (biology). — Coquillett 1910: 539 (type data).

141

Euaresta

Epocbra canadensis: Loew 1873: 238, 329 (taxonomy). — W. A. Snow 1894: 159 (taxonomy). — Harvey 1896: 111 (review, Me.). — Doane 1898: 71 (taxonomy, biology). — Felt 1898: 160 (biology). — Harvey 1898: 25 (description, biology). — Piper and Doane 1898: 7 (description, biology). - Doane 1899: 178 (distribution). — Aldrich 1905: 603 (in catalog). — Aldrich 1909: 72 (taxonomy). — Banks 1912: 33 (description of larva). — Paine 1912: 139 (biology). - Lovett 1913: 135 (biolo¬ gy). — Viereck 1913: 567 (parasite). — Hendel 1914b 89 (type data). — Whitney 1916: 152 (review, Calif.). — Severin 1917a: 177 (review, Me.). — Severin 1918a: 201 (review, Calif.). — Cole and Lovett 1921: 325 (Oreg.). — V. T. Phillips 1923: 133 (review). — Johnson 1925b: 261 (N. Engl.). - Bird 1927: 128 (Man.). — Cole 1927: 444 (male genitalia). — Mitchener 1927: 38 (biology). — Leonard 1928: 851 (N.Y.). — Keifer 1930: 579 (description of larva). — Gambrell 1931: 226 (biology). — Janes and Thomas 1932: 103 (Utah). - Curran 1934: figs. 24, 63 (figures of wing and head). — Jones 1937: 1 (review). — Knowlton and Harmston 1937: 145 (Utah). - Pickett 1937: 59 (male genitalia). — Essig 1938: 603, 605 (review). Hatch 1938: 197 (bibliography). — Strickland 1938: 204 (Alta.). — Hering 1940c: 5 (taxonomy). — V. T. Phillips 1946: 41, 114 (description, biology larva; hosts). — Forsell 1947: 167 (Puget Sound). — Morrison 1950: 46 (biology). — Hennig 1952: 219 (in catalog immature stages). — Peterson 1953: 333,335 (figures of larva). Strickland 1953: 277 (ptilinal armature). — Frick et al. 1954: 59 (trapping). — Foxlee 1957: 36 (B.C.). - Essig 1958: 603, 605 (review). — Christenson and Foote 1960: 171 (biology). — Foote and Blanc 1963: 16 (review, Calif.). - Wasbauer 1964: 8 (review). — Foote 1965a: 677 (in catalog). — Wasbauer 1972: 112 (hosts).— Muesebeck 1980: 36 (parasite). Trypeta (Epocbra) canadensis: Osten Sacken 1878: 189 (in catalog). Epocbra lunifera Hering 1940c: 5 (holotype female, BMNH; Sumner, Wash.). Foote 1959c: 59 (synonymy). recognition.

distribution.

As

for

the genus.

Map 12. E. canadensis is not known to occur south of the United

States. hosts.

E. canadensis is apparently restricted to Ribes spp.

Genus Euaresta Loew Euaresta Loew 1873: 296 (type species, Trypeta festiva Loew 1873: 296, by designa¬ tion of Coquillett 1910: 540). - Williston 1896: 123 (in key). - Coquillett 1899c: 268 (in key). - Aldrich (part) 1905: 612 (in catalog). - Cresson 1907: 99 (taxon¬ omy). - Williston 1908: 287 (in key). - Coquillett 1910: 540 (type designation). Cresson 1914a: 276 (taxonomy). - Hendel 1914b: 96 (in key). - V. T. Phillips (part) 1923: 123, 145 (in key; review, e. U.S.). - Hendel 1928: 368 (taxonomy). -

142

Systematic Treatment of the Genera F. H. Benjamin 1934: 49 (taxonomy). - Aczel 1939: 124 (in key). - Aczel 1940: 235 (in key). - Quisenberry 1950: 11 (revision). - Aczel 1952d: 151 (synonymy). - Foote and Blanc 1959: 149 (taxonomy). - R. H. Foote 1960e: 72 (taxonomy). R. FI. Foote 1960f: 146 (taxonomy). - Foote and Blanc 1963: 17 (review, Calif.). B.A. Foote 1965: 105 (hosts). - R.FF Foote 1965a: 665 (in catalog). - Wasbauer 1972: 113 (hosts). - Cogan and Munro 1983: 545 (in catalog). - R. H. Foote 1984: 87 (in catalog). — B.A. Foote 1984: 53 (hosts). Trypeta (Euaresta): Osten Sacken (part) 1878: 194 (in catalog). Trypeta [misidentification]: Aldrich (part) 1905: 604 (in catalog). Urellia Robineau-Desvoidy [misidentification]: Aldrich (part) 1905: 613 (in catalog). Tepbritis (Euaresta): Cresson (part) 1907: 99 (taxonomy). Camaromyia Hendel 1914b: 95 (type species, Trypeta bullans Wiedemann 1830: 506, by original designation). - Hendel 1914c: 63 (taxonomy). - Hendel 1928: 368 (taxonomy). — Foote 1984: 74 (in catalog). Tepbritis Latreille [misidentification]: Curran (part) 1932b: 4 (in key). — Curran (part) 1934: 291 (in key). Euaresta (Euaresta): F. H. Benjamin 1934: 49 (review, Fla.). Euaresta (Setigeresta) Benjamin 1934: 50 (type species, Trypeta aequalis Loew 1862c: 86, by original designation). Setigeresta Benjamin: Aczel 1940: 235 (in key). recognition.

Euaresta is one of five U.S. and Canadian genera having the fol¬

lowing combination of characters: generally small flies with a dark wing filled with discrete, fairly large, rounded hyaline spots; a flat scutellum; two pairs of frontal bristles; two pairs of orbital bristles, the posterior pair yellowish and reclinate; two pairs of scutellar bristles; and a single pair of dorsocentral bristles situated very close behind the transverse suture. The genus is separable from Gonioxyna, Paroxyna, and Dioxyna by its nongeniculate labellum, and from Tepbritis (and all other North American genera) by the fact that the fore femora of the male are distinctly enlarged and the male terminalia (fig. 93) are broadened and have conspicuous striations in posteroventral view. The wing pattern of this genus resembles that of many others that have a dark wing with many hyaline spots, but the general pattern holds for most of the species. The wing pattern of Dyseuaresta may be mistaken for that of Euaresta, but in the former genus, there is only one pair of scutellar bristles, and large rounded hyaline spots are present on either side of vein r-m well separated from that vein by dark areas. discussion.

Eight species of Euaresta occur in the United States and Canada,

one of which, bullans, was probably introduced from South America. At least four other species occur south of the United States. Foote (1984) recognized Camaromyia as a separate genus, but it is considered a synonym of Euaresta here and by Foote (1980). All known hosts of the genus are species of ragweed (Ambrosia) or cocklebur (Xantbium). Several Euaresta species have been investigated as biological control agents for weedy species of these plant genera.

143

Euaresta

a

a

a

188 a

Figures 184-191. Right wings, Euaresta spp. 184, aequalis (Lw.); 185, festiva (Lw.); 186, bullans (Wied.); 187, tapetis (Coq.); 188, jonesi Curr.; 189, bellula Snow; 190, stigmatica Coq.; 191, bella (Lw.).

Key to U.S. and Canadian Species of Euaresta 1. Ground color of thorax yellow .2 Ground color of thorax dark brown to black, often obscured by whitish or yellowish tomentum.3 2. Width of gena at least 2 times width of 1st flagellomere (fig. 192, a); hyaline spot near apex of cell r4+5 separated from wing margin by a distinct brown band (fig. 184, a) .aequalis (Loew) Width of gena about equal to width of 1st flagellomere (fig. 193, a); hyaline area at apex of cell r4+5 broadly joined to wing margin (fig. 185, a) .. .festiva (Loew)

144

Systematic Treatment of the Genera

Figures 192, 193. Lateral view, left side of head, Euaresta spp. 192, aequalis (Lw.); 193, festiva (Lw.). Figures 194, 195. Dorsal view, ovipositor sheath, Euaresta spp. 194, stigmatica Coq.; 195, bella (Lw.).

3. Cell r4+5 without a bulla (fig. 186, b); usually 3 hyaline spots in cell rt distad of pterostigma (fig. 186, a); arista brown at apex, yellow at base, with central portion distinctly white, male with pedicel and first flagellomere dark brown .bullans (Wiedemann) Cell r4+5 usually with at feast a small bulla (fig. 187, c); if bulla absent, arista brown with yellowish base; 2 or 3 hyaline spots in cell r1 distad of pterostigma (fig. 187, b); antenna of both sexes entirely yellowish. .4 4. Wing with a narrow brown band from center of cell c to posterior wing margin (fig. 187, a), the hyaline area adjoining it distally usually inter¬ rupted (fig. 187, d) .tapetis (Coquillett) Wing without such markings .5 5. Hyaline spot in cell r2+3 immediately anterior to vein r-m broadly joined across vein R2+3 to hyaline area apicad of pterostigma (fig. 189, b); usually only 2 hyaline spots in cell r, distad of pterostigma, third subapical spot, if present, not crossing vein R2 f 3 .6 Hyaline spot in cell r2+3 immediately anterior to vein r-m distinctly rounded, narrowly joined to, or completely separated from, hyaline area apicad of pterostigma (fig. 190, b); 3 hyaline spots in cell r, distad of pterostigma, subapical spot usually crossing vein R, , ,.7

145

Euaresta

6. Cell br with a large, subapical, nearly quadrate hyaline spot extending completely across cell and broadly touching vein R4+5 (fig. 188, a) ... .jonesi Curran Cell br with a rounded subapical hyaline spot which is separated from vein R4+5 by infuscation of varying extent (fig. 189, a) .bellula Snow 7. Wing relatively narrow; pterostigma usually with a distinct brown spot at its center (fig. 190, a); oviscape at least 0.62 mm long, tapering (fig. 194, a); more than half of abdominal tergite 5 without pruinosity, shiny . .stigmatica Coquillett Wing relatively wide; pterostigma never with a discrete brown spot at its middle (fig. 191, a); oviscape less tapering, no longer than 0.52 mm (fig. 195, a); abdominal tergite 5 mostly pruinose, matte.bella (Loew)

Euaresta aequalis (Loew) (Figs. 18, 184, 192; Map 13) Trypeta aequalis Loew 1862c: 86 (male syntype, MCZ; Ill.). - Marlatt 1890: 168 (biology). - Marlatt 1891a: 40 (biology). - Marlatt 1891b: 80 (biology). - Marlatt 1891c: 312 (biology). Euaresta aequalis: Loew 1873: 309, 330 (taxonomy). - W. A. Snow 1894: 171 (taxonomy). - Coquillett 1899c: 264 (taxonomy). - Doane 1899: 191 (taxonomy, distribution). - Aldrich 1905: 612 (in catalog). - Washburn 1905: 118 (Minn.). Tucker 1907: 105 (Colo.). — Johnson 1910: 803 (N.J.). — Cole 1912: 158 (Calif.). -C. C. Adams 1915: 49, 189 (Ill.). - Peterson 1916: 240 etseq. (figures of head). Cole and Lovett 1921: 326 (Oreg.). -V. T. Phillips 1923: 147 (review). - Johnson 1925b: 264 (N. Engl.). - Cole 1927: 444 (male genitalia). - Leonard 1928: 853 (N.Y.). — Currie 1932: 191 (oviposition). - Janes and Thomas 1932: 103 (Utah). Petch and Maltaise 1932: 57 (Que.). - Bates 1934b: 1 (morphology of larva). Knowlton and Harmston 1937: 145 (Utah). - Brimley 1938: 384 (N. Car.). Essig 1938: 605 (taxonomy). - V. T. Phillips 1946: 43, 114 (description of larva, biology, host). - Quisenberry 1950: 17 (review). - Hennig 1952: 211 (in catalog immature stages). — Cleveland and Hamilton 1958: 213 (Ind.). — Essig 1958: 605 (taxonomy). — Foote and Blanc 1963: 18 (review, Calif.). — B.A. Foote 1965: 106 (host). - R.H. Foote 1965a: 665 (in catalog). - Lipp and Schulz 1970: 27 (dam¬ age). - Wasbauer 1972: 113 (hosts). - Hilgendorf and Goeden 1982: 152 (host). Hilgendorf and Goeden 1983: 404 (host). — B. A. Foote 1984: 53 (host). Trypeta (Euaresta) sp.: Osten Sacken 1877: 345 (Colo., taxonomy). Trypeta (Euaresta) aequalis: Osten Sacken 1878: 194 (in catalog). Tephritis aequalis: Coquillett 1899c: 264 (taxonomy). - Johnson 1900b: 688 (N.J.). - Rau 1922: 62 (biology). - Hendrickson 1930: 143 (Iowa). - Strickland 1938: 204 (Alta.). Trypeta signalis [error]: L. O. Howard 1901: fig. 101 (illustration). Tephritis gemella Coquillett 1902b: 181 (female holotype, NMNH; Las Vegas Hot Springs, N. Mex.). - C. F. Baker 1904: 30 (Calif.). - Aldrich 1905: 612 (in catalog). Ensina aequalis: F. H. Snow 1903: 219 (Kans.).

Systematic Treatment of the Genera

Map 13. Distribution of Euaresta aequalis.

Trypeta gemella: Woodworth 1913: 137 (Calif.). Camaromyia aequalis: Hendel 1914c: 63 (taxonomy). Euaresta (Setigeresta) aequalis: F. H. Benjamin 1934: 50 (type species designation). Tephritis arcualis Walker [nomen nudum]: Foote 1964d: 325 (D.C., synonymy). E. aequalis shares with festiva the unique character of a yellow thoracic and abdominal integument beneath the tomentum present there, whereas the ground color in all other species of Euaresta discussed herein is dark brown to recognition.

black. E. aequalis is generally larger than festiva, and individuals of both species are generally larger than those of the dark-bodied N.A. Euaresta species. In festiva, the hyaline area at the apex of cell r4+5 is broadly joined to the wing margin (fig. 185, a),

147

Euaresta

whereas in aequalis this area forms a rounded spot completely isolated from the wing margin by the union of the two dark apical rays surrounding it (fig. 184, a). A distinctive feature of aequalis is the very broad gena (fig. 192, a) which, in contrast to the narrower condition in festiva (fig. 193, a), is more than two times as wide as the first flagellomere. distribution.

Map 13. E. aequalis does not occur south of the United States.

Xanthium strumarium (= X. pungens) is the only known host; several varieties were recorded as hosts by Wasbauer (1972). host.

Euaresta bella (Loew) (Figs. 191, 195; Map 14) Trypeta bella Loew 1862c: 88 (male and female syntypes, MCZ; DC., N.Y.). — Loew 1873: 311 (review). — Cockerell 1898: 155 (N. Mex.). Acinia bella: Loew 1862c: 89 (nomenclature). Euaresta bella: Loew 1873: 311, 330 (taxonomy). — F. H. Snow 1894: 171 (distribu¬ tion). — Johnson 1895b: 337 (Fla.). — Doane 1899: 191 (distribution). - Chagnon 1901: 14 (Canada). — F. FI. Snow 1903: 219 (Kans.). — C. F. Baker 1904: 31 (Calif.). — F. FI. Snow 1904: 345 (Ariz.). - Aldrich 1905: 612 (in catalog). — Washburn 1905: 118 (Minn.). — Tucker 1907: 105 (Ala.). — Johnson 1910: 803 (N.J.). - Johnson 1913b: 84 (Fla.). - Hendel 1914c: 71 (review). - Winn and Beaulieu 1915: 154 (Que.). — Sturtevant 1918: 36 (Ala.). - Britton 1920: 204 (Conn.). - V. T. Phillips 1923: 147 (review). - Johnson 1925b: 264 (N. Engl.). Sturtevant 1925: 215 (seminal receptacles). — Johannsen 1926: 159 (N.Y.). — Johnson 1927: 218 (Mt. Desert). - Leonard 1928: 853 (N.Y.). - Curran 1930: 77 (N.Y.). - Flendrickson 1930: 144 (Iowa). — Curran 1932a: 9 (taxonomy). - Janes and Thomas 1932: 103 (Utah). - Petch and Maltaise 1932: 57 (Que.). - Bates 1933b: 170 (distribution). - Brimley 1938: 384 (N. Car.). - Aczel 1949: 281 (in catalog). - Quisenberry 1950: 34 (revision). - Aczel 1952a: 129 (in catalog). Aczel 1952d: 150 (synonymy). - Steyskal 1957: 94 (eye color). - Cleveland and Hamilton 1958: 213 (Ind.). - Oatman et al. 1964: 980 (Wis.). - R. H. Foote 1965a: 665 (in catalog). - Novak et al. 1967: 147 (host, biology). - Harris and Piper 1970: 134 (host). - Wasbauer 1972: 113 (host). - Morihara and Balsbaugh 1976: 692 (S. Dak., host). - Batra 1979: 118 (behavior). - Batra et al. 1981: 74 (host). - Berlocher 1984b: 354 (electrophoresis). - B. A. Foote 1984: 53 (host). Trypeta (Euaresta) bella: Osten Sacken 1878: 194 (in catalog). Euraesta [error] bella: Johnson 1900b: 688 (N.J.). Euaresta bella Fitch [error]: Hendrickson 1928: 138 (Iowa). Euaresta (Euaresta) bella: F. H. Benjamin 1934: 50 (review, Fla.). Tepbritis bella: Knowlton and Harmston 1937: 145 (Utah). Tephritidae: Ametangelo 1974: 15 (biology). Euarestia [error] bella: Elzinga and Broce 1986: 208 (labellar morphology).

Systematic Treatment of the Genera

Map 14. Distribution of Euaresta bella.

The wing pattern of bella closely resembles those of stigmatica, jonesi, and bellula in that the bulla is always present and the basal half to two-thirds recognition.

of cell r4+5 is dark in most specimens. In jonesi and bellula, there usually are two marginal hyaline spots present in cell rt; if a third, apical one is present, it is small and does not cross vein R2+3. The basal marginal spot broadly crosses this vein. In bella and stigmatica, the basal marginal spot in cell rt is always interrupted at the vein, whereas the subapical spot crosses it broadly. E. bella can be distinguished from stigmatica and the other two species by its high degree of abdominal pruinosity; in the other three species tergite 5 is mostly shiny.

Euaresta distribution. Map 14. E. bella also has been reported from Mexico, the Bahamas, and Barbados, but it is not known to occur farther south. host. Ambrosia artemisiifolia is the only known host.

Euaresta bellula Snow (Fig. 189; Map 15) Trypeta (Euaresta) stelligera Coquillett 1894: 74 (male holotype, NMNH; Los An¬ geles County, Calif.). — Quisenberry 1950: 30 (synonymy). Euaresta bellula Snow 1894: 172 (female lectotype, UKL; Ariz.). - Doane 1899: 192

• o

Euaresta bellula Euaresta festiva

Map 15. Distribution of Euaresta bellula and E. festiva.

150

Systematic Treatment of the Genera

(distribution). - F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 613 (in catalog). Woodworth 1913: 137 (Calif.). - Curran 1932a: 9 (taxonomy). - Forsell 1947: 167 (Wash.). — Quisenberry 1950: 30 (revision). — R. H. Foote 1960b: 254 (syn¬ onymy). - Byers et al. 1962: 180 (type data). - R. H. Foote 1962: 173 (type designation). - Foote and Blanc 1963: 19 (review, Calif.). - Oatman et al. 1964: 980 (Wis.). - B.A. Foote 1965: 106 (host). - R.H. Foote 1965a: 665 (in catalog). Flarris and Piper 1970: 134 (host). - Goeden 1971: 47 (host). - Wasbauer 1972: 113 (host). - Goeden and Ricker 1974b: 837 (hosts). - Goeden and Ricker 1976a: 49 (hosts). - Goeden and Ricker 1976b: 926 (host). - Berlocher 1984b: 354 (electrophoresis). — B. A. Foote 1984: 53 (host). Trypeta stelligera: Aldrich 1905: 605 (in catalog). Euaresta stelligera: Berlocher 1984b: 354 (electrophoresis, proposed resurrection from synonymy). The wing pattern of bellula closely resembles those of jonesi, stigmatica, and bella. See the Recognition section of bella to distinguish the latter two recognition.

species from bellula. In contrast to jonesi, the hyaline spot in cell br is always small, rounded anteriorly, and always separated from vein R4+5 by a dark area. distribution.

Map 15. E. bellula also occurs in Mexico along the coast of the

Gulf of California. hosts.

Ambrosia appears to be the favored host.

discussion.

Berlocher (1984b) presented strong electrophoretic evidence that

bellula comprises two distinct populations; he stated that his study material from Oregon, in addition to having physiological differences, corresponds mor¬ phologically to Coquillett’s description of stelligera (Coquillett 1894). Whether stelligera as defined by Berlocher should stand as a distinct species is a question still open to further investigation. We have not recognized it for the practical reason that this handbook relies mainly on morphological characters and the fact that only two populations of what Berlocher suggests is a species complex have been analyzed biochemically. A study of the morphological characters proposed by Berlocher to separate stelligera and bellula should be investigated in a broader sample of electrophoretically analyzed populations.

Euaresta bullans (Wiedemann) (Figs. 25, 186; Map 16) Trypeta bullans Wiedemann 1830: 506 (syntype(s), sex not stated, NMW; Mon¬ tevideo, Uruguay). Euaresta adspersa Coquillett in C. F. Baker 1904: 30 (male and female syntypes, NMNF1; Stanford University, Stanford, Calif.). — Woodworth 1913: 137 (Calif.). — Quisenberry 1950: 24 (synonymy). Camaromyia bullans: Hendel 1914b: 95 (type designation). — Aczel 1949: 295 (in

151

Euaresta

Map 16. Distribution of Euaresta bullans, E. jonesi, E. stigmatica, and E. tapetis.

catalog). — Aczel 1950: 130 (in catalog). — Hennig 1952: 211 (catalog immature stages). — Foote 1984: 74 (in catalog). Tephritis wolffi Cresson 1931: 5 (male holotype, ANSP; Pomona, Calif.). — Quisenberry 1951: 60 (taxonomy, synonymy). Euaresta (Camaromyia) bullans: Malloch 1933: 274 (review). — F. H. Benjamin 1934: 50, 58 (taxonomy). Euaresta bullans: Quisenberry 1950: 24 (revision). — Aczel 1952d: 150 (synonymy). — Munro 1957: 1010 (distribution, biology). — Foote and Blanc 1963: 20 (review, Calif.). — B. A. Foote 1965: 106 (host). — R. FI. Foote 1965a: 665 (in catalog). — Wasbauer 1972: 113 (hosts). — Hilgendorf and Goeden 1982: 152 (host). - Cogan and Munro 1983: 545 (in catalog). — Flilgendorf and Goeden 1983: 404 (host). — Goeden 1986: 326 (host). E. bullans closely resembles jonesi, bella, and bellula, especially in wing pattern, but in contrast to the pattern of those three species, that of bullans has the following characters: no visible bulla is present; a large rounded hyaline spot lies close to and on either side of vein r-m; three large rounded hyaline spots, equidistantly spaced, are present in cell r1 distad of the pterostigma; the arms of the apical Y-shaped mark are expanded markedly at the wing margin; and a comparatively wide hyaline area extending across the wing proximal to the pterostigma restricts the extent of the dark area immediately proximal to it. The dark brown color of the antenna of the male also distinguishes bullans from all of its congeners. recognition.

152

Systematic Treatment of the Genera

distribution.

Map 16. In the New World, bullans is absent from Mexico,

Central America, and northern South America but it occurs from there south to Brazil. It is found in Europe, the Middle East, and Asia (as Camaromyia bullans) (Foote 1984) and has been introduced into South Africa and Australia (Hardy and Foote 1989). hosts.

Xanthium spinosum is the only confirmed host. A record of X.

strumarium is questionable.

Euaresta festiva (Loew) (Figs. 185, 193; Map 15) Trypeta festiva Foew 1862c: 86 (male and female syntypes, MCZ; Pa.). — Foew 1873: 309 (review). — Coquillett 1910: 540 (type designation). Euaresta festiva: Foew 1873: 309, 330 (taxonomy). — Keen 1885: 55 (Pa.). — Snow 1894: 171 (distribution). — Doane 1899: 191 (distribution). — F. H. Snow 1903: 219 (Kans.). — Aldrich 1905: 613 (in catalog). — Washburn 1905: 118 (Minn.). — Tucker 1907: 105 (Kans.). - Johnson 1910: 803 (N.J.). - Hendel 1914b: 96 (type data). — Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 147 (review). — Johnson 1925b: 264 (N. Engl.). — Feonard 1928: 853 (N.Y.). - Hendrickson 1930: 144 (Iowa). — Petch and Maltaise 1932: 57 (Que.). — Brimley 1938: 384 (N. Car.). — Aczel 1939: 128 (taxonomy). — Quisenberry 1950: 19 (revision). — B. A. Foote 1965: 106 (host). - R. H. Foote 1965a: 665 (in catalog). — Novak et al. 1967: 147 (host, biology). — Harris and Piper 1970: 134 (host). — Wasbauer 1972: 113 (host). — Batra 1979: 118 (behavior). - Berlocher 1984b: 354 (electrophoresis). — B. A. Foote 1984: 53 (host). Trypeta (Euaresta) festiva: Osten Sacken 1878: 194 (in catalog). Euraesta [error] festiva: Johnson 1900b: 688 (N.J.). Tephritidae: Ametangelo 1974: 15 (biology). Only two of the Euaresta species discussed herein, festiva and aequalis, exhibit a yellow integument beneath the tomentum on the thorax and recognition.

abdomen. In all other North American species of the genus, this ground color is dark brown to black. E. festiva is a large Euaresta species with a scutum about 4.0 mm long. In contrast to the wing pattern of aequalis (fig. 184), the hyaline area at the apex of cell r4+5 in festiva broadly adjoins the apical margin of that cell (fig. 185, a). In addition to other pattern differences shown in figs. 184 and 185, the gena of festiva is comparatively narrow, being hardly wider than the 1st flagellomere (fig. 193, a), whereas in aequalis it is quite broad (fig. 192, a). DISTRIBUTION.

host.

Map 15.

Ambrosia trifida is the only known host.

153

Euaresta

Euaresta jonesi Curran (Fig. 188; Map 16) Euaresta jonesi Curran 1932a: 9 (female holotype, AMNH; DeLake, Oreg.). — Quisenberry 1950: 27 (revision). — B.A. Foote 1965: 106 (host). — R.H. Foote 1965a: 665 (in catalog). - Harris and Piper 1970: 134 (host). — Wasbauer 1972: 113 (hosts). - Arnaud and Owen 1981: 149 (type data). Tephritis jonesi: Curran 1934: 286 (figure of wing). The wing pattern of jonesi most closely resembles those of bella, bellula, and stigmatica. See the Recognition section of bella to distinguish it and stigmatica from jonesi. In contrast to bellula as well as the other two species, in jonesi a large quadrate hyaline spot occupies cell br near its apex, extends completely across recognition.

that cell, and is broadly based on veins R4+5 and M. distribution.

host.

Map 16. E. jonesi does not occur south of the United States.

Ambrosia chamissonis is the only known host.

Euaresta stigmatica Coquillett (Figs. 190, 194; Map 16) Euaresta stigmatica Coquillett 1902b: 180 (male and female syntypes, NMNH; Flagstaff, Ariz.). - Aldrich 1905: 613 (in catalog). - Quisenberry 1950: 33 (revi¬ sion). - Foote and Blanc 1963: 21 (review, Calif.). - B.A. Foote 1965: 106 (host). R.H. Foote 1965a: 665 (in catalog). - Harris and Piper 1970: 134 (host). Wasbauer 1972: 113 (hosts). - Goeden and Ricker 1974a: 831 (hosts). - Goeden and Ricker 1974b: 835 (host). - B. A. Foote 1984: 53 (hosts). recognition.

E. stigmatica invariably may be recognized among all the Eu¬

aresta species discussed herein by the presence near the center of the pterostigma of a discrete blackish spot (fig. 190, a) which extends wholly or incompletely across that cell and is surrounded by a hyaline field, rendering it a striking feature of the pattern. This character, combined with the comparatively light brown color of the wing pattern, makes this species usually recognizable without the aid of a microscope. In dorsal view, the dark brown to black oviscape tapers to its apex rather strongly, in contrast to those of many other Euaresta species. See the Recognition section of bella for additional characters to distinguish that species, bellula, and jonesi from stig¬ matica. DISTRIBUTION.

hosts.

Map 16.

B. A. Foote (1984) listed four species of Ambrosia as hosts.

154

Systematic Treatment of the Genera

Euaresta tapetis (Coquillett) (Fig. 187; Map 16) Trypeta (Euaresta) tapetis Coquillett 1894: 75 (male and female syntypes, NMNH; N. Mex.). Trypeta tapetis: Cockerell 1898: 155 (N. Mex.). Euaresta tapetis [error]: Doane 1899: 191 (distribution). Euaresta tapsetus [error]: F. H. Snow 1903: 219 (Kans.). Euaresta tapetis: Aldrich 1905: 613 (in catalog). — Cresson 1907: 105 (N. Mex.). — Tucker 1907: 105 (Colo.). — Quisenberry 1950: 22 (revision). — Foote and Blanc 1963: 22 (review, Calif.). — Foote 1965a: 665 (in catalog). recognition.

Among the wing patterns of all the Euaresta species discussed

herein, that of tapetis is unique in that an irregular and narrow but unbroken and distinct dark band extends across the wing from the costa at the center of cell 2nd c to the posterior margin (fig. 187, a). The hyaline area immediately distad of it (fig. 187, d) is usually uninterrupted as well, and together with the hyaline wing base, it lends prominence to the dark band. Despite the small size of this species, it usually can be recognized by the prominent band without the aid of magnification. The bulla (fig. 187, c) often is missing, but the aforementioned band distinguishes it from bullans, festiva, and aequalis, the only other Euaresta species occasionally or always lacking a bulla. DISTRIBUTION.

host.

Not

Map 16.

known.

Genus Euarestoides Benjamin Urellia Robineau-Desvoidy [misidentification]: Loew (part) 1873: 322, 328 (taxon¬ omy). Trypeta (Urellia) [misidentification]: Osten Sacken 1878: 194 (in catalog). Tephritis Latreille [misidentification]: Coquillett (part) 1899c: 265 (taxonomy). Aldrich (part) 1905: 611 (in catalog). Trypanea Schrank [misidentification]: V. T. Phillips (part) 1923: 123, 147 (in key, review, e. U.S.). Trupanea (Euarestoides) Benjamin 1934: 57 (type species, Trypeta abstersa Loew 1862a: 221, by original designation). Euarestoides: Hering 1941a: 164 (revision). — Quisenberry 1951: 59 (taxonomy). — Foote 1958: 288 (revision). — Foote 1959a: 16 (taxonomy). — Foote 1960e: 72 (taxonomy). - Foote 1960f: 146 (taxonomy). - Foote 1962: 175 (in key to spe¬ cies). - Foote and Blanc 1963: 22 (review, Calif.). - Foote 1965a: 669 (in catalog). - Foote 1967b: 24 (in catalog). - Wasbauer 1972: 113 (hosts.). - Foote 1980: 28 (taxonomy, in key).

155

Euarestoides Small, dark flies with three pairs of frontal bristles; two pairs of orbital bristles, the posterior pair reclinate; two pairs of scutellar bristles; scutellum flat; one pair of dorsocentral bristles situated close behind the transverse suture. The apical half of the wing has a dark stellate mark (fig. 196, b), and the apical hyaline spot in cell r4+5 is at least as long as wide (fig. 196, c). This starlike mark is supplemented by an extensive yellowish or lighter brown area with a reticulate pattern in the basal half of the wing. The wing of Euarestoides most closely resem¬ bles those of the majority of species of Trupanea, but the basal half of the wing, in contrast to that genus, is almost completely filled with a reticulate pattern and is further distinguished from Trupanea by the presence of two pairs of scutellar bristles. This genus also resembles Neotephritis in having a reticulate wing pattern, three recognition.

pairs of frontal bristles, and two pairs of scutellar bristles, but the difference in wing patterns distinguishes it adequately. The most recent revision of Euarestoides (Foote 1958) provided additional detail concerning three of the four species discussed herein. Euarestoides probably extends south from the United States through Mexico, Central, and South America, but numerous specimens we have seen suggest that it may be one of a complex of closely related genera occurring south of the United States that requires further study. A Mexican species, dreisbachi, was described by Foote (1958). discussion.

Figures 196-199. Right wings, Euarestoides spp. 196, acutangulus (Thoms.); 197, abstersus (Lw.); 198, arnaudi Foote; 199, flavus (Adams). Figures 200, 201. Dorsal view, terminal abdominal segments and oviscape, Euarestoides spp. 200, acutangulus (Thoms.); 201, arnaudi Foote.

Systematic Treatment of the Genera

Key to U.S. and Canadian Species of Euarestoides 1. Brown color of wing pattern of uniform intensity throughout, the pattern in proximal half fainter than in distal half but not distinctly yellow (fig. 196); oviscape yellow with blackened base and apex (fig. 200); western Canada and U.S.acutangulus (Thomson) Pattern in proximal half of wing distinctly yellow in contrast to the dark brown of distal half, the line of demarcation rather abrupt (figs. 197— 199); oviscape entirely yellow, brown or black, not bicolored.2 2. Dark area in distal half of cell r1 with a small hyaline spot close to its center (fig. 197, b); eastern North America.abstersus (Loew) This dark area with the hyaline spot distinctly closer to the termination of vein R2+3 than to its center (figs. 198, a; 199, b); western and south¬ western U.S.3 3. Oviscape dark brown to black, 1.5-2.0 times as long as last 2 abdominal tergites (fig. 201); wing pattern as in fig. 198.arnaudi Foote Oviscape evenly yellow, about 2.0 times as long as last 2 abdominal tergites; wing pattern as in fig. 199 .flavus (Adams)

Euarestoides abstersus (Loew) (Fig. 197; Map 17) Trypeta abstersa Loew 1862a: 221 (female holotype, NMW; “Americae septentrionalis”). - Loew 1864: 91 (taxonomy). - Loew 1873: 322 (review).

•

Euarestoides abstersus

°

Euarestoides arnaudi

A

Euarestoides flavus

Map 17. Distribution of Euarestoides abstersus, E. arnaudi, and E. flavus.

157

Euarestoides

Urellia abstersa: Loew 1873: 324, 330 (taxonomy). - Johnson 1895b: 337 (Fla.). Coquillett 1899c: 265 (taxonomy, synonymy). — C. F. Adams 1904: 450 (in key). Johnson 1909: 113 (distribution). — Johnson 1910: 803 (N.J.). — Johnson 1913b: 84 (Fla.). Trypeta (Urellia) abstersa: Osten Sacken (part) 1878: 194 (in catalog). Euaresta abstersa: Coquillett 1899c: 265 (taxonomy). Trypanea abstersa: Johnson 1925b: 264 (N. Engl.). — Leonard 1928: 853 (N.Y.). — Johnson 1930: 151 (Nantucket). — Curran 1932b: 6 (in key). — Aczel (part) 1949: 292 (in catalog). - Flennig 1952: 212 (in catalog of immature stages). Tephritis abstersa: Cresson 1931: 3 (review). Trupanea (Euarestoides) abstersa: F. H. Benjamin (part) 1934: 58 (review, Fla.). Trupanea abstersa: Brimley 1938: 384 (N. Car.). — Ouellet 1941: 130 (Que.). — V. T. Phillips 1946: 83, 125 (description, biology larva; host). Euarestoides abstersa: Quisenberry 1951: 59 (taxonomy). — Foote 1958: 290 (revi¬ sion). Euarestoides abstersus: Foote 1965a: 669 (in catalog). — Wasbauer 1972: 113 (hosts). — Hilgendorf and Goeden 1982: 152 (host). Some of the older literature pertaining to abstersus is confused with that of acutangulus because these two species were long regarded as conspecific. The informa¬ tion presented in those references is too scanty for the identification of the actual species involved, thus we have not been able to study the material they represent. They are as follows: Loew 1873: 323, Observation 1 (Trypeta abstersa: we have not seen any Cuban species assignable to Euarestoides, and the identification here must remain tenta¬ tive). Wulp 1900: 426 (Urellia abstersa: the Mexico City record is probably a different species). Aldrich 1905: 611, 613 (Urellia abstersa, Tephritis acutangula: New Jersey and Key West records probably abstersus, the remainder acutangulus). Flendel 1914: 81 (Trypanea abstersa: the description fits neither abstersus nor acu¬ tangulus). V. T. Phillips 1923: 148, fig. 60 (Trypanea abstersa: the New York specimens are probably abstersus, the remainder acutangulus, as is the wing figure). Curran 1932b: 6 (Trypanea abstersa: the source of this information is not given). Aczel 1949: 292, 293 (Euarestoides abstersa, E. acutangula: abstersa is a mixed entry and acutangula a misidentification). Aczel 1951: 307 (Euarestoides abstersa: this is probably an incorrect entry referring to some other species). recognition. The wing pattern of abstersus very closely resembles that of acu¬ tangulus, but in contrast to the latter, the light markings in the basal half of the wing are definitely yellowed, and the hyaline spots therein are smaller and not as heavily coalesced, leaving a reticulate rather than striped pattern. The hyaline spot in the

Systematic Treatment of the Genera apex of cell r4+5 is less elongated than in acutangulus, and the hyaline spot close to the apex of cell rx is more centrally located in the surrounding dark area. E. abstersus resembles arnaudi and flavus in that the dark ray connecting the pterostigma and vein r-m (fig. 197, a) in these three species is narrower and much more discrete at its posterior termination than in acutangulus (fig. 196, a), but abstersus is distinguished from the two western species by its distribution, by the position of the hyaline spot near the apex of cell r,, and by the absence of a distinct whitish hyaline spot of varying size and shape at the base of cell r4+5 (fig. 198, b). distribution.

Map 17. The exact western limits of the distribution of abstersus

are not known. host.

Not

known.

E. abstersus is widely distributed throughout the Bahamas. We have seen abstersus-Yike flies from Mexico, but they may represent a different species more closely related to arnaudi or flavus (or both). discussion.

Euarestoides acutangulus (Thomson) (Figs. 105, 196, 200; Map 18) Trypeta acutangulus Thomson 1869: 583 (male holotype, NRS; Calif.). - Coquillett 1899c: 265 (synonymy). Tephritis acutangula: Loew 1873: 335 (taxonomy). - Woodworth 1913: 137 (Calif.). - Cresson 1931: 4 (review). - Knowlton and Harmston 1937: 145 (Utah). Trypeta (Tephritis) acutangula: Osten Sacken 1878: 194 (in catalog). Trypeta abstersa Loew [misidentification]: Cockerell 1898: 155 (N. Mex.). Cockerell 1902: 347 (N. Mex.). Urellia abstersa: [misidentification]: Doane 1899: 192 (distribution). - F. H. Snow 1903: 219 (Kans.). - Snow 1904: 345 (Ariz.). - Tucker 1907: 105 (Colo.). Aldrich 1913: 218 (Great Basin). Euaresta abstersa [misidentification]: Johnson 1903b: 106 (N. Mex.). - Cresson 1907: 106 (N. Mex., Tex.). — Cole 1912: 158 (Calif.). — Woodworth 1913: 137 (Calif.). Tephritis abstersa [misidentification]: Janes and Thomas 1932: 103 (Utah). Trupanea acutangula: Schwitzgebel and Wilbur 1943: 5 (biology). Euarestoides acutangula: Quisenberry 1951: 59 (taxonomy). - Foote 1958: 291 (revision). - Foote and Blanc 1963: 23 (review, Calif.). - Lipp and Schulz 1970: 27 (damage). Euarestoides acutangulus: Foote 1965a: 669 (in catalog). - Harris and Piper 1970: 134 (host). - Wasbauer 1972: 114 (hosts). - Goeden and Ricker 1974a: 831 (hosts). - Goeden and Ricker 1975: 304 (hosts). - Goeden and Ricker 1976a: 49 (hosts). - Goeden and Ricker 1976b: 926, 927, 929 (hosts). - Goeden and Ricker 1976c: 1174 (hosts). - Piper 1976: 381 (biology). - Hilgendorf and Goeden 1982:

Euarestoides

• Euarestoides acutangulus ★ Euarestoides acutangulus (state record only)

Map 18. Distribution of Euarestoides acutangulus. 152 (host). - Goeden 1986: 326 (host). - Goeden and Ricker 1986a: 42 (host). W. A. Palmer 1987: 192 (hosts). See also the additional references listed under abstersus, which include some allusions to acutangulus as well. E. acutangulus is the only species in the genus whose wing pattern is wholly brown (fig. 196), in contrast to the three other U.S. and Canadian species in which the basal half of the wing is distinctly yellow in contrast to the bold, dark preapical starlike mark. It is also the only U.S. and Canadian species of Euarestoides in which the hyaline spots in the basal half of the wing are so large and coalesce so RECOGNITION.

Systematic Treatment of the Genera broadly that the dark areas appear as irregular bands rather than reticulation. In contrast to all other species, the proximal third and extreme apex of the oviscape are blackened dorsally, and the intervening space is yellow (fig. 200). distribution.

Map 18. E. acutangulus is by far the most widespread Eu-

arestoides species occurring in North America. The eastern limit of its range in the United States is not definitely known. It also occurs in Tamaulipas and Juarez in northern Mexico. hosts.

Eight plant genera (Ambrosia, Anaphalis, Arnica, Chrysanthemum, Gail-

lardia, Helenium, lva, and Xanthium) were listed as hosts by Wasbauer (1972). Goeden and Ricker (1974a) reared acutangulus from Ambrosia acanthicarpa, Goeden and Ricker (1976a) reared it from A. dumosa, and Goeden (1986d) reared it from flower heads of Ambrosia cbamissonis. Additional host information was pre¬ sented by several authors (indicated in the synonymy above).

Euarestoides arnaudi Foote (Figs. 198, 201; Map 17) Euarestoides arnaudi Foote 1958: 291 (female holotype, NMNH; Canada Road behind Belmont, San Mateo County, Calif.). - Foote 1962: 175 (in key). - Foote and Blanc 1963: 26 (review, Calif.). - Foote 1965a: 669 (in catalog). - Arnaud 1979: 330 (type data). - Goeden 1983: 399 (host). - Goeden 1986: 326 (host). E. arnaudi, abstersus, and flavus are distinguished from acu¬ tangulus by the narrow, discrete ray connecting the pterostigma with vein r-m (figs. recognition.

196, a; 197, a) and by the yellow coloration of the dark areas in the basal half of the wing. In contrast to abstersus, both arnaudi and flavus have a whitish hyaline spot of varying size and shape in the base of cell r4+5 (fig. 198, b), the hyaline spot near the apex of cell

(figs. 197, b; 198, a) is not as centered in its surrounding dark area,

and the wide yellow band from the pterostigma to the apex of cell dm is unbroken in its anterior half. The only distinct character separating arnaudi from flavus is the shining black oviscape in the former species; it is entirely yellow in flavus. distribution.

Map 17. E. arnaudi does not occur south of the United States.

Goeden (1983, 1986) reared arnaudi from Baccbaris glutinosa and from B. pilularis ssp. consanguinea. hosts.

discussion.

See the discussion of flavus.

Euarestoides flavus (Adams) (Fig. 199; Map 17) Urellia flava Adams 1904: 450, 451 (female holotype, UKF; Bill Williams Fork, Ariz.). - F. H. Snow 1904: 345 (Ariz.). - Byers et al. 1962: 181 (type data).

161

Euleia Euarestoides flava: Foote 1962: 175 (type data, in key). — Foote and Blanc 1963: 26 (review, Calif.). Euarestoides flavus: Foote 1965a: 669 (in catalog). — Wasbauer 1973: 114 (host). Like abstersus and arnaudi, flavus is distinguished from acutangulus by the narrow dark discrete ray connecting the pterostigma with vein r-m and by the yellow coloration of the darker areas in the basal half of the wing. In contrast to abstersus, both flavus and arnaudi have a whitish hyaline spot of varying size and shape in the base of cell r4+5 (fig. 198, b), the hyaline spot near the apex of cell rx is not as well centered in the surrounding dark area, and the wide yellow band from the pterostigma to the apex of cell dm is unbroken in its anterior half (fig. 199, a). E. flavus has a distinctly and entirely yellow oviscape; that of arnaudi is entirely recognition.

shining black dorsally (see discussion below). distribution.

Map 17. E. flavus is not known to occur south of the United

States. host.

The only known host is Baccbaris viminea, as indicated by Wasbauer

(1972). We have examined a large series of “ arnaudi-flavus” reared from species of Baccbaris in southern California by R. D. Goeden. Our exhaustive exam¬ ination of this material failed to reveal any morphological differences between these two species other than the color of the oviscape. Therefore the status of arnaudi as a species distinct from flavus should be considered tentative until further evidence is discussion.

made available.

Genus Euleia Walker Euleia Walker 1835: 81 (type species, Musca onopordinis Fabricius 1775: 787, by monotypy) (= beraclei (Linnaeus)). - Hering 1940c: 16 (taxonomy). - Foote 1959b: 145 (review). - Foote and Blanc 1963: 27 (review, Calif.). - Foote 1965a: 677 (in catalog). - Sabrosky 1971: 85, 87 (correction, N.A. catalog). - Wasbauer 1972: 114 (hosts). - Foote 1984: 88 (in catalog). Acidia Robineau-Desvoidy [misidentification]): Loew (part) 1873: 235 (taxonomy). - Aldrich (part) 1905: 603 (in catalog). - Malloch 1923: 216 (taxonomy). - V. T. Phillips (part) 1923: 121, 128 (in key; review, e. N.A.). - Johnson 1925b: 261 (N. Engl.). - Curran 1932b: 3 (in key). Trypeta (Acidia): Osten Sacken (part) 1878: 189 (in catalog). Pbilopbylla Rondani [misidentification]: Hendel 1914b: 89 (in key, world). - Hendel 1927: 97 (revision, Palearctic Region). Trypeta Meigen [misidentification]: Curran 1932b: 3 (in key). - Curran 1934: 287, 289 (in key).

Systematic Treatment of the Genera Myoleja Rondani [misidentification]: Bates 1933a: 50 (synonymy). - Curran 1934: 289 (in key). recognition.

The species of Euleia superficially resemble those of Strauzia, Ep-

ochra, Trypeta, and the yellow-bodied Rhagoletis species. Euleia is distinguishable from Strauzia by the presence of two pairs of orbital bristles; from Epochra by the presence of presutural bristles, two pairs of orbitals, and the relatively short anten¬ nae; from Trypeta by the presence of a distinct rounded spot in the distal half of cell dm and the more distal position of vein r-m along cell dm; and from Rhagoletis by the apically rounded 1st flagellomere. Specimens of Euleia and Trypeta frequently have tiny dark spots on the integument, especially on the posterior half of the scutum and on the scutellum. This character never occurs in the other three genera. discussion.

The genus Euleia is essentially Holarctic, having eight Palearctic

species in addition to the two discussed here. The type species, heraclei (L.), is the celery fly of Europe, the Near East, and North Africa. Both heraclei and the North American fratria are economically important species throughout their respective ranges. The species of Euleia whose habits are known, including both North Ameri¬ can species, are leaf miners, an unusual habit among tephritids. Among the New World genera of Tephritidae, Euleia is probably most closely related to Trypeta, the species of which also are leaf miners.

Key to U.S. and Canadian Species of Euleia Wing with 2 hyaline incisions (fig. 202, a) separated by an uninterrupted dark band in posterior distal quarter of wing; gena about Vs or less as high as eye (fig. 204, a).fratria (Loew) Wing with a single wide hyaline area present in distal posterior quarter of wing, without a dark band dividing it completely into 2 parts (fig. 203, a) gena at least V4 as high as eye (fig. 205, a) .uncinata (Coquillett)

Euleia fratria (Loew) Parsnip leaf miner (Figs. 65, 202, 204; Map 19) Trypeta fratria Loew 1862c: 67 (female holotype, MCZ; U.S.). - Loew, 1873: 234 (review). - Coquillett 1895: 381 (biology). - Curran 1934: 238 (figure of wing). Trypeta liogaster Thomson 1869: 578 (holotype male, NRS; Calif.). - Loew 1873: 337, 340 (synonymy, description). - Doane 1899: 178 (synonymy). - Aldrich 1905: 603 (in catalog). Acidia fratria: Loew 1873: 235, 329 (taxonomy). - Doane 1899: 178 (taxonomy). Johnson 1900b: 687 (N.J.). - Aldrich 1905: 603 (in catalog). - Chittenden 1909: 9 (biology, distribution). - Johnson 1910: 801 (N.J.). - Banks 1912: 31 (descrip¬ tion, larva). - Woodworth 1913: 135 (Calif.). - Winn and Beaulieu 1915: 153

163

Euleia

Figures 202, 203. Right wings, Euleia spp. 202, fratria (Lw.).; 203, uncinata (Coq.). Figures 204, 205. Lateral view, head, Euleia spp. 204, fratria (Lw.); 205, uncinata (Coq.).

(Que.). - Britton 1920: 213 (Conn.). - V. T. Phillips 1923: 129 (review).-Johnson 1925a: 97 (Harris collection). - Johnson 1925b: 261 (N. Engl.). - Leonard 1928: 851 (N.Y.). - Petch and Maltaise 1932: 56 (Que.). - Essig 1938: 602 (review, w. N.A.) - V. T. Phillips 1946: 28, 105 (biology; description larva). - Forsell 1947: 167 (Puget Sound). - Essig 1958: 602 (review, w. N.A.). Trypeta (Acidia) fratria: Osten Sacken 1878: 189 (in catalog). Acidia frateria [error]: Frost 1924: 32 (synonymy). Acidia heraclei (misidentification): Frost 1924: 32 (hosts; description leaf mine; distribution). - Needham et al. 1946: 345 (note, biology). Alcidia [error] fratria: Woodworth 1913: 137 (Calif.). Philophylla fratria: Hendel 1927: 99 (taxonomy). - Hendel 1935: 53 (taxonomy). Hennig 1952: 219 (in catalog, immature stages). Euleia fratria: Foote 1959b: 146 (review). - Foote and Blanc 1963: 27 (review, Calif.) - Foote 1965a: 677 (in catalog). - Tauber and Toschi 1965a: 369 (biology, reproductive behavior). - Bateman 1972: 497 (ecology). - Steyskal 1972d: 130 (taxonomy). — Wasbauer 1972: 114 (hosts). recognition. E. fratria is distinguished from uncinata in having, in profile, a less convex face and a slightly less acute 1st flagellomere apically. The gena (fig. 204, a) of fratria is at most only one-fifth as high as the eye, whereas that of uncinata is at least one-fourth as high. The most distinctive differences between the two species are in the wing pattern. In fratria, two hyaline areas immediately behind the dark apical

164

Systematic Treatment of the Genera

Map 19. Distribution of Euleia fratria and E. uncinata.

costal band extend to the wing margin and are separated by an unbroken dark band (fig. 202, a); in uncinata (fig. 203, a), a single hyaline area reaches the posterior margin of the wing proximal to the dark mark on vein M, which is fused with the apical costal band at the wing apex. DISTRIBUTION.

Map 19.

E. fratria was reported as infesting parsnip in the United States as early as 1891, but reports of damage to that plant have been sporadic ever since. Five or six other plants in the families Apiaceae and Asteraceae are known hosts. hosts.

165

Eurosta

Euleia uncinata (Coquillett) (Figs. 203, 205; Map 19) Acidia uncinata Coquillett 1899c: 260 (female holotype, NMNH; Ft. Wrangel, St.Paul’s I., Alaska). — Aldrich 1905: 603 (in catalog). — Cole 1921: 175 (descrip¬ tion male, type data). — Malloch 1923: 217 (description pupa, adult; biology). — Hennig 1952: 219 (in catalog, immature stages). — Arnaud 1979: 329 (type data). Euleia uncinata: Foote 1959b: 149 (review). - Foote 1965a: 677 (in catalog). recognition. The characters that distinguish uncinata from fratria will be found in the Recognition section under the latter species (see figs. 202—205). DISTRIBUTION. Map 19. host. Not known. discussion. In contrast to fratria, uncinata is of no known economic impor¬ tance.

Genus Eurosta Loew Eurosta Loew 1873: 280 (type species, Acinia solidaginis Fitch 1855a: 771, by designation of Coquillett 1910: 543). - Williston 1896: 122 (in key). - Coquillett 1899c: 268 (in key). - Aldrich (part) 1905: 609 (in catalog). - Williston 1908: 287 (in key). - Hendel 1914b: 94 (in key). - V. T. Phillips 1923: 122, 142 (in key, review). - F. FI. Benjamin 1934: 26 (review, Fla.). - Curran 1932b: 2, 4 (in key, taxonomy). - Curran 1934: 293 (in key). - Foote 1965a: 662 (in catalog). Wasbauer 1972: 114 (hosts). - Steyskal and Foote 1977: 151 (key, known species). - Novak and Foote 1980: 205 (biology, immatures). - Ming 1989: 1 (revision). Trypeta {Eurosta): Osten Sacken 1878: 192 (in catalog). Durosta [error]: M. T. Cook 1907: 1 (description of galls). Eurostina Curran 1932b: 4 (type species, Trypeta latifrons Loew 1862c: 89, by original designation). - F. H. Benjamin 1934: 36 (taxonomy, synonomy). - Curran 1934: 293 (in key). - Foote 1961: 28 (synonymy). - Arnaud and Owen 1981: 149 (type data). Members of Eurosta are characteristically dark, stout-bodied flies of medium size; older specimens usually have a distinctive greasy appearance, mak¬ ing color patterns and setation difficult to study. The wing is quite dark with light or hyaline spots that sometimes coalesce to form larger light areas (figs. 207, 208, 213, 214). The frons is very broad and setulose, there are one or two pairs of reclinate orbital bristles, two to four pairs of frontal bristles, all the postoculars are whitish, and the first flagellomere is very short. The dorsocentral bristles are located closer to recognition.

Systematic Treatment of the Genera the supra-alars than to the transverse suture, and the apical scutellar bristles are usually absent or, if present, much shorter than the basal pair. Extra (supernumer¬ ary) bristles sometimes occur at or near the location of normal body bristles in Eurosta, but this condition is not of any known taxonomic value. It is difficult to confuse Eurosta with any other North American fruit fly genus; the characters given in the key distinguish it adequately. discussion.

All members of Eurosta feed inside cavities they form within the

roots, crowns, or stems of goldenrod, usually causing the formation of distinct, often strongly swollen galls. The early literature is replete with descriptions of these galls. Stoltzfus (1989) reported the presence in Solidago canadensis of occasional individu¬ als of solidaginis, the larvae of which feed inside stem cavities but do not cause the formation of galls. Until recently, the only taxonomic treatment of the genus was presented by Steyskal and Foote (1977) in the form of a key and the description of one new species. Ming (1989) revised Eurosta formally and included a cladistic study of relationships within and outside the genus. We are deeply indebted to her for making that revision available to us for inclusion in this handbook.

Key to U.S. and Canadian Species of Eurosta (Adapted from Ming 1989) 1. Scutum with patches of darker, brown to black setulae among the lightcolored setulae; wing apex usually with radiate pattern or at least large hyaline spots (figs. 206-208) .2 Scutal setulae unicolorous, golden to whitish; wing pattern variable. .3 2. One pair scutellar bristles; cell be almost completely hyaline (fig. 206, a); light area in cell cua1 scarcely extending into cell dm (fig. 206, c); dorsal surface of abdomen with distinct, broad, light-colored median stripe .. .latifrons (Loew) Two pairs scutellar bristles; cell be entirely infuscated or with small basal hyaline spot (figs. 207, a; 208, a); light area in cell cuaj often extending into cell dm or br (figs. 207, c; 208, b); dorsal surface of abdomen often patchy brown and gray, often a narrow, light-colored median stripe pre¬ sent .cribrata (Wulp) 3. Apex of wing with narrow, crescentic hyaline mark (fig. 209, c), or if broken into several small spots by darkening at middle of cell r4 + 5 or at apices of veins R4+5 or M, spots are extremely small (fig. 211) and/or wider than long and broadly touching wing margin .4 Apex of wing with large, semicircular hyaline or reticulate area or with several large, rounded hyaline spots often narrowly touching, or isolated from, wing margin (figs. 213-215).6 4. Large, distinct spot present at apex of vein At + CuA2, longer than wide (fig. 209, b), usually devoid of any dark markings, margins sharply de¬ fined .fenestrata Snow

Figures 206-215. Right wings, Eurosta spp. 206, latifrons (Lw.); 207, cribrata (Wulp); 208, cribrata (Wulp), variant; 209, fenestrata Snow; 210, floridensis Foote; 211, comma (Wied.); 212, comma (Wied.), variant; 213, solidaginis solidaginis (Fitch); 214, solidaginis fascipennis Curr.; 215, lateralis (Wied.).

167

Systematic Treatment of the Genera

This distinct spot present (fig. 210, c) or reduced (fig. 212, c); a lighter, variable-sized area in anal lobe composed of numerous small pale spots .5 5. Light area at apex of vein A^ + CuA2 conspicuous, extending over half the width of cell cua3 (fig. 210, c); hyaline spot at apex of pterostigma an inverted triangle divided by a median bar (fig. 210, a). .floridensis Foote Light area at apex of vein Aj + CuA1 poorly developed (fig. 211, c); if evident, at most extending only V4 distance across cell cuaj (fig. 212, c); hyaline spot at apex of pterostigma variable (figs. 211, a; 212, a) . .comma (Wiedemann) 6. Wing with extensive hyaline regions: at apex of pterostigma, at apex of wing, and at apices of cells m and cuaj (figs. 213, 214); hyaline area in cell cua: triangular, broadly touching posterior wing margin and extend¬ ing into cell dm or beyond; usually 2 pairs of orbital bristles . .solidaginis (Fitch) a. Hyaline mark at apex of pterostigma confined to cell r2 + 3 or de¬ scending posteriorly (fig. 213, a), but distinctly separated from hyaline area in cell m (fig. 213, c). Eastern U.S., extending west¬ ward .solidaginis solidaginis (Fitch) b. Hyaline mark at apex of pterostigma descending into cell m and fully connected with hyaline area there (fig. 214, a, c), or con¬ nected by 2-4 hyaline spots to form a distinct transverse band. Western U.S., extending eastward . .solidaginis fascipennis Curran Dark color of wing extensive and hyaline areas reduced; hyaline spot or spots in cell cua3 narrowly ovate, attaining posterior wing margin nar¬ rowly or not at all (fig. 215 c); wing apex with 3-6 small, widely sepa¬ rated hyaline spots (fig. 215, b); usually only 1 pair orbital bristles . .. .lateralis (Wiedemann)

Eurosta comma (Wiedemann) (Figs. 211, 212; Map 20) Trypeta comma Wiedemann 1830: 478 (female lectotype, here designated; NMW; Ky.). - Osten Sacken 1858: 79 (in catalog). - Loew 1862c: 93 (review). - Loew 1873: 280, 336 (review). Trypeta alvea Walker 1849: 1027 (male holotype, BMNH; U.S.). - Hardy 1959: 208, 234 (synonymy). - Foote 1964d: 318 (synonymy, taxonomy). Trypeta dertona Walker 1849: 1028 (female holotype, BMNH; U.S.). - Hardy 1959: 213, 234 (synonymy). - Foote 1964d: 320 (synonymy). Eurosta comma: Loew 1873: 280, 329 (taxonomy). - W. A. Snow 1894: 169 (tax¬ onomy). - Doane 1899: 186 (taxonomy). - Johnson 1900b: 688 (N.J.). - Aldrich 1905: 609 (in catalog). - Crevecour 1906: 95 (Kans.). - Daecke 1910: 342 (biolo¬

gy)- — Johnson 1910: 802 (N.J.). — Stebbins 1910: 50 (biology, description of gall). - Felt 1918: 198 (description of galls). - Britton 1920: 204 (Conn.). - V. T. Phillips 1923: 144 (review). - Johnson 1925b: 263 (N. Engl.). - Sturtevant 1925: 215 (seminal receptacles). - Cole 1927: 445 (male genitalia). - Leonard 1928: 852

169

Eurosta

(N.Y.). - Hendrickson 1930: 143 (Iowa). - Curran 1934: 292 (figure of head). Brimley 1938: 384 (N. Car.). — Strickland 1938: 204 (Alta.). - Felt 1940: 322 (description of gall). - V. T. Phillips 1946: 45, 114 (description, biology larva; hosts). - Hennig 1952: 211 (in catalog of immature stages). - Strickland 1953: 277 (ptilinal armature). - Foote 1965a: 663 (in catalog). - Wasbauer 1972: 114 (hosts). - Steyskal and Foote 1977: 149, 151 (in key, taxonomy, hosts). - Novak and Foote 1980: 207 (in key to larvae). - Ming 1989: 29 (revision). Eurosta elsa Daecke 1910: 342 (female lectotype, NMNH; Richmond Hill, Long I., N.Y.). - Felt 1918: 198 (description of gall). - Britton 1920: 204 (Conn.). - V. T. Phillips 1923: 144 (review). - Johnson 1925b: 264 (N. Engl.). - Curran 1930: 77 (N.Y.). - Felt 1940: 322 (description of gall). - V. T. Phillips 1946: 114 (hosts). Foote 1965a: 663 (in catalog). - Novak et al. 1967: 147 (hosts, gall). - Wasbauer 1972: 115 (hosts). - Steyskal and Foote 1977: 149, 151 (taxonomy, lectotype designation, host, in key). - Novak and Foote 1980: 207, 211 (biology; hosts; description of larva, pupa; in key to larvae). - Ming 1989: 29 (synonymy). E. comma shares with fenestrata and floridensis a wing pattern characterized by the presence of a narrow apical hyaline crescent (as in fig. 209, c), which is sometimes broken into small hyaline spots by the presence of dark markings at the ends of the radial veins (fig. 210, b). In all other species of Eurosta, the hyaline spots at the wing apex project deeply into the wing disk (fig. 206, b). These three species are also characterized by the lack of a well-developed hyaline area in the posterior margin of cell m. In comma, The hyaline area at the apex of vein Aj + recognition.

170

Systematic Treatment of the Genera

CuA2 in the wing pattern is usually small (figs. 211, c; 212, c) compared with those of fenestrata and floridensis, although this character intergrades among these spe¬ cies. The hyaline spot at the apex of the pterostigma in comma is usually wide at the wing margin and is divided by a narrow dark mark, but may be reduced (compare figs. 211, a and 212, a). distribution.

Map 20.

E. comma has been reported from Solidago juncea and S. rugosa. It also has been found associated with Aster lateriflorus, questionably as a true host. hosts.

E. comma, fenestrata, and floridensis form a species complex in which some individuals or series may be difficult to identify. Further careful study of discussion.

host plants and perhaps genetic or biochemical analyses are needed to define the species of this complex more precisely. Ming (1989) concluded that elsa is a synonym of comma based primarily on the extensive variation she observed in the hyaline area at the apex of the pterostigma. The adult and immature stages were described in detail by Ming (1989); that author and Novak and Foote (1980) provided biological information. The lectotype female of comma is accompanied by labels with “kentucky” and “comma Wied.” in Wiedemann’s handwriting, a minute pink square, a white square with “52,” a label with “comma det. Low” written by a later NMW worker, and one with “Coll. Winthem.” Although it was described from an unstated number of females in Wiedemann’s collection, we consider this a valid syntype because of the Wiedemann label. Two specimens from the old NMW general collection are possibly from the same original series as the lectotype but are doubtful syntypes.

Euros ta cribrata (Wulp) (Figs. 69, 207, 208; Map 21) Trypeta cribrata Wulp 1867: 125 (male and female syntypes, RNHL; Wis.). — Foote 1965a: 663 (in catalog). — Ming 1989: 45 (synonymy). Eurosta reticulata Snow 1894: 170 (male lectotype, UKL; Mont.). — Coquillett 1899c: 264 (synonymy). - Doane 1899: 186 (distribution). - Doane 1900: 47 (taxonomy). — Aldrich 1905: 609 (in catalog). — Crevecoeur 1906: 95 (Kans.). — Thompson 1907: 71 (description of gall). - Stebbins 1910: 52 (biology, descrip¬ tion of gall). - Felt 1918: 197 (description of gall). - Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 144 (review). - Johnson 1925b: 263 (N. Engl.). - F. H. Benjamin 1934: 28 (review, Fla.). — Felt 1940: 322 (description of gall). — V. T. Phillips 1946: 114 (hosts). — Flennig 1952: 211 (in catalog immature stages). — Byers et al. 1962: 180 (type data). — Foote 1962: 177 (type designation). — Foote 1965a: 663 (in catalog). - Novak et al. 1967: 147 (hosts, gall). - Wasbauer 1972: 115 (hosts). - Steyskal and Foote 1977: 152 (in key). - Novak and Foote 1980: 207, 211, 217 (biology, description of immature stages, in key to larvae, hosts). Ming 1989: 45 (synonymy).

171

Eurosta

Eurosta latifrons (state record only)

Map 21. Distribution of Eurosta cribrata, E. fenestrata, and E. latifrons.

Eurosta conspurcata Doane 1899: 186 (male lectotype, WSU; Pullman, Wash.). Coquillett 1899c: 264 (synonymy). - Doane 1900: 47 (taxonomy). - Johnson 1900b: 688 (N.J.). - Aldrich 1905: 609 (in catalog).-Johnson 1910: 802 (N.J.). Britton 1920: 204 (Conn.). - V. T. Phillips 1923: 144 (review). - Johnson 1925b: 263 (N. Engl.).-Leonard 1928: 852 (N.Y.). - Benjamin 1934: 28 (nomenclature). - Brimley 1938: 384 (N. Car.). - Foote 1965a: 663 (in catalog). - Foote 1966b: 123 (type designation). - Steyskal and Foote 1977: 152 (in key). - Zack 1984: 32 (type data). - Ming 1989: 45 (synonymy). Eurosta cribrata: Ming 1989: 45 (revision).

172

Systematic Treatment of the Genera

recognition.

E. cribrata and latifrons are the only species of Eurosta with

mixed light and dark patches of scutal setulae. Their wing patterns are similar in having a pair of comparatively large hyaline spots apicad of the pterostigma and distinct, rounded hyaline spots at the wing apex (fig. 206, b). Cell be of latifrons (fig. 206, a) is almost entirely hyaline, whereas that of cribrata (figs. 207, a; 208, a) has a small hyaline spot or is completely dark. This species is the only Eurosta which consistently has two pairs of scutellar bristles; occasional specimens of floridensis and solidaginis also have two pairs. DISTRIBUTION.

Map 21.

E. cribrata has been reared from Solidago sempervirens in Florida and from S. juncea in Ohio (Novak and Foote 1980). It was collected on Solidago hosts.

missouriensis by Ming (1989). discussion.

We follow Ming (1989) in considering conspurcata and reticulata as

synonyms of cribrata, primarily on the basis of variations in wing pattern.

Eurosta fenestrata Snow (Fig. 209; Map 21) Eurosta fenestrata Snow 1894: 169 (female holotype, UKL; Ariz.). — Aldrich 1905: 609 (in catalog). — Johnson 1909: 113 (distribution). - Johnson 1913b: 84 (Fla.). — Byers et al. 1962: 180 (type data). - Foote 1962: 174 (type data). — Foote 1965a: 663 (in catalog). — Steyskal and Foote 1977: 151 (in key). — Ming 1989: 54 (revision). recognition.

The wing patterns of comma and floridensis resemble that of

fenestrata in that a narrow hyaline crescent, sometimes broken into small hyaline spots, occupies the apex of the wing (fig. 209, c). E. fenestrata and floridensis may be distinguished from comma by the extensive development of the hyaline area at the apex of vein Aj + CuA2 (figs. 209, b; 210, c). This area in fenestrata is usually longer than wide, and the margins are relatively distinct and in sharp contrast to the dark area surrounding it. The comparable area in floridensis is more or less quadrangular, the margins are somewhat diffuse, and irregular dark markings often occupy its central portion. The anal lobe of fenestrata is largely dark with marginal light spots (fig. 209, a), whereas that of floridensis is usually more extensively hyaline with dark marginal spots (fig. 210, d). DISTRIBUTION.

host.

Map 21.

Not known. Ming (1989) suggested that Solidago is probably the host

genus for fenestrata, based largely on the host relationships of the other members of this species group. discussion.

See

discussion of

comma.

173

Eurosta

Eurosta floridensis Foote (Fig. 210; Map 20) Eurosta floridensis Foote in Steyskal and Foote 1977: 148 (female holotype, NMNH; Fla.). — Novak and Foote 1980: 211 (host). — Ming 1989: 60 (revision). Eurosta comma: F. H. Benjamin 1934: 28 (host). E. floridensis shares with comma and fenestrata the distinctive narrow hyaline area in the apex of the wing, in floridensis often broken into small hyaline spots by the darkening at the tips of veins R4+5 and M or in the middle of cell recognition.

r4+5 (fig- 210, b). Both floridensis and fenestrata have a well-developed hyaline area at the apex of vein + CuA2 and thereby are distinguished from comma. In flor¬ idensis, this hyaline area is more or less quadrangular, somewhat darkened centrally, and its margins are diffuse (fig. 210, c), whereas in fenestrata (fig. 209, b), this area is longer than wide and has sharp edges that contrast with the surrounding dark area. These two species may be further distinguished by the markings in the anal lobe; in floridensis, that cell is usually largely hyaline with marginal dark spots (fig. 210, d). distribution.

host.

Map 20.

E. floridensis has been reared in large numbers from Solidago fistulosa, the

only known host. discussion.

Florida records of comma represent this species. See also the discus¬

sion of comma.

Eurosta lateralis (Wiedemann) (Fig. 215; Map 20) Trypeta lateralis Wiedemann 1830: 479 (male syntypes, NMW; probably Fla.). Hardy 1968: 146 (type data, synonymy). Trypeta donysa Walker 1849: 1007 (male holotype, BMNH; probably Fla.). - Foote 1964a: 61 (description, synonymy). - Foote 1964d: 321 (synonymy). - Hardy 1968: 146 (synonymy). Eurosta nicholsoni Benjamin 1934: 27 (male holotype, originally NMNH, lost; 1.25 mi s. Titusville, Fla.). - V. T. Phillips 1946: 114 (hosts). - Hennig 1952: 211 (in catalog immature stages). — Foote 1964a: 61 (synonymy). — Foote 1964d: 321 (synonymy). - Foote 1965a: 663 (in catalog). - Hardy 1968: 146 (synonymy). Eurosta donysa: Wasbauer 1972: 115 (host). Eurosta lateralis: Steyskal and Foote 1977: 152 (in key). - Ming 1989: 64 (revision). E. lateralis may be distinguished from all other Eurosta by the (usually) single pair of orbital bristles, by the extensively dark wing, and by the nature of the hyaline area in cell cua1? which is distinct and, in contrast to those of any other species, rarely attains the posterior border of the wing (fig. 215, c). The dark area dividing the two hyaline spots immediately apicad of the pterostigma is recognition.

174

Systematic Treatment of the Genera

wider than in most other species (fig. 215, a), and the hyaline spots at the wing apex are irregular in shape and do not project quite as deeply into the wing as in many other species (fig. 215, b). distribution.

HOST.

Map 20.

Benjamin (1934) indicated lateralis (as nicholsoni) occurs in small round

stem galls of a Solidago sp. in Florida. discussion.

Ming (1989) described lateralis in detail, and Benjamin (1934) pre¬

sented a brief description of the adult and larva of this species under the name nicholsoni.

Eurosta latifrons (Loew) (Fig. 206; Map 21) Trypeta latifrons Loew 1862c: 89 (female holotype, MCZ; “Carolina”). — Loew 1873: 283 (review). Eurosta latifrons: Loew 1873: 283, 329 (taxonomy). — W. A. Snow 1894: 171 (taxonomy). — Aldrich 1905: 609 (in catalog). — Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 144 (review). — Johnson 1925b: 263 (N. Engl.). — Johnson 1927: 218 (Mt. Desert). - Brimley 1938: 384 (N. Car.). - Foote 1961: 28 (taxonomy). Foote 1965a: 663 (in catalog). — Steyskal and Foote 1977: 152 (in key). — Ming 1989: 70 (revision). Trypeta (Eurosta) latifrons: Osten Sacken 1878: 192 (in catalog). Eurostina latifrons: Curran 1932a: 2 (type data). Eurostina confusa Curran 1934: 293 [nomen nudum]. — F. FI. Benjamin 1934: 27 (synonymy). - Foote 1961: 28 (synonymy). — Arnaud and Owen 1981: 149 (type data). recognition.

E. latifrons resembles several other Eurosta species in having well-

developed hyaline spots immediately apicad of the pterostigma and at the wing apex (fig. 206). E. latifrons and cribrata differ from other Eurosta in having some black scutal setulae. The former may be distinguished from cribrata by the number of scutellar bristles and the almost entirely hyaline cell be (fig. 206, a). It is one of the few Eurosta species in which the marginal spots in the apices of cells r2+3 and r4+5 are of more or less equal size and equally spaced (fig. 206, b), and the bulla is larger and more prominent than in any other member of the genus. The light area at the apex of vein A1 -I- CuA2 is not well developed in this species; it consists of several partially coalesced hyaline spots (fig. 206, c). DISTRIBUTION.

host.

Not

Map 21.

known.

175

Euro st a

Eurosta solidaginis (Fitch) Goldenrod gall fly (Figs. 67, 213, 214; Map 22) Ortalis nuphera Say [nomen nudum]: T. W. Harris 1835a: 80. Tepbritis asteris Harris 1841: 417 (syntypes, sex not stated, MCZ; Mass.) (preoc¬ cupied Haliday 1838). — T. W. Harris 1842: 417 (taxonomy). — T. W. Harris 1852: 97 (biology). — B. D. Walsh 1866: 114 (taxonomy, synonymy). — Flint 1890: 620 (biology). — Aldrich 1905: 609 (in catalog). — F. H. Benjamin 1934: 27 (nomencla¬ ture). Acinia solidaginis Fitch 1855a: 771 (male lectotype, here designated, NMNH; N.Y.). — Fitch 1856: 67 (taxonomy). - Coquillett 1910: 543 (type species designation).

Map 22. Distribution of Eurosta solidaginis.

176

Systematic Treatment of the Genera

Trypeta asteris: Osten Sacken 1858: 79 (in catalog). — Loew 1862c: 58 (taxonomy). — Osten Sacken 1862: 99 (taxonomy). - Osten Sacken 1869: 299 (gall). — Loew 1873: 335 (synonymy).

Trypeta solidaginis: Osten Sacken 1858: 80 (in catalog). — Loew 1862c: 82 (review). — B. D. Walsh 1866: 114 (synonymy, host). - Loew 1873: 280 (review). Ashmead 1889: 636 (parasite). - Fyles 1894: 120 (description of gall, adult, parasite). - C. F. Baker 1895: 174 (description of gall). - Snyder 1898: 99 (emer¬ gence from gall). - M. T. Cook 1903: 422 (description of gall). - Field 1904: 309 (biology). - M. T. Cook 1905: 806, 844 (description of gall). - Jarvis 1907: 68 (description of gall). - M. T. Cook 1910: 32 (description of gall). - Girault 1915: 132 (description of adult, gall).

Eurosta asteris: Loew 1873: 99 (synonymy). - Johnson 1925a: 97 (Harris collec¬ tion). — Johnson 1927: 218 (Mt. Desert).

Eurosta solidaginis: Loew 1873: 280, 329 (taxonomy). - Osten Sacken 1880: 53 (host). — Keen 1885: 55 (Pa.). — Beutenmueller 1892: 274 (description of gall). Brodie 1892: 137 (description of gall). - W. A. Snow 1894: 169 (taxonomy). Harrington 1895: 197 (description of gall). — Johnson 1895b: 303 (Fla.). — Doane 1899: 186 (taxonomy).—Johnson 1900b: 688 (N.J.). — Chagnon 1901: 14 (Can.). — F. H. Snow 1903: 219 (Kans.). - Aldrich 1905: 609 (in catalog). - Washburn 1905: 118 (Minn.). - Thompson 1907: 71 (biology).—Johnson 1910: 703 (N.J.). — Stebbins 1910: 51 (description of gall). — Cosens 1912: 297 (description of gall). — Johannsen 1912: 9 (parasite). — Johnson 1913b: 84 (Fla.). — Hendel 1914b: 94 (type data). - Ping 1915: 161 (biology, description of gall). - Winn and Beaulieu 1915: 153 (Que.). - Viereck 1917: 238 (parasite). - Felt 1918: 197 (description of gall). — Britton 1920: 204 (Conn.). — Cole and Lovett 1921: 326 (Oreg.). — V. T. Phillips 1923: 144 (review). — Curran 1925: 128 (biology). — Johnson 1925a: 97 (Harris collection). — Johnson 1925b: 263 (N. Engl.). — Leonard 1928: 253 (N.Y.). — Gronemann 1930: 27 (description of gall). — W. T. Davis 1931: 120 (predators). — Curran 1932b: 2 (taxonomy). — Petch and Maltaise 1932: 57 (Que.). — Bates 1934c: 1 (morphology). - F. H. Benjamin 1934: 27 (synonymy). - Gahan 1934: 116 (parasite). - Hughes 1934: 119 (parasites, description of gall). - Houard 1935: 172 (description of gall). — Brimley 1938: 384 (N. Car.). — Essig 1938: 605 (note). - Felt 1940: 321 (description of gall). — Milne 1940: 101 (biology). — Carstenson and Jaques 1942: 525 (Iowa). — Burks 1943: 534 (parasite). — V. T. Phillips 1946: 47, 115 (description, biology larva; host). — Weigel and Dilks 1950: 134 (biology). — Krogstad 1951: 23 (biology, Minn.). — Uhler 1951: 1 (biology). — Hennig 1952: 211 (in catalog immature stages). — Judd 1953: 294 (gall, biology). — Peterson 1953: 331, 333, 335, 337 (figures of larva). — Strickland 1953: 277, 296 (ptilinal armature). — Essig 1958: 605 (taxonomy). - Miller 1959: 246 (re¬ view). — Uhler 1961: 215 (parasites, predators). — Winterringer 1961: 39, 40 (biology, description of galls). — Foote 1965a: 663 (in catalog). — Judd 1968: 78 (gall, parasite). — Capek 1971: 434 (description of gall). — Wasbauer 1972: 115 (hosts). — Cane and Kurczewski 1976: 976 (biology, parasites). — Steyskal and Foote 1977: 151 (in key). - Laing and Heraty 1978: 12 (biology). — Hartnett and Abrahamson 1979: 910 (plant-insect interactions). - Stinner and Abrahamson

177

Eurosta

1979: 918 (plant-insect interactions). — Novak and Foote 1980: 207, 211 (in key to larvae, hosts). — Gilbert and Kurczewski 1986: 29 (soil nutrients). — Vasey and Ritter 1987: 452 (morphology). — Walton 1988: 654 (galls). — Ming 1989: 77 (revision). — Stoltzfus 1989: 50 (nongail-forming larvae).

Trypeta (Eurosta) asteris: Osten Sacken 1878: 192 (in catalog). Durosta [error] solidaginis: M. T. Cook 1907: 1 (description of gall). Eurosta asteri [error]: Johnson 1930: 151 (Nantucket). recognition.

E. solidaginis is one of the most easily recognized species in the

genus in having three or four widely separated light areas in the wing. One of these is a prominent subtriangular hyaline spot at the apex of vein

+ CuA2, extending

into cell dm (fig. 213, d) and occasionally into cell br (fig. 214, d). A second area consists of a group of rounded hyaline spots at the wing apex (fig. 214, b), sometimes coalesced into a larger hyaline area. A third light area consists of a costal spot immediately apicad of the pterostigma. This light area descends at least into cell r2+3 and often into cell r4+5 (fig. 213, a), but in eastern specimens (subspecies solidaginis), it is distinctly separated from a group of rounded hyaline spots in the posterior margin of cell m (fig. 213, c), whereas in western specimens (subspecies fascipennis), the spots are most narrowly separated, forming a continuous or almost continuous hyaline band (fig. 214, a, c). No other species of Eurosta presents this combination of wing characters (see additional comments below). DISTRIBUTION.

Map 22. In this map, the distribution records of solidaginis soli¬

daginis and s. fascipennis have not been recorded separately. hosts.

Wasbauer (1972) indicated that the following species of Solidago are

known to be hosts of solidaginis: altissima, canadensis var. hargeri, gigantea, gram-

ini folia, rugosa, serotina, and ulmifolia. Curran (1925) also reported S. serotina var. salebrosa. See also the Host sections of the two subspecies (below). discussion.

E. solidaginis is one of the earliest American tephritids to be recog¬

nized taxonomically as evidenced by the early date of its original description and the large body of literature referring to it. According to the International Code of Zoological Nomenclature (Ride et al. 1985), names published as varieties before 1961 are to be accorded subspecific status; therefore, we follow Ming (1989) in treating the name fascipennis Curran 1923 as a subspecies.

Eurosta solidaginis solidaginis (Fitch) (Figs. 67, 213) RECOGNITION.

As discussed in the Recognition section of solidaginis, four dis¬

tinctive light areas, well separated from each other, are present in the wing. In addition to the areas at the apex of vein At + CuA2 and at the wing tip, those at the

178

Systematic Treatment of the Genera apex of the pterostigma and at the posterior border of vein M are distinctly separated except for occasional small hyaline droplets in the dark area between them (fig. 213, a, c). These spots are connected in s. fascipennis (fig. 214). distribution.

The distribution of this subspecies has not been mapped sepa¬

rately. According to Ming (1989), this subspecies is widely distributed in the eastern United States and Canada and occurs west to North and South Dakota, Minnesota, and Iowa and south to Texas. hosts.

It is assumed that all of the plants listed in the host section of solidaginis

(above) that occur within the range of this subspecies are hosts.

Eurosta solidaginis fascipennis Curran (Fig. 214)

Eurosta solidaginis var. fascipennis Curran 1923: 302 (male holotype, CNC; Ot¬ tawa, Ont., Canada [error; see Curran 1925]). - Curran 1925: 128 (host, taxon¬ omy). - Foote 1965a: 663 (in catalog). - Wasbauer 1972: 116 (hosts). - Steyskal and Foote 1977: 151 (in key). - Arnaud and Owen 1981: 149 (type data).

Eurosta solidaginis var. subfasciata Curran 1923: 302 (male holotype, CNC; Ver¬ non, B.C.). - Foote 1965a: 663 (in catalog). - Wasbauer 1972: 116 (host). Steyskal and Foote 1977: 151 (in key).-Arnaud and Owen 1981: 149 (type data). - Ming 1989: 77 (synonymy).

Eurosta solidaginis var.: Curran 1925: 129 (taxonomy). Eurosta solidaginis fascipennis: Strickland 1938: 204 (Alta.). — Ming 1989: 82 (revision).

Eurosta solidaginis sub fascipennis [error]: Strickland 1938: 204 (Alta.). recognition.

The wing of this subspecies contains only three well-defined light

areas (fig. 214) (see Recognition section of solidaginis). The light area at the apex of the pterostigma and that on the posterior border of vein M are joined in a distinct diagonal band across the wing, the band sometimes with small dark spots that do not interrupt its appearance. distribution.

The distribution of this subspecies has not been mapped sepa¬

rately. According to Ming (1989), this subspecies occurs from Washington east to Minnesota and from Alberta south to Colorado.

Solidago serotina var. salebrosa, to date the only known host of this sub¬ species, was reported by Wasbauer (1972). Although this form is known from Soli¬ host.

dago sp., its host relationships throughout its range have not been studied. discussion.

In his original description of fascipennis, Curran (1923) erred in

recording the type locality as Ottawa. In a subsequent paper (Curran 1925), he

179

Eutreta

indicated that the specimens were reared in Ontario but probably were actually collected at an unknown locality in the western United States. Specimens from the midwestern United States are difficult to name because the diagonal hyaline band is intermediate between the two subspecies. For that reason, a better understanding of host relationships and possibly biochemical analyses are needed to clarify the status of these two subspecies.

Genus Eutreta Loew Icaria Schiner 1868: 276 (type species, Trypeta sparsa Wiedemann 1830: 492, by original designation) (preoccupied Saussure, 1853). — Coquillett 1910: 543, 555 (type data).

Eutreta Loew 1873: 276 (type species, Trypeta sparsa Wiedemann 1830: 492, desig¬ nated by Coquillett 1910: 543). - Williston 1896: 122 (in key). - Coquillett 1899c: 268 (in key). — Aldrich (part) 1905: 608 (in catalog). — Williston (part) 1908: 287 (in key). - Coquillett 1910: 543 (type designation). - Cresson 1914a: 276 (taxonomy). — Hendel 1914b: 93 (in key). — V. T. Phillips 1923: 122, 140 (in key, review). - Banks 1926: 44 (key to species). - Curran 1932b: 4, 15 (in key, review). - Curran 1934: 291 (in key). - Foote and Blanc 1963: 27 (review, Calif.). - Foote 1965a: 661 (in catalog). - Stoltzfus and Foote 1965: 263 (biology). Furniss and Barr 1975: 20 (gall parasites). - Stoltzfus 1977: 369 (revision). Freidberg 1981: 93 (mating behavior, review). - Freidberg 1984: 129 (review, galls).

Trypeta (Eutreta): Osten Sacken 1878: 191 (in catalog). Trypeta Meigen [misidentification]: Aldrich (part) 1905: 604 (in catalog). Eutreta (Phasmatocepbala) Hering 1937: 297 (type species, Eutreta rhinopbora Hering, by original designation). - Stoltzfus 1977: 374 (synonymy).

Eutreta (Uncaculeus) Stoltzfus 1977: 390 (type species, Trypeta diana Osten Sacken, by original designation).

Eutreta (Setosigena) Stoltzfus 1977: 395 (type species, Trypeta rotundipennis Loew, by original designation). recognition.

These dark-winged flies range in size from about 3 mm to over 8

mm. Most species are quickly recognized as Eutreta by their dark brown to black wings which contain a spattering of small, light-colored round spots and, except in males of simplex and decora, have an apical white crescent. Other distinctive traits of

Eutreta are as follows: the dark color of the wing extends completely to the base; head with three pairs of frontal bristles and two pairs of orbitals, the posterior pair reclinate and white or yellowish; two pairs of scutellar bristles; one pair of dorsocentral bristles, situated closer to the transverse suture than to the level of the supraalar bristles; the frons at its vertex at least as wide as one eye; the scutellum not swollen or polished; vein R4+5 bare or with only a few setae dorsally (in Nearctic species); and the apex of cell cup elongate at its lower corner. The femora are usually

Systematic Treatment of the Genera

concolorous with the thorax; the tibiae and tarsi are commonly lighter but in some species also are concolorous with the thorax. The color of the abdominal tergites may match that of the dark scutum, or in some species such as diana, some segments may be bright red. The 17 species recorded from America north of Mexico vary in the degree of coalescence of the light spots of the wing. In species such as hespera, most of the light spots of the central and basal areas are coalesced, but in diana, all of them are well separated. The male of divisa has two white diagonal bars across the wing; these are entirely lacking in the female.

discussion.

Eutreta includes more than 30 species which occur from southern

Canada to Brazil. One species from Mexico, the lantana gall fly (E. xantbochaeta Aldrich), has been introduced into Hawaii for the control of Lantana camara. All species whose biology is known form galls. Artemisia species are the only known hosts of species of the subgenus Uncaculeus, but a broad range of Asteraceae is attacked by the species of the subgenus Eutreta. Several Neotropical species are unusual for New World Tephritinae in that Verbenaceae rather than Asteraceae are host plants. Stoltzfus (1977) revised the genus, dividing it into three subgenera based on the degree to which the oral margin is produced, the presence or absence of large hyaline spots along the costal margin, and the length of the oviscape compared with that of the abdominal tergites. These subgenera are reflected in our key to genera as follows: Uncaculeus, couplets 1-9; Setosigena, couplets 10-11; Eutreta, couplets 12-16. We concur in Stoltzfus’ recognition of the genus as one entity and his transfer of baccharis Coquillett from Eutreta to Tephritis (see discussion of that genus). Stoltzfus (1977) provided a key to the subgenera and species, wing photographs, line drawings of genitalia, and distribution maps for most of the species. Metatephritis

fenestrata, whose wing has an apical hyaline mark but lacks medial hyaline spots, may be related to the subgenus Uncaculeus (see discussion section of Metatephritis).

Key to Subgenera and Species of U.S. and Canadian Eutreta 1. In profile, oral margin only slightly produced (figs. 234, a; 235, c); costal margin brown, without hyaline spots (fig. 216, a); face without a pair of rounded dark spots; aculeus dorsally at base of tip with transverse row of small barbs (subgenus Uncaculeus).2 In profile, oral margin distinctly produced, the fronto-orbital angle usually 90° or less (fig. 238, b); costal margin usually with hyaline spots or lighter areas (fig. 225, b); face with or without spots; aculeus without dorsal row of barbs .20 2. Antenna as long as face, or nearly so (fig. 235, b).3 Antenna distinctly shorter than face (fig. 238, a) .4 3. Fronto-orbital plate darker than remainder of frons; face yellowish to silvergray tomentose; parafacial spot small (fig. 235, a); wing pattern of male as in fig. 216; western U.S.oregona Curran Fronto-orbital plate same color as frons; face shining black or reddish black in male and tomentose with whitish, yellowish, or reddish ground color

a

Figures 216-225. Right wings, Eutreta spp. 216, oregona Curr., male; 217, longicornis Snow, male; 218, modocorum Blanc, female; 219, divisa Stoltzfus, male; 220, divisa Stoltz., female; 221, diana (O.S.), female; 222, decora Stoltz., female; 223, pollinosa Curr., female; 224, coalita Blanc, female; 225, fenestra Stoltz., female.

181

Systematic Treatment of the Genera in female; parafacial spot large; wing pattern as in fig. 217; northcentral U.S., southcentral Canada.longicornis Snow 4. Wings black.5 Wings brown.7 5. Pleuron pinkish amber; wing pattern as in fig. 218. .modocorum Blanc Pleuron black.6 6. Antenna dark brown to black; male with 2 prominent diagonal hyaline bars at center of wing (fig. 219, a); female with few spots in posterior half of wing (fig. 220, a); northwestern U.S., southwestern Canada. .divisa Stoltzfus Antenna yellow to light brown; male without prominent hyaline bars in center of wing; female with spots nearly as numerous in posterior half as in anterior half of wing (fig. 221); western U.S., southwestern Canada .diana (Osten Sacken) 7. Wing dark brown, all hyaline spots distinct; male without apical hyaline crescent.8 Wing brown, some of discal hyaline spots coalesced; male with apical hyaline crescent.9 8. Wing with hyaline spots relatively large; aculeus tip slender, elongate, with¬ out strong lateral barbs; California, Colorado ....... simplex Thomas Wing with hyaline spots relatively small (fig. 222); aculeus tip short, broad, with strong lateral barbs; Utah .decora Stoltzfus 9. Cells be and c lighter posteriorly than anteriorly (fig. 223, a); spots in cell dm fused into a yellowish brown streak in middle of wing (fig. 223, b); anteapical brown band about as wide as apical hyaline area (fig. 223, c); western U.S., southwestern Canada.pollinosa Curran Cells be and c evenly dark (fig. 224, a); discal spots large, poorly delimited, fused, especially in cell r2+3 but not forming a distinctive yellowish brown streak; anteapical brown band distinctly wider than apical hyaline area (fig. 224, c); California.coalita Blanc 10. Wing broad; marginal hyaline spots large along both anterior and posterior borders (figs. 225, b; 226, c); oviscape shorter than last 2 abdominal tergites (fig. 236); most setae of oral margin, palpi, and postgenae whitish and inflated, several black setae along oral margin and apically on palpi (fig. 238, c); face without rounded pair of dark spots (subgenus Setosigena) .11 Wing width and spots variable; oviscape at least as long as last two tergites (fig. 237); setae of oral margin and lower postgena mostly brown to black; face with pair of dark spots except in intermedia (subgenus Eutreta).12 11. Wing with many coalesced hyaline spots at center of disk (fig. 225, a); oral margin, antenna, postgena, pleuron, and coxae tinged with violet; apical hyaline crescent complete (fig. 225, c); Arizona .fenestra Stoltzfus Wing with hyaline spots usually distinct; no large hyaline area present in center of wing (fig. 226, a); violet color absent from areas described above; apical hyaline crescent usually interrupted along veins R4 + 5 and M by extension of brown color of wing (fig. 226, b); Texas, eastern U.S. .rotundipennis

(Loew)

183

Eutreta a

Figures 226-233. Right wings, Eutreta spp. 226, rotundipennis (Lw.), male; 227, simplex Thom., female; 228, intermedia Stoltzfus, female; 229, frontalis Curr., male; 230, hespera Banks, male; 231, angusta Banks, female; 232, caliptera (Say), male; 233, novaeboracensis (Fitch), male.

12. Face without dark spots; costal margin with only 1 hyaline spot, at apex of vein Rt; light spots in wing small; Arizona .intermedia Stoltzfus Face with a pair of large dark brown to black spots (fig. 241, c) .13 13. Base of wing with hyaline spots coalesced, especially in anal lobe and cell cua! (fig. 230, a) .14 Hyaline spots of wing usually separated, not forming a single light area near base of wing (fig. 233, c) .15 14. Costa without hyaline spots beyond pterostigma (fig. 229, a); frons with lateral margins gradually narrowing from vertex to antenna (fig. 240, a); parafacial spot small and indistinct (fig. 242, a); northeastern U.S., south¬ eastern Canada.frontalis Curran

184

Systematic Treatment of the Genera

Figures 234, 235. Lateral view of head, Eutreta spp. 234, diana (O.S.); 235, oregona Curr. Figures 236, 237. Dorsal view of abdomen, Eutreta spp. 236, rotundipennis (Lw.).; 237, hespera Banks. Figures 238, 239. Lateral view of head, Eutreta spp. 238, rotundipennis (Lw.); 239, novaeboracensis (Fitch).

Costa with hyaline spots beyond pterostigma (fig. 230, b); frons with lateral margins markedly narrowing near antenna (fig. 241, b); parafacial spot dark brown, distinct (figs. 241, a; 243, a); western U.S. and Canada .. .hespera Banks 15. Subapical unpunctated brown band narrower than apical hyaline crescent (fig. 231, b); costa usually with at least 1 hyaline spot beyond pterostigma (fig. 231, a); southwestern U.S.angusta Banks Subapical unpunctated brown band as wide as apical hyaline crescent (figs. 232, c; 233, b); costa without hyaline spots beyond pterostigma or spots very obscure (fig. 232, b) .16

Figure 240. Dorsal view of head, Eutreta frontalis Curr. Figure 241. Frontal view of head, Eutreta hespera Banks. Figures 242, 243. Lateral view of head, Eutreta spp. 242, frontalis Curr.; 243, hespera Banks.

185

186

Systematic Treatment of the Genera

16. Body dark brown to grayish brown; hyaline spot on costa at pterostigma narrow, not extending into cell (fig. 232, a); wing length 4.2-5.7 mm; eastern U.S.caliptera (Say) Body often slightly reddish to yellow brown; hyaline spot on costa at pterostigma broader, usually extending as a small wedge into cell r1 (fig. 233, a); wing length 5.0-7.5 mm; eastern U.S., southeastern Canada .novaeboracensis (Fitch)

Eutreta angusta Banks (Fig. 231; Map 23)

Eutreta sparsa (Wiedemann) [misidentification]: Loew 1873: 275 (taxonomy). W. A. Snow 1894: 167 (taxonomy). - Osten Sacken 1877: 345 (taxonomy). Doane 1899: 184 (taxonomy). - C. F. Baker 1904: 30 (Nev.). - Cresson 1907: 198 (Tex.). - Painter 1935: 83 (biology). - Felt 1940: 260 (gall).

Eutreta angusta Banks 1926: 44 (female lectotype, MCZ; Tex.). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 377 (revision, lectotype designation). - Steck 1981: 18 (host). — Steck and Wharton 1985: 296 (description of larva).

Eutreta pacifica Curran 1932b: 17 (male holotype, AMNH; Pomona, Calif.). Curran 1934: 286 (figure of wing). - Foote and Blanc 1963: 30 (review, Calif.). Foote 1965a: 661 (in catalog). - Harris and Piper 1970: 134 (hosts). - Frick 1972: 629 (biology). - Wasbauer 1972: 116 (hosts). - Goeden and Ricker 1976c: 1174

o

Eutreta angusta Eutreta rotundipennis

ft

Eutreta rotundipennis (state record only)

Map 23. Distribution of Eutreta angusta and E. rotundipennis.

187

Eutreta (host). - Stoltzfus 1977: 377 (synonymy). - Arnaud and Owen 1981: 150 (type data). recognition.

E. angusta one of the largest of our species of Eutreta, ranging in

body length from about 3.75 mm to over 6 mm. The wings and body are brown to grayish brown. The very large paired brownish black abdominal tergal spots cover most of the abdominal dorsum, leaving only a narrow central gray longitudinal stripe. The legs are concolorous with the body but are slightly lighter. E. angusta is similar to caliptera and novaeboracensis but differs from those species in that the subapical solid brown band of its wing is narrower than the apical whitish hyaline crescent (fig. 231, b), and several amber light spots are present along the costa bordering cell r2+3. In both caliptera and novaeboracensis, the spotless brown subapical band is wider than the widest part of the apical hyaline crescent, and there are no light spots along the costa bordering cell r2+3. The wing spots of angusta are small, round, silvery gray, and not coalesced, even near the wing base. There is a pair of large black spots on the face and one on the parafacial between the compound eye and the antennal base. Other recognition characters are given in the key. distribution. Map 23.

E. angusta also occurs in Mexico, Guatemala, and Be¬

lize.

E. angusta has been reared from species of Ambrosia, Callistephus, Chry¬ santhemum, Ratibida, Senecio, and Vernonia. See Stoltzfus (1977) for additional hosts.

details.

Eutreta caliptera (Say) (Fig. 232; Map 24)

Trypeta caliptera Say 1830: 187 (female neotype, NMNH; Lafayette, Ind.). - Osten Sacken 1858: 79 (in catalog). — Loew 1862c: 58 (synonymy). — Loew 1873: 336 (synonymy). — Aldrich 1905: 608 (in catalog). — Stoltzfus 1977: 379 (neotype designation).

Trypeta calyptera [error or emendation]: Leconte 1859: 370 (taxonomy). Eutreta sparsa (Wiedemann) [misidentification]: Thompson 1907: 71 (gall). — Painter (part) 1935: 83 (biology, host). — Felt (part) 1940: 300 (gall). - Blanton 1952: 75 (host). - Novak et al. 1967: 147 (biology, hosts).

Eutreta caliptera: Stoltzfus 1977: 379 (revision). See additional references in the synonymy of E. novaeboracensis. recognition.

E. caliptera is similar to novaeboracensis but is smaller: the body

length is a little over 4 mm to about 6.5 mm and the wing length is 4.2—5.7 mm; in

novaeboracensis, the body length is usually over 6.5 mm and the wing length 5—7.5 mm. The blade angle of the aculeus is less than 46° in caliptera and greater than 48°

Systematic Treatment of the Genera

• Eutreta caliptera o Eutreta bespera

Map 24. Distribution of Eutreta caliptera and E. hespera.

in novaeboracensis. The whitish wing spot near the terminus of vein Rt is very small in caliptera (fig. 232, a); it is larger in novaeboracensis and extends farther posteri¬ orly into cell rv E. caliptera differs from angusta in that the spotless brown subapical wing band is wider than the apical hyaline crescent. Other separation characters are given in the key to species.

distribution.

Map 24.

E. caliptera forms stem galls on many genera of Asteraceae, including Ambrosia, Bidens, Chrysanthemum, Helianthus, and Vernonia. HOSTS.

189

Eutreta

■Q

• Eutreta coalita o Eutreta modocorum ▲ Eutreta oregona

• Eutreta decora o Eutreta fenestra ▲ Eutreta simplex

Map 25. Distribution of Eutreta coalita, E. decora, E. fenestra, E. modocorum, E. oregona, and E. simplex.

Eutreta coalita Blanc (Fig. 224; Map 25) Eutreta coalita Blanc in Foote and Blanc 1979: 162 (female holotype, NMNH; 5 mi. e. Smith Meadow, 9 Mile Cyn., Tulare County, Calif.). recognition. E. coalita, an extremely rare species, is known only from the holotype and is unique in having the light wing spots poorly delimited and coalesced in some areas (fig. 224, b), but these spots do not form a central basal yellowish streak as in pollinosa (fig. 223, b). In coalita, the subapical spotless brown band is distinctly wider than the apical hyaline area, whereas in pollinosa this band is about as wide as the apical hyaline area. E. coalita lacks the dark parafacial spot between the eye and the antennal base. The first and second costal cells are lighter anteriorly and posteriorly. See the key to species for other differences between coalita and other similar Eutreta species such as decora. distribution. Map 25. host. Not known.

Eutreta decora Stoltzfus (Fig. 222; Map 25) Eutreta decora Stoltzfus 1977: 391 (female holotype, NMNH; Green Cyn., Utah).

Systematic Treatment of the Genera

recognition.

The wing of decora (fig. 222) is dark brown with very small,

widely scattered light spots which are located almost entirely in the anterior half of the disk. The light-colored antennae do not attain the oral margin. The occiput and postgena are grayish brown. E. pollinosa is similar to decora but has a lighter brown wing; the light spots are more numerous, larger, and those near the center and base are coalesced into a yellow-brown streak. The wing of coalita differs from that of decora by its much larger and more poorly defined light spots which extend into cell cua! and the anal lobe. E. decora is a rare species. The male has not been described previously, but two males in the UKL collection resemble the females except in lacking the apical hyaline crescent. Thus decora, like simplex, appears to be sexually dimorphic in this character.

distribution.

HOST.

Not

Map 25.

known.

Eutreta diana (Osten Sacken) (Figs. 108, 221, 234; Map 26) Trypeta (Eutreta) diana Osten Sacken 1877: 347 (female holotype, MCZ; Mo.). Osten Sacken 1878: 191 (in catalog). Eutreta diana: Osten Sacken 1880: 53 (host). - Townsend 1893a: 10 (host). - W. A. Snow 1894: 168 (taxonomy). - Doane 1899: 184 (distribution, host). - Aldrich 1905: 608 (in catalog). - Thompson 1907: 71 (biology). - Cole and Lovett 1921: 324 (review). - V. T. Phillips 1923: 141 (figure of wing). - Curran 1932b: 19 (review). - Knowlton and Harmston 1937: 145 (Utah). - Strickland 1938: 204 (Alta.). — V. T. Phillips 1946: 115 (host). — Foote 1962: 178 (taxonomy). — Foote and Blanc 1963: 28 (review, Calif.). - Fronk et al. 1964: 575 (hosts, associates, galls). - Foote 1965a: 661 (in catalog). - Wasbauer 1972: 116 (hosts). - Stoltzfus 1977: 391 (revision). - Foster and Benbow 1978: 48 (galls, biology). - Steck 1981: 18 (host). - Benbow and Foster 1982: 19 (biology, Texas). - Steck and Wharton 1985: 296 (description of larva). — Goeden 1990a: 24 (life history, hosts). Eutreta diana var. tricolor Snow 1894: 168 (female lectotype, UKL; Mont.). Aldrich 1905: 608 (in catalog). - Byers et al. 1962: 180 (type data). - Foote 1962: 178 (type designation). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 391 (synonymy). Eutreta tricolor: C. F. Baker 1904: 30 (Nev.). Gymnocarena tricolor: Ffuber 1927: 48 (parasite). Eutreta diana tricolor: Curran 1932b: 19 (synonymy). - Wasbauer 1972: 116 (host). Eutreta jonesi Curran 1932b: 19 (female holotype, AMNH; Antelope Mt., Harney County, Oreg.). - Knowlton and Harmston 1937: 145 (Utah). - Foote 1965a: 661 (in catalog). - Wasbauer 1972: 116 (host). - Stoltzfus 1977: 391 (synonymy). Arnaud and Owen 1981: 150 (type data). “Trypetid”: Felt 1940: 334 (gall).

191

Eutreta

E. diana is a small, black-winged species with a yellowish, pink, red, or black abdomen. The pleuron is black. The oral margin is not produced. The costa along the margin of cell is void of any light or hyaline spots (fig. 221). It differs from oregona and longicornis by its light yellow to amber antenna, which recognition.

does not attain the oral margin; in oregona and longicornis, the antenna is dark gray to black and is long enough to reach or surpass the oral margin. E. divisa is similar to diana but the male differs by having two prominent diagonal white stripes across the wing (fig. 219, a). Females of divisa can be distinguished from those of diana in lacking small rounded hyaline spots in the posterior half of the wing (especially in cell cuaj and the anal lobe); in the female of diana, these spots are uniformly spread over the wing to the posterior margin.

Systematic Treatment of the Genera distribution.

Map 26.

According to Goeden (1990a), E. diana forms galls on six species and two varieties of Artemisia. A. tridentata is the principal host in California. hosts.

discussion.

E. diana is one of our most commonly collected Eutreta species.

Benbow and Foster (1982) gave an account of the biology of diana on Artemisia filifolia in northwestern Texas. Goeden (1990a) has shown that, in that location, the biology of diana is quite different from that in California and suggests that the Texas population may represent a sibling species which should be studied further.

Eutreta divisa Stoltzfus (Figs. 219, 220; Map 27) Eutreta divisa Stoltzfus 1977: 393 (male holotype, originally in NMNH, destroyed; Regina, Idaho). The male of divisa differs from all other species of Eutreta occur¬ ring north of Mexico by its two striking diagonal white wing stripes (fig. 219, a). Females are very similar to those of diana but differ in lacking the very small round recognition.

hyaline spots in the posterior portion of the wing (see Recognition section for diana). In divisa, the oral margin is not produced, and the antenna is brown to dark brown and rarely, if ever, attains the oral margin. Its wing lacks amber or hyaline spots on the costa distad of the pterostigma. The wing and pleuron are black. distribution.

host.

Map 27.

E. divisa has been reared from galls on Artemisia tridentata.

Eutreta fenestra Stoltzfus (Fig. 225; Map 25) Eutreta fenestra Stoltzfus 1977: 395 (female holotype, badly damaged, NMNH; Ramsey Cyn., Huachuca Mts., Ariz.). In fenestra, as in rotundipennis, most of the setae of the oral margin, postgena, and palpus are whitish and inflated. The wing is broad and has recognition.

large hyaline spots spread rather uniformly along the anterior and posterior margins (fig. 225). The oviscape is shorter than the two preceding abdominal tergites. The hyaline spots of the wing are not uniform in size. E. fenestra differs from rotundipen¬ nis in having many of the hyaline wing spots coalesced in cell br and adjacent cells; by its complete hyaline crescent, and by the violet tinge on the antenna, oral margin, postgena, pleuron, and coxae. In rotundipennis, the hyaline wing spots are not coalesced (fig. 226, a), the hyaline crescent is interrupted by the extension of the

193

Eutreta

Map 27. Distribution of Eutreta divisa and E. longicornis.

brown wing color along veins R4+5 and M to the wing margin, and the violet color is absent in the locations indicated above. distribution.

host.

Not

Map 25.

known.

discussion.

E. fenestra and rotundipennis were included in his subgenus Set-

osigena by Stoltzfus (1977).

Systematic Treatment of the Genera

Eutreta frontalis Curran (Figs. 229, 240, 242; Map 26) Eutreta frontalis Curran 1932b: 16 (female holotype, AMNH; Valley of Black Mts., N. Car.). — Foote 1965a: 661 (in catalog). — Novak et al. 1967: 147 (host). — Wasbauer 1972: 116 (host). - Stoltzfus 1977: 381 (revision). - Arnaud and Owen 1981: 150 (type data). In the wing of frontalis (fig. 229), the hyaline spots are coalesced near the base, and the costa is without light spots distad of the light area in cell rj. recognition.

The second costal cell is almost totally dark and lacks any large hyaline spots. The oral margin of the head is distinctly produced, and the face has a pair of large dark spots. The frons gradually narrows from the vertex to the antennae, and the parafacial spot is small and indistinct. Most of the setae on the second antennal segment are whitish (some may be brown). E. hespera is similar but has one or more light spots on the costa distad of the hyaline area at the base of cell r1? the second costal cell has two large hyaline spots, the frons is suddenly narrowed near the antenna, and the parafacial spot is dark brown and distinct. E. intermedia and simplex differ in lacking the pair of dark spots on the face. E. angusta, caliptera, and novaeboracensis differ from frontalis in lacking any coalescence of the hyaline wing spots. distribution.

Map 26.

E. frontalis forms crown galls on Aster spp. See Stoltzfus (1977) for details of the biology of this species. hosts.

Eutreta hespera Banks (Figs. 230, 237, 241, 243; Map 24) Eutreta sparsa (Wiedemann) [misidentification]: Woodworth 1913: 137 (Calif.). Eutreta hespera Banks 1926: 44 (male lectotype, MCZ; Manitou, Colo.). - Foote and Blanc 1963: 29 (review, Calif.). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 382 (revision, lectotype designation). See possible additional references in the synonymy of E. novaeboracensis. Of the Eutreta species in America north of Mexico, hespera is similar only to frontalis in that the hyaline spots are coalesced in the central basal part of the wing disk (fig. 230, a). It differs from frontalis by having one or more light recognition.

spots on the costa distad of the hyaline area at the base of cell rl5 by having two large hyaline spots in the second costal cell, by its distinct and dark brown parafacial spot, and by the abrupt narrowing of its frons at the antennal bases. Some specimens of angusta may be confused with those of hespera, but the subapical spotless band of the wing of angusta is narrower than the apical hyaline crescent, whereas in hespera it is wider.

Eutreta distribution.

host.

Map 24.

Stoltzfus (1977) recorded a single male reared from goldenrod.

Eutreta intermedia Stoltzfus (Fig. 228; Map 28) Eutreta intermedia Stoltzfus 1977: 382 (female holotype, UKL; 17 mi. n. Altacomulco, Mex.). Eutreta navajorum Blanc in Blanc and Foote 1987: 427 (female holotype, NMNH; Sunnyside Cyn., Fluachuca Mts., Cochise County, Ariz.). New synonymy.

▲ Eutreta intermedia • °

Eutreta novaeboracensis Eutreta pollinosa

Map 28. Distribution of Eutreta intermedia, E. novaeboracensis, and E. pollinosa.

196

Systematic Treatment of the Genera

recognition.

E. intermedia is a medium-sized Eutreta with a dark brown body

and a dark brownish to black wing. In profile, the oral margin projects forward, although not as strongly as in the other species in the subgenus Eutreta. It might be confused with some species of the subgenus Uncaculeus, which also lack a pair of dark spots on the face. E. simplex, which often has large hyaline spots in cell rl5 especially may be confused with it. It differs from intermedia in that large hyaline areas are sometimes present in the costal cell, the spots of the wing are comparatively large (fig. 227), and the male lacks an apical hyaline crescent. The aculeus and male genitalia of these species also distinguish them. distribution.

Map 28. E. intermedia is known from the United States to central

Mexico. host.

Not

known.

Eutreta longicornis Snow (Fig. 217; Map 27) Eutreta longicornis Snow 1894: 166 (male holotype, UKL; Mont.). - Aldrich 1905: 608 (in catalog). - Cole and Lovett 1921: 320 (Oreg.). - Curran 1932b: 15 (in key). - Strickland 1938: 204 (Alta.). - Byers et ai. 1962: 180 (type data). - Foote 1962: 175 (type data). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 394 (revision). Eutreta facialis Curran 1932b: 17 (female holotype, AMNH; Huntley, Mont.). Strickland 1946: 166 (Alta.). - Stoltzfus 1977: 394 (synonymy). - Arnaud and Owen 1981: 150 (type data). As the name implies, the antenna of longicornis is distinctively long, reaching or surpassing the oral margin; the same is true also of oregona. Both recognition.

are small black-bodied and black-winged flies. The orbital plates of oregona are contrasting dark brown or black, whereas those of longicornis are concolorous with the frons. The face of the male of longicornis is shining black or reddish black; that of the female is tomentose with whitish, yellowish, or reddish ground color but may be blackish along the facial ridges. The face of oregona is silver-gray to yellowish tomen¬ tose in both sexes. The parafacial spot of longicornis is large and black, whereas that of oregona is small (fig. 235, a), often brownish, and not prominent. distribution.

Map 27.

E. longicornis forms galls on the lateral branches of Artemisia cana. For details, see Stoltzfus (1977). host.

Eutreta modocorum Blanc (Fig. 218; Map 25) Eutreta modocorum Blanc in Blanc and Foote 1987: 425 (female holotype, NMNH; 4 mi. e. Davis Cr., Modoc County, Calif.).

197

Eutreta

recognition.

E. modocorum resembles diana and divisa. It differs from diana

by its almost total lack of spots in the wing (fig. 218), by a prominent white spot near the apex of vein AjCuA^ by its pinkish amber pleuron and pink head, and by the totally amber-yellow legs. E. modocorum differs from divisa by the absence of two prominent diagonal white slash marks on the wing of the male, and it differs in both sexes by its pinkish amber pleuron in contrast to the black pleuron of divisa. distribution.

Map 25. E. modocorum is known only by two specimens from

the type locality.

host.

E. modocorum was reared from stem tip galls on Artemisia sp.

Eutreta novaeboracensis (Fitch) (Figs. 233, 239; Map 28)

Acinia novaeboracensis Fitch 1855a: 771 (female syntype, NMNH; N.Y.; neotype, CUI; Sixmile Cr. above Ithaca Res., 0.2 mi. nw. Burns Rd., Tompkins County, N.Y.). - Fitch 1856: 67 (taxonomy). - Aldrich 1905: 608 (in catalog). - Stoltzfus 1977: 385 (neotype designation).

Trypeta novaeboracensis: Osten Sacken 1858: 79 (in catalog). - Loew 1862c: 59 (synonymy). - Loew 1873: 337 (synonymy).

Trypeta sparsa (Wiedemann) [misidentification]: Loew 1862c: 78 (review). - Couper 1864: 20 (Que.). — Wulp 1883: 54 (Que.). - Chagnon 1901: 14 (Can.). Washburn 1906: 80 (Minn.).

Eutreta sparsa [misidentification]: Loew 1873: 274 (taxonomy). - Wulp 1883: 54 (Que.). - Chagnon

1901:

14 (Canada). - Washburn 1906:

80 (Minn.). -

Thompson 1907: 71 (biology). - Stebbins 1910: 50 (gall, biology). - Winn and Beaulieu 1915: 153 (Que.). - Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 140 (biology, distribution). - Johnson 1925a: 97 (Harris collection). - Johnson 1925b: 263 (N. Engl.). — Johannsen 1926: 159 (N.Y.). - Johnson 1927: 218 (Mt. Desert). — Curran 1930: 77 (N.Y.). - Curran (part) 1932b: 15 (in key). - Petch and Maltaise 1932: 56 (Que.). — Felt (part) 1940: 322 (gall). — Foote (part) 1965a: 661 (in catalog). — Wasbauer (part) 1972: 117 (hosts).

Trypeta (Eutreta) novaeboracensis: Osten Sacken 1878: 192 (in catalog). Eutreta sp.: Stoltzfus and Foote 1965: 263 (biology).

Eutreta novaeboracensis: Stoltzfus 1977: 384 (revision). The following references to E. sparsa are applicable to caliptera (Say), hespera Banks, novaeboracensis (Fitch), or other species of Eutreta. Further studies are needed to make the proper assignments.

Trypeta (Eutreta) sparsa (Wiedemann) [misidentification]: Loew 1873: 336 (taxon¬ omy). — Osten Sacken 1877: 345 (taxonomy). — Osten Sacken 1878: 192 (in catalog).

Eutreta sparsa [misidentification]: Doane 1899: 184 (taxonomy). — Johnson 1900b: 688 (N.J.). — Coquillett 1904: 30 (Calif., Nev.). - F. H. Snow (part) 1904: 167

Systematic Treatment of the Genera

(taxonomy). - F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 608 (in catalog). Cresson 1907: 101 (N. Mex.). - Johnson 1910: 802 (N.J.). - Cole 1912: 158 (Calif.). - Thomas 1914: 426 (taxonomy). - Felt 1918: 198 (gall). - Banks 1926: 44 (taxonomy). - Cole 1927: 444 (male genitalia). - Leonard 1928: 852 (N.Y.). Brimley 1938: 384 (N. Car.).-V. T. Phillips 1946: 115 (hosts). - Strickland 1946: 166 (Alta.). — Steyskal 1957: 94 (eye color).

recognition.

E. novaeboracensis is one of the largest species of Eutreta in Amer¬

ica north of Mexico. It has a reddish to yellow-brown body and legs and brownish black wings which are well covered with small, round, grayish white spots (fig. 233). The face has a pair of large black spots, and the oral margin is distinctly produced. This species differs from angusta by having the unpunctured subapical wing band as wide as or wider than the apical hyaline crescent and by lacking amber-hyaline spots (or having very obscure spots) on the costa distad of the terminus of vein

It is

quite similar to caliptera but differs from that species by its greater wing length (5.07.5 mm in novaeboracensis and 4.2-5.7 mm in caliptera), by the greater extension of the hyaline costal spot into cell r1 (fig. 233, a), and by its reddish to yellow-brown body color (caliptera is dark brown to grayish brown). In the female of novaebora¬

censis, the lateral blade angle of the aculeus is more than 48°, whereas in caliptera this angle is less than 46°. E. novaeboracensis differs from frontalis in that the round hyaline wing spots are distinct, but in frontalis they are coalesced, especially in cell

cuaA and the anal lobe. distribution.

host.

Map 28.

E. novaeboracensis feeds as a larva in the rhizomes of Solidago altissima.

See Stoltzfus (1977) for details.

discussion.

The neotype designation by Stoltzfus (1977) was unnecessary be¬

cause a female syntype is in the NMNFL An application to the International Com¬ mission of Zoological Nomenclature must be made to determine which specimen should be considered the valid type.

Eutreta oregona Curran (Figs. 216, 235; Map 25)

Eutreta oregona Curran 1932b: 18 (male holotype, AMNH; Blitzen R., Oreg.) Foote and Blanc 1963: 30 (review, Calif.). - Foote 1965a: 661 (in catalog). Wasbauer 1972: 116 (host). - Stoltzfus 1977: 394 (revision). - Arnaud and Owen 1981: 150 (type data).

RECOGNITION.

E. oregona, a very small, grayish black Eutreta, is characterized by

its long antenna which nearly attains the oral margin (fig. 235, b), and by the oral margin not being produced (fig. 235, c). It closely resembles longicornis in these characters but is distinguished from that species in that its orbital plates are dark

199

Eutreta

brown to black, contrasting against the light brown color of the frons; in longicornis, the orbital plate and frons are concolorous. The face of oregona in both sexes is silver-gray to yellow tomentose, whereas in the male of longicornis, the face is shining black or reddish black and in the female is tomentose with a whitish or reddish ground color; sometimes the facial ridges are black. The parafacial spot is small and usually brown and insignificant in oregona (fig. 235, a), whereas in longicornis it is large, black, and prominent. distribution. Map 25.

host.

E. oregona forms galls on Artemisia tridentata.

Eutreta pollinosa Curran (Fig. 223; Map 28)

Eutreta pollinosa Curran 1932b: 18 (male holotype, AMNH; Antelope Mt., Harney County, Oreg.). - Foote and Blanc 1963: 32 (review, Calif.). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 395 (revision). - Arnaud and Owen 1981: 150 (type data).

recognition.

E. pollinosa is a small, light brown species with an antenna that

does not reach the nonproduced oral margin. It has grayish brown wings with large hyaline spots which are coalesced in cells bm and dm (fig. 223). E. decora differs from pollinosa in having a much darker brown wing and fewer hyaline wing spots, none of which are coalesced. E. coalita differs in that the hyaline wing spots have indistinct borders, some being fused but not forming a central hyaline streak as in

pollinosa, and the second costal cell is uniformly dark; this cell in pollinosa is lighter brown in a narrow line along the posterior border (fig. 223, a). distribution. Map 28.

host.

E. pollinosa has been reared from galls on Artemisia tridentata.

Eutreta rotundipennis (Loew) (Figs. 226, 236, 238; Map 23)

Trypeta rotundipennis Loew 1862c: 79 (male holotype, MCZ; Tex.). — Loew 1873: 276 (review).

Eutreta rotundipennis: Loew 1873: 276, 329 (taxonomy). — Aldrich 1905: 608 (in catalog). — Crevecoeur 1906: 95 (Kans.). — V. T. Phillips 1923: 141 (review). — Curran 1932b: 16 (in key). — Brimley 1938: 384 (N. Car.). - Foote 1965a: 661 (in catalog). — Stoltzfus 1977: 396 (revision). recognition. The wings of

rotundipennis and fenestra are distinctive in their

broad width and by the large hyaline spots uniformly spaced along the anterior and

200

Systematic Treatment of the Genera

posterior margins (figs. 225, 226). In both species, many of the setae of the gena, postgena, and palpus are whitish and inflated (fig. 238), the oral margin is produced, and the oviscape is shorter than the two preceding abdominal tergites (fig. 236). In

rotundipennis, the hyaline wing spots in cell br and adjacent cells are distinct, whereas in fenestra they are coalesced. The apical hyaline wing crescent of rotun¬

dipennis is interrupted by an extension of the brown wing color along the apical veins to the wing margin (fig. 226, b), and the oral margin, postgena, pleuron, and coxae are tinged with violet. In fenestra, the hyaline crescent is complete, and there is no violet tinge on the oral margin, postgena, pleuron, or coxae. distribution. Map 23.

host. Not known.

E. rotundipennis and fenestra were placed by Stoltzfus (1977) in his subgenus Setosigena. discussion.

Eutreta simplex Thomas (Fig. 227; Map 25)

Eutreta simplex Thomas 1914: 425 (female holotype, MCZ; Sunset, Colo.). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 388 (revision). - Goeden 1990b: 33 (life history, host). recognition. The male of

simplex lacks the apical hyaline crescent, distinguish¬

ing it from all other Eutreta except males of decora. This crescent is present in the female but is small and does not extend anteriorly far enough to attain vein R2+3. The oral margin of E. simplex is at most weakly produced, and there are no dark spots on the face. E. intermedia differs in having the oral margin more strongly produced and the apical hyaline crescent well developed in both sexes, reaching anteriorly past vein R2+3. The hyaline wing spots of simplex (fig. 227) are much larger than those of decora (fig. 222). distribution. Map 25.

HOST. Goeden (1990b) reared

simplex from stem galls on Artemisia ludoviciana,

the only known host, in California. discussion. The life history and hosts of

simplex were recorded by Goeden

(1990b).

Genus Gonioxyna Hendel Gonioxyna Hendel 1927: 160 (type species, Gonioxyna magniceps Hendel 1927: 161, by original designation). - Foote and Blanc 1979: 163 (review, New World).

200

Systematic Treatment of the Genera

posterior margins (figs. 225, 226). In both species, many of the setae of the gena, postgena, and palpus are whitish and inflated (fig. 238), the oral margin is produced, and the oviscape is shorter than the two preceding abdominal tergites (fig. 236). In

rotundipennis, the hyaline wing spots in cell br and adjacent cells are distinct, whereas in fenestra they are coalesced. The apical hyaline wing crescent of rotun¬

dipennis is interrupted by an extension of the brown wing color along the apical veins to the wing margin (fig. 226, b), and the oral margin, postgena, pleuron, and coxae are tinged with violet. In fenestra, the hyaline crescent is complete, and there is no violet tinge on the oral margin, postgena, pleuron, or coxae. distribution. Map 23.

host. Not known.

E. rotundipennis and fenestra were placed by Stoltzfus (1977) in his subgenus Setosigena. discussion.

Eutreta simplex Thomas (Fig. 227; Map 25)

Eutreta simplex Thomas 1914: 425 (female holotype, MCZ; Sunset, Colo.). - Foote 1965a: 661 (in catalog). - Stoltzfus 1977: 388 (revision). - Goeden 1990b: 33 (life history, host). recognition. The male of

simplex lacks the apical hyaline crescent, distinguish¬

ing it from all other Eutreta except males of decora. This crescent is present in the female but is small and does not extend anteriorly far enough to attain vein R2+3. The oral margin of E. simplex is at most weakly produced, and there are no dark spots on the face. E. intermedia differs in having the oral margin more strongly produced and the apical hyaline crescent well developed in both sexes, reaching anteriorly past vein R2+3. The hyaline wing spots of simplex (fig. 227) are much larger than those of decora (fig. 222). distribution. Map 25.

HOST. Goeden (1990b) reared

simplex from stem galls on Artemisia ludoviciana,

the only known host, in California. discussion. The life history and hosts of

simplex were recorded by Goeden

(1990b).

Genus Gonioxyna Hendel Gonioxyna Hendel 1927: 160 (type species, Gonioxyna magniceps Hendel 1927: 161, by original designation). - Foote and Blanc 1979: 163 (review, New World).

201

Gonioxyna recognition. Closely allied to

Dioxyna, Oxyna, and Paroxyna, the distinctive

genus Gonioxyna includes species with a geniculate proboscis and several other characters found in those genera. The wing pattern has a large dark spot in the vicinity of the pterostigma in addition to smaller scattered spots. As in Oxyna and

Paroxyna, the aedeagus has small spines immediately basal to the distiphallus. All members of the genus may be recognized by the shape of the wing (fig. 244); the costa is bowed anteriorly between the base of the pterostigma and the apex of vein R2+3, which causes cell rl to be distinctly wider than cell r2+3 at the apex of the pterostigma. It is curious to note that only males have been found, suggesting the possibility that the bowed costa is a secondary sexual character and that the species now placed in Gonioxyna represent males of species belonging to other genera.

Map 29. Distribution of Gonioxyna fuscata, Gymnocarena diffusa, G. flava, and G. tricolor.

202

Systematic Treatment of the Genera

Figure 244. Right wing, Gonioxyna fuscata Foote.

discussion. Korneyev (1989) synonymized this genus with

Campiglossa Ron-

dani, a change made too recently for inclusion in this handbook. The sole U.S. species does not appear to belong to Campiglossa, however, because it has a setulose frons. The classification of this and the other two New World species (one Mexican, one Guatemalan) needs to be reevaluated. Individuals are rarely collected, and their habits and hosts are unknown. Other species in the genus were discussed by Foote and Blanc (1979).

Gonioxyna fuscata Foote (Figs. 91, 244; Map 29)

Gonioxyna fuscata Foote in Foote and Blanc 1979: 166 (male holotype, NMNH; Calif.). recognition. In contrast to the wing patterns of the other New World species of

Gonioxyna, that of fuscata is characterized by a very dark area in the anterior half of the disk not punctuated by hyaline spots (fig. 244). In this respect, it most resembles that of Paroxyna opacipennis (Foote) (fig. 320), but the latter has a different arrange¬ ment of hyaline spots in the wing apex, two distinct hyaline spots in cell

imme¬

diately distad of the pterostigma, and a completely dark cell dm. In fuscata, the posterior orbital bristle usually is absent. distribution. Map 29.

host. Not known. Adults have been collected from

Artemisia tridentata and

Cbrysotbamnus sp.

Genus Gymnocarena Hering Spilograpba Loew: Aldrich (part) 1905: 604 (in catalog). Euaresta Loew: Aldrich (part) 1905: 612 (in catalog). Oedicarena Loew: Curran 1934: 289 (in key). Gymnocarena Hering 1940a: 4 (type species, Oedicarena diffusa Snow 1894: 161, by original designation). - Foote 1960h: 112 (revision). - Foote 1965a: 675 (in catalog). - Wasbauer 1972: 117 (hosts). - Blanc and Foote 1987: 428 (revision).

203

Gymnocarena Gymnocarena, strikingly like Mylogymnocarena in many re¬ spects, comprises yellow-bodied, yellow-haired species in which the dorsocentral bristles are situated at about the level of the supra-alars (see fig. 23, b); the head in lateral view has a wide parafacial, receding face, swollen postgena, and hairless frons; and crossvein r-m is situated apicad of the middle of cell dm. From those of Mylogymnocarena, these species are distinguished by having cell with two hyaline spots apicad of the pterostigma and one in the outer half of cell r4+5, the background color of the wing completely or in part yellowish, crossveins r-m and dm-cu usually nearly parallel, and all of the femora distinctly enlarged. recognition.

The species of Gymnocarena generally occur below 3,000 feet east of the Continental Divide in the United States. The genus also is found in Mexico, where its exact distribution is not known. discussion.

Key to Known Species of Gymnocarena 1. Background color of wing uniformly light yellow, the hyaline markings sometimes difficult to distinguish; apical-most hyaline marks in cells 2+3 and r4 + 5 joined across vein R4 + 5 (fig. 245, a).diffusa (Snow) Apical half of wing dark brown, basal half faded brown or yellowish; apical-most hyaline marks in cells r2+3 and r4+5 separated, not at all joined across vein R4+5 (figs. 246, b; 247, b).2 2. Hyaline mark at apex of pterostigma ending above vein r-m (fig. 246, a); apical third of cell dm included in the dark brown area anterior to it (fig. 246, c); face concave with a rather sharp central carina. .tricolor (Doane) Hyaline mark at apex of pterostigma ending distinctly distad of anterior end of vein r-m (fig. 247, a); apical third of cell dm and posterior half of vein

Figures 245—247. Right wings, Gymnocarena spp. 245, diffusa (Snow); 246, tricolor (Doane); 247, flava Foote.

Systematic Treatment of the Genera r-m situated in a yellow area resembling that in basal half of wing (fig. 247, c); lower half of face swollen, in profile clearly visible beyond parafacial .flava Foote

Gymnocarena diffusa (Snow) (Figs. 63, 245; Map 29) Oedicarena diffusa Snow 1894: 161 (female lectotype, UKL; Kans.). - Coquillett 1899c: 261 (taxonomy). — Doane 1899: 179 (taxonomy). — F. H. Snow 1903: 219 (Kans.). — Curran 1934: 290 (figure, head). - Knowlton and Harmston 1937: 145 (Utah). - Byers et al. 1962: 180 (type data). - Beirne 1971: 60 (economic impor¬ tance). — Arnaud 1979: 330 (type data). Straussia diffusa: Coquillett 1899c:. 261 (taxonomy). — Essig 1938: 602 (note). Spilographa diffusa: Aldrich 1905: 604 (in catalog). - Washburn 1905: 19 (Minn.). Strauzia diffusia [error]: Cresson 1907: 100 (N. Mex.). Gymnocarena diffusa: Hering 1940a: 4 (type data). - Foote 1960h: 113 (revision). Foote 1962: 174 (type designation). - Foote 1965a: 676 (in catalog). - Lipp and Schulz 1970: 27 (damage). - Kamali and Schulz 1971: 85 (artificial diet). Wasbauer 1972: 117 (hosts). - Kamali and Schulz 1973: 288 (biology). - Kamali and Schulz 1974: 695 (immature stages). - Hilgendorf and Goeden 1981: 105 (host). — Blanc and Foote 1987: 428 (revision). recognition.

The wing pattern of diffusa is a very light yellow color, hence the

hyaline markings do not contrast well with it and are sometimes difficult to distin¬ guish. The hyaline spots in cells r2+3 and r4+5 are joined across vein R4+5 to form a continuous curved hyaline band (fig. 245, a) in contrast to the situation in flava and tricolor, in which these hyaline spots are completely separated. distribution.

host.

Map 29.

Helianthus annuus is the only known host.

discussion.

G. diffusa is by far the most common species of the genus. Lipp and

Schulz (1970) indicated that diffusa is of actual and potential economic importance on Helianthus, and Kamali and Schulz (1973) presented the results of a rather detailed investigation of the biology of this species in North Dakota. Kamali and Schulz (1974) described and illustrated the egg, larva, and puparium. Mature larvae appear to be unique among Tephritidae in showing heavy intersegmental pigmenta¬ tion.

Gymnocarena flava Foote (Fig. 247; Map 29) Gymnocarena flava Foote in Blanc and Foote 1987: 430 (female holotype, NMNH; Manson, Iowa).

205

Hexacbaeta The basal half of the wing disk is light yellow, whereas most of the apical half is dark brown with hyaline markings; the yellow area of the wing base extends apicad to include part of crossvein r-m and all of cell dm, including crossvein dm-cu. The hyaline mark at the apex of the pterostigma terminates apicad of the anterior end of crossvein r-m (fig. 247, a) in contrast to that of tricolor (fig. 246, a), recognition.

in which it terminates directly above crossvein r-m. The lower half of the face is somewhat swollen, enough so that it is plainly visible in lateral view beyond the parafacial. distribution.

host.

Map 29.

Not known.

Gymnocarena tricolor (Doane) (Fig. 246; Map 29) Euaresta tricolor Doane 1899: 191 (male lectotype, WSU; So. Dak.). - Aldrich 1905: 613 (in catalog). — Foote 1966b: 125 (type designation). — Zack 1984: 32 (type data). Tephritis tricolor: Coquillett 1899c: 264 (taxonomy). Gymnocarena tricolor: Quisenberry 1950: 10 (taxonomy). - Foote 1960h: 113 (review). - Foote 1965a: 676 (in catalog). - Blanc and Foote 1987: 429 (review). The hyaline mark at the apex of the pterostigma in tricolor termi¬ nates at the apex of crossvein r-m (fig. 246, a) rather than apicad of it as in flava (fig. 247, a). In tricolor, the apical third of cell dm and all of crossvein r-m lie in a uniform dark area. In lateral view, the face of tricolor is distinctly concave so that its lower half cannot be seen beyond the parafacial. recognition.

distribution.

host.

Not

Map 29.

known.

Genus Hexachaeta Loew Hexacbaeta Loew 1873: 219 (type species, Trypeta eximia Wiedemann 1830: 477, by designation of Coquillett 1910: 552). - Lima and Leite 1952: 297 (revision). Lima 1953a: 153 (Texas record). - Lima 1953b: 562 (Texas record, review). Lima 1954: 280 (Texas record, review). - Sabrosky 1967: 121 (addition, N.A. catalog). Hexacbaeta is unique among all genera occurring in America north of Mexico in having three pairs of scutellar bristles (see fig. 26). Individuals recognition.

205

Hexacbaeta The basal half of the wing disk is light yellow, whereas most of the apical half is dark brown with hyaline markings; the yellow area of the wing base extends apicad to include part of crossvein r-m and all of cell dm, including crossvein dm-cu. The hyaline mark at the apex of the pterostigma terminates apicad of the anterior end of crossvein r-m (fig. 247, a) in contrast to that of tricolor (fig. 246, a), recognition.

in which it terminates directly above crossvein r-m. The lower half of the face is somewhat swollen, enough so that it is plainly visible in lateral view beyond the parafacial. distribution.

host.

Map 29.

Not known.

Gymnocarena tricolor (Doane) (Fig. 246; Map 29) Euaresta tricolor Doane 1899: 191 (male lectotype, WSU; So. Dak.). - Aldrich 1905: 613 (in catalog). — Foote 1966b: 125 (type designation). — Zack 1984: 32 (type data). Tephritis tricolor: Coquillett 1899c: 264 (taxonomy). Gymnocarena tricolor: Quisenberry 1950: 10 (taxonomy). - Foote 1960h: 113 (review). - Foote 1965a: 676 (in catalog). - Blanc and Foote 1987: 429 (review). The hyaline mark at the apex of the pterostigma in tricolor termi¬ nates at the apex of crossvein r-m (fig. 246, a) rather than apicad of it as in flava (fig. 247, a). In tricolor, the apical third of cell dm and all of crossvein r-m lie in a uniform dark area. In lateral view, the face of tricolor is distinctly concave so that its lower half cannot be seen beyond the parafacial. recognition.

distribution.

host.

Not

Map 29.

known.

Genus Hexachaeta Loew Hexacbaeta Loew 1873: 219 (type species, Trypeta eximia Wiedemann 1830: 477, by designation of Coquillett 1910: 552). - Lima and Leite 1952: 297 (revision). Lima 1953a: 153 (Texas record). - Lima 1953b: 562 (Texas record, review). Lima 1954: 280 (Texas record, review). - Sabrosky 1967: 121 (addition, N.A. catalog). Hexacbaeta is unique among all genera occurring in America north of Mexico in having three pairs of scutellar bristles (see fig. 26). Individuals recognition.

Systematic Treatment of the Genera

somewhat resemble those of the larger species of Anastrepha in having a yellowish body and prominent dark bands on a clear wing field, but most Hexachaeta species have prominently dark or black leg, thoracic, and abdominal markings, and vein M is not curved forward at its apex. distribution.

The genus Hexachaeta, comprising about 25 species, is restricted

to the New World. It is frequently found in McPhail traps baited for Anastrepha all the way from Mexico to Brazil. Our records show the U.S. distribution to be con¬ fined to the Rio Grande Valley of Texas, but the species have not been seen there since about 1940 despite intensive trapping for Anastrepha species. discussion.

The genus consists of three species groups that are well defined by

wing pattern and male and female genitalic characters. It is remarkable that all three of these groups are represented in U.S. records, each by one species. The works by Lima and Leite cited above constitute virtually a complete revision of the genus. Four species have been reported to breed in the fruit of Sorocea and/or Helicostylis (Moraceae); one of these, eximia (Wiedemann), also has been bred from the fruit of Aegiphila (Verbenaceae) (Lima 1935, Lima and Leite 1952, d’Araujo e Silva et al. 1968). To this we may add data for valida Lima, reared from fruit of Sorocea affinis in Panama (data from NMNH specimens), and for fallax, reared from twigs of a Morus species.

Figures 248—250. Right wings, Hexachaeta spp. 248, obscura Hend.; 249, fallax Lima; 250, seabrai Lima.

Key to U.S. Species of Hexachaeta 1. Apical third of wing with only 1 dark transverse band, which covers vein dm-cu (fig. 248, b), and is separated from the basal dark area by an

Hexachaeta uninterrupted hyaline band between rm and dm-cu (fig. 248, a) . .obscura Hendel Apical third of wing with 2 transverse dark bands, the one covering vein dm-cu connected with the basal dark area in cell dm (figs. 249, b; 250, b)

.2 2. One triangular spot at anterior wing margin distad of pterostigma (fig. 249, a) .fallax Lima Two triangular hyaline spots at anterior wing margin distad of pterostigma (fig. 250, a) .seabrai Lima

Hexachaeta fallax Lima (Fig. 249; Map 30) Hexachaeta fallax Lima 1954: 280 (female holotype, NMNH; Cano Saddle, Gatun Lake, Pan.). - Foote 1967a: 1327 (Tex., taxonomy). - Sabrosky 1967: 121, 123 (addition, N.A. catalog). Hexachaeta n. sp.: Lima 1954: 280 (Texas record). Of the three species of Hexachaeta treated here, fallax is the only one with a single, deep triangular hyaline incision at the costa immediately apicad of the pterostigma (fig. 249, a), combined with the presence of two narrow dark bands in the apical third of the wing. Two hyaline areas also interrupt the anterior margin of the wing distad of the pterostigma in obscura, but in that species, the more apical hyaline mark is a band completely crossing the wing, and only one dark band is present in the apical third of the wing. The aculeus tip is complex with deep notches recognition.

and serrations. distribution.

Map 30. H. fallax is known only from Panama, Mexico, and

Texas. This is the species whose aculeus tip was figured by Baker et al. (1944: 124, fig. 69b). They reported it to damage mulberry (Morus) twigs in Morelos, Mexico. Specimens (with puparia) from that study are deposited in the NMNH. host.

Several other Neotropical species with similar wing patterns and genitalia are known, but none occurs north of Central America. discussion.

Hexachaeta obscura Hendel (Fig. 248; Map 30) Hexachaeta amabilis var. obscura Hendel 1914c: 25 (male syntypes, SMT, NMW; Urubamba R., Peru, and San Bernardino, Paraguay). Hexachaeta obscura: Lima 1953a: 153 (Texas record). - Foote 1967a: 1327 (Tex., taxonomy). — Sabrosky 1967: 121 (addition, N.A. catalog). — Hardy 1968: 118 (type data).

208

Systematic Treatment of the Genera

▲

Hexachaeta fallax

A

Hexachaeta obscura

■

Hexachaeta seabrai

•

Icterica circinata

★

Icterica circinata (state record only)

o

Icterica seriata

☆

Icterica seriata (state record only)

Map 30. Distribution of Hexachaeta fallax, H. obscura, H. seabrai, Icterica circinata, and I. seriata.

Among the three species of Hexachaeta discussed here, the wing pattern of obscura is the only one showing the combination of the dark narrow recognition.

subapical band (fig. 248, b), an even-sided hyaline band extending completely across the middle of the wing (fig. 248, a), and a single hyaline triangle immediately apicad of the pterostigma. The aculeus tip has deep notches like fallax but lacks the serra¬ tions found in the latter species. distribution.

Map 30. H. obscura is apparently widespread throughout the

tropical and subtropical areas of the New World and occurs from Texas south to Brazil.

209

Icterica host. Not known. discussion. A thorough study of series of specimens of obscura from representa¬ tive locations throughout its range may show that several distinct species are in¬ cluded under this name.

Hexachaeta seabrai Lima (Figs. 26, 250; Map 30) Hexachaeta seabrai Lima 1953b: 562 (female holotype, IOC; Parque Sooretama, Linhares, Espirito Santo, Brazil). - Foote 1967a: 1327 (Tex., taxonomy). Sabrosky 1967: 121, 123 (additions, N.A. catalog). recognition. In contrast to the wing patterns of the preceding two species, that

of seabrai has two relatively narrow hyaline triangles along the costa immediately distad of the pterostigma (fig. 250, a). As in fallax, there are two dark stripes in the distal third of the wing, but no even-sided hyaline band extends across the wing as in obscura. The aculeus tip is relatively simple, with the margin entire. distribution. Map 30. The distribution of seabrai south of the United States is

not well known; it also has been collected from Guatemala, Panama, and Brazil. host. Not known. discussion. H. seabrai is one of several Neotropical species with similar wing

patterns and genitalia, but as far as known, all of the others occur south of Central America.

Genus Icterica Loew Icterica Loew 1873: 287 (type species, Trypeta seriata Loew 1862c: 84, by designa¬ tion of Coquillett 1910: 555). — Loew 1873: 328 (in key). - Coquillett 1899c: 268 (in key). - C. F. Adams 1904: 449 (key to species). - Aldrich 1905: 610 (in catalog). - Williston 1908: 287 (in key). - Coquillett 1910: 555 (type species designation). - Hendel 1914b: 95 (in key). - Phillips 1923: 123 (in key). - V. T. Phillips 1923: 144 (review). - Curran 1932b: 3 (in key). - Curran 1934: 287 (in key). - Steyskal and Foote 1963: 166 (taxonomy). - R.H. Foote 1965a: 664 (in catalog). - B.A. Foote 1967: 1296 (biology). — Wasbauer 1972: 117 (hosts). Foote and Freidberg 1980: 31 (type data). Trypeta (Icterica): Osten Sacken 1878: 193 (in catalog). recognition. The two North American species are modest-sized flies with

largely unmarked yellow-brown bodies and legs; yellow to brown body and leg

209

Icterica host. Not known. discussion. A thorough study of series of specimens of obscura from representa¬ tive locations throughout its range may show that several distinct species are in¬ cluded under this name.

Hexachaeta seabrai Lima (Figs. 26, 250; Map 30) Hexachaeta seabrai Lima 1953b: 562 (female holotype, IOC; Parque Sooretama, Linhares, Espirito Santo, Brazil). - Foote 1967a: 1327 (Tex., taxonomy). Sabrosky 1967: 121, 123 (additions, N.A. catalog). recognition. In contrast to the wing patterns of the preceding two species, that

of seabrai has two relatively narrow hyaline triangles along the costa immediately distad of the pterostigma (fig. 250, a). As in fallax, there are two dark stripes in the distal third of the wing, but no even-sided hyaline band extends across the wing as in obscura. The aculeus tip is relatively simple, with the margin entire. distribution. Map 30. The distribution of seabrai south of the United States is

not well known; it also has been collected from Guatemala, Panama, and Brazil. host. Not known. discussion. H. seabrai is one of several Neotropical species with similar wing

patterns and genitalia, but as far as known, all of the others occur south of Central America.

Genus Icterica Loew Icterica Loew 1873: 287 (type species, Trypeta seriata Loew 1862c: 84, by designa¬ tion of Coquillett 1910: 555). — Loew 1873: 328 (in key). - Coquillett 1899c: 268 (in key). - C. F. Adams 1904: 449 (key to species). - Aldrich 1905: 610 (in catalog). - Williston 1908: 287 (in key). - Coquillett 1910: 555 (type species designation). - Hendel 1914b: 95 (in key). - Phillips 1923: 123 (in key). - V. T. Phillips 1923: 144 (review). - Curran 1932b: 3 (in key). - Curran 1934: 287 (in key). - Steyskal and Foote 1963: 166 (taxonomy). - R.H. Foote 1965a: 664 (in catalog). - B.A. Foote 1967: 1296 (biology). — Wasbauer 1972: 117 (hosts). Foote and Freidberg 1980: 31 (type data). Trypeta (Icterica): Osten Sacken 1878: 193 (in catalog). recognition. The two North American species are modest-sized flies with

largely unmarked yellow-brown bodies and legs; yellow to brown body and leg

Systematic Treatment of the Genera bristles; two to four frontal bristles, two reclinate orbital bristles, two pairs of scutellar bristles; the single pair of dorsocentral bristles in a transverse line through the supra-alars; frons with fine setae; vein r-m distinctly distad of the midpoint of the cell dm; a flat scutellum; and the border of the wing distinctly brown with hyaline spots, the disk with a central area of massively fused yellowish spots, making that area appear as if covered by irregular-shaped darker brown spots against a yellowish background. This type of wing pattern is unique in the Tephritidae and distinguishes the species of this genus from any others occurring in Canada and the United States. discussion. Korneyev (1989) transferred the Palearctic species previously placed in lcterica to Orotava Frey, lcterica now includes only the two North American species discussed here. Their relationships are not well understood. Both species breed in flowers of Bidens. As far as known, lcterica does not occur south of the United States, although several Neotropical species previously have been misplaced in the genus (Foote 1980).

Key to North American Species of lcterica Base and apex of pterostigma yellow, center with a dark spot (fig. .circinata Pterostigma darkened along almost its entire posterior border (fig. .seriata

251, a) (Loew) 252, a) (Loew)

lcterica circinata (Loew) (Fig. 251; Map 30) Trypeta circinata Loew 1873: 288 (male and female syntypes, MCZ; N.Y.). lcterica circinata: Loew 1873: 330 (taxonomy). - Johnson 1900b: 688 (N.J.). - C. F. Adams 1904: 449 (in key). - Aldrich 1905: 610 (in catalog). - G. M. Greene 1909: 443 (N.J.). - Johnson 1910: 802 (N.J.). - V. T. Phillips 1923: 145 (review). Leonard 1928: 853 (N.Y.). - Curran 1934: 288 (figure wing). - Steyskal and Foote 1963: 166 (taxonomy). - R.H. Foote 1965a: 664 (in catalog). - B.A. Foote 1967: 1299 (biology, description of immature stages). - Novak et al. 1967: 147 (host). Wasbauer 1972: 117 (host). Trypeta (lcterica) circinata: Osten Sacken 1878: 193 (in catalog).

a

a

Figures 251, 252. Right wings, lcterica spp. 251, circinata (Lw.); 252, seriata (Lw.).'

211

Jamesomyia I. circinata may be recognized by the markings within the pterostigma, which consist essentially of a yellowish area on each end of a central dark spot (fig. 251, a). In seriata, by contrast, at least one of the yellowish areas does not extend completely across the pterostigma, leaving vein Rt almost completely bordered with dark brown (fig. 252, a). The border of the wing in circinata is not as dark as in seriata, and the four hyaline spots in cell rl along the costa apicad of the pterostigma are much more discrete and distinct. recognition.

distribution.

host.

Map 30.

Bidens cernua was the only host listed by Wasbauer (1972).

Icterica seriata (Loew) (Figs. 58, 71, 252; Map 30) Trypeta seriata Loew 1862c: 84 (male syntype, MCZ; “Middle States”). — Loew 1873: 287 (review). Icterica seriata: Loew 1873: 287, 330 (taxonomy). - W. A. Snow 1894: 171 (taxon¬ omy). — Doane 1899: 188 (distribution).-Johnson 1900b: 688 (N.J.). — Chagnon 1901: 14 (Can.). — C. F. Adams 1904: 449 (in key). — Aldrich 1905: 610 (in catalog). — Greene 1909: 443 (N.J.). — Coquillett 1910: 555 (type designation). — Johnson 1910: 802 (N.J.). — Flendel 1914b: 95 (type data). - Winn and Beaulieu 1915: 154 (Que.). — Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 145 (re¬ view). — Johnson 1925b: 264 (N. Engl.). — Petch and Maltaise 1932: 57 (Que.). Steyskal and Foote 1963: 166 (taxonomy). - R.H. Foote 1965a: 664 (in catalog). B.A. Foote 1967: 1298 (biology, hosts, description immature stages). - Novak et al. 1967: 147 (hosts). — Wasbauer 1972: 117 (hosts). — Freidberg 1981: 93 (review, mating behavior). Trypeta (Icterica) seriata: Osten Sacken 1878: 193 (in catalog). Icterica sericata [error]: Curran 1934: 29 (figure of head). recognition.

See recognition characters for circinata.

distribution.

Map 30.

hosts.

I. seriata is known only from several species of Bidens.

Genus Jamesomyia Quisenberry Trypeta (Tepbritis) (part): Osten Sacken 1878: 193 (in catalog). Tephritis (part): Aldrich 1905: 611 (in catalog). Euribia (part): V. T. Phillips 1923: 123 (in key); 149 (review). Jamesomyia Quisenberry 1949b: 49 (type species, Trypeta geminata Loew 1862a: 220 (1864: 90), by original designation). — Foote 1965a: 663 (in catalog). — Wasbauer 1972: 117 (hosts).

211

Jamesomyia I. circinata may be recognized by the markings within the pterostigma, which consist essentially of a yellowish area on each end of a central dark spot (fig. 251, a). In seriata, by contrast, at least one of the yellowish areas does not extend completely across the pterostigma, leaving vein Rt almost completely bordered with dark brown (fig. 252, a). The border of the wing in circinata is not as dark as in seriata, and the four hyaline spots in cell rl along the costa apicad of the pterostigma are much more discrete and distinct. recognition.

distribution.

host.

Map 30.

Bidens cernua was the only host listed by Wasbauer (1972).

Icterica seriata (Loew) (Figs. 58, 71, 252; Map 30) Trypeta seriata Loew 1862c: 84 (male syntype, MCZ; “Middle States”). — Loew 1873: 287 (review). Icterica seriata: Loew 1873: 287, 330 (taxonomy). - W. A. Snow 1894: 171 (taxon¬ omy). — Doane 1899: 188 (distribution).-Johnson 1900b: 688 (N.J.). — Chagnon 1901: 14 (Can.). — C. F. Adams 1904: 449 (in key). — Aldrich 1905: 610 (in catalog). — Greene 1909: 443 (N.J.). — Coquillett 1910: 555 (type designation). — Johnson 1910: 802 (N.J.). — Flendel 1914b: 95 (type data). - Winn and Beaulieu 1915: 154 (Que.). — Britton 1920: 204 (Conn.). — V. T. Phillips 1923: 145 (re¬ view). — Johnson 1925b: 264 (N. Engl.). — Petch and Maltaise 1932: 57 (Que.). Steyskal and Foote 1963: 166 (taxonomy). - R.H. Foote 1965a: 664 (in catalog). B.A. Foote 1967: 1298 (biology, hosts, description immature stages). - Novak et al. 1967: 147 (hosts). — Wasbauer 1972: 117 (hosts). — Freidberg 1981: 93 (review, mating behavior). Trypeta (Icterica) seriata: Osten Sacken 1878: 193 (in catalog). Icterica sericata [error]: Curran 1934: 29 (figure of head). recognition.

See recognition characters for circinata.

distribution.

Map 30.

hosts.

I. seriata is known only from several species of Bidens.

Genus Jamesomyia Quisenberry Trypeta (Tepbritis) (part): Osten Sacken 1878: 193 (in catalog). Tephritis (part): Aldrich 1905: 611 (in catalog). Euribia (part): V. T. Phillips 1923: 123 (in key); 149 (review). Jamesomyia Quisenberry 1949b: 49 (type species, Trypeta geminata Loew 1862a: 220 (1864: 90), by original designation). — Foote 1965a: 663 (in catalog). — Wasbauer 1972: 117 (hosts).

212

Systematic Treatment of the Genera

Figure 253. Right wing, Jamesomyia geminata (Lw.).

recognition.

The basal one-fifth of the wing (fig. 253) in this monotypic genus

is hyaline, the remainder dark and peppered with indistinctly bordered hyaline spots somewhat reminiscent of Eurosta and Eutreta. The genus most closely resembles Xenochaeta, from which it differs by the presence of four pairs of light-colored frontal bristles, the wing pattern, and other characters found in the key. All the head hairs are light-colored. A parafacial spot is present. The dorsocentral bristles are situated almost exactly in a line between the supra-alar bristles, and together with the other scutal bristles, they arise out of small dark spots that contrast markedly with the lighter, brownish scutal tomentum. jamesomyia resembles Xenochaeta and is superficially similar to Xanthomyia. We have placed it in the tephritine tribe Eutretini, which contains Eutreta and seven other Nearctic genera. discussion,

Jamesomyia geminata (Loew) (Figs. 73, 76, 253; Map 31) Trypeta geminata Loew 1862a: 220 (1864: 90) (female syntypes, originally NMW, probably lost; Pa.). - Loew 1873: 298 (review). Tephritis geminata: Loew 1873: 300, 330 (taxonomy). - Johnson 1900b: 688 (N.J.). -Aldrich 1905: 612 (in catalog). - Johnson 1910: 802 (N.J.). - Johnson 1925b: 264 (N. Engl.). — Hering 1944b: 17 (taxonomy). Trypeta (Tephritis) geminata: Osten Sacken 1878: 193 (in catalog). Euribia geminata: V. T. Phillips 1923: 150 (review). Tephritis geminatus: Petch and Maltaise 1932: 57 (Que.). Jamesomyia geminata: Quisenberry 1949b: 50 (review). - Quisenberry 1951: 60 (taxonomy). - Foote 1965a: 633 (in catalog). - Novak et al. 1967: 147 (host). Wasbauer 1972: 117 (host). recognition.

As for the genus.

DISTRIBUTION.

Map 31.

host.

Novak et al. (1967) bred this species from flowers of Lactuca canadensis.

No recent work has been done on any aspects of geminata. It is rarely found in general collecting and is rare in fruit fly collections. discussion.

213

Metatephritis

Map 31. Distribution of Jamesomyia geminata, Metatephritis fenestrata, Mylogymnocarena apicata, and M. bicolor.

Genus Metatephritis Foote Metatephritis Foote 1960h: 110 (type species, Metatephritis fenestrata Foote 1960h: 110, by original designation). — Foote 1965a: 669 (in catalog). — Wasbauer 1972: 118 (hosts). — Freidberg 1984: 129 (gall biology). - Blanc and Foote 1987: 431 (taxonomy). Metatephritis is a curious anomaly among North American gen¬ era. Structurally, it resembles Neotephritis and Euarestoides in having three pairs of frontal and two pairs of scutellar bristles, but its wing pattern, instead of numerous recognition.

213

Metatephritis

Map 31. Distribution of Jamesomyia geminata, Metatephritis fenestrata, Mylogymnocarena apicata, and M. bicolor.

Genus Metatephritis Foote Metatephritis Foote 1960h: 110 (type species, Metatephritis fenestrata Foote 1960h: 110, by original designation). — Foote 1965a: 669 (in catalog). — Wasbauer 1972: 118 (hosts). — Freidberg 1984: 129 (gall biology). - Blanc and Foote 1987: 431 (taxonomy). Metatephritis is a curious anomaly among North American gen¬ era. Structurally, it resembles Neotephritis and Euarestoides in having three pairs of frontal and two pairs of scutellar bristles, but its wing pattern, instead of numerous recognition.

214

Systematic Treatment of the Genera

Figure 254. Right wing, Metatephritis fenestrata Foote.

hyaline spots in a dark field as in those two genera, consists of a large median hyaline area surrounded by a dark border (fig. 254). discussion.

Metatephritis may be the senior synonym of Eutreta subgenus Un-

caculeus. For the present, its status as a separate genus rests principally upon its different wing pattern, which includes an apical hyaline band but lacks the small hyaline spots found in Eutreta. Discovery of the female of fenestrata would help to confirm this possible relationship.

Metatephritis fenestrata Foote (Fig. 254; Map 31) Metatephritis fenestrata Foote 1960h: 110 (male holotype, NMNH; Cody, Wyo.). — Fronk et al. 1964: 575 (galls, hosts, associates). — Foote 1965a: 669 (in catalog). — Wasbauer 1972: 118 (hosts). — Steck 1981: 18 (host). — Blanc and Foote 1987: 431 (taxonomy). recognition.

The recognition characters of fenestrata are as discussed under the

genus. In one specimen from California, two hyaline spots are present in addition to those shown in fig. 254. One of these occurs near the center of the anal lobe contiguous with the wing margin, the other is centered on the posterior margin of cell cual between the termination of vein CuA! and the next proximal hyaline area. DISTRIBUTION.

Map 31.

M. fenestrata arbuscula ssp. nova. host.

forms galls on the vegetative tips of new growth on

Artemisia

M. fenestrata is extremely rare; it is known at present by only three specimens from Wyoming and one from California. discussion.

Genus Mylogymnocarena Foote Mylogymnocarena Foote 1960h: 111 (type species, Urellia apicata Thomas 1914: 428, by original designation). - Foote 1965a: 669 (in catalog). - Blanc and Foote 1987: 430 (revision).

215

Mylogymnocarena Strikingly similar to those of Gymnocarena, the species of My¬ logymnocarena are yellow-bodied and yellow-haired and have heads of similar shape in profile (see discussion of Gymnocarena for details). In contrast to the former genus, members of Mylogymnocarena have a uniform brown wing pattern, some¬ times fading to hyaline basally; only one hyaline area apicad of the pterostigma in cell r^ crossveins r-m and dm-cu lying at an angle to each other so that the posterior extensions of each converge at or near the posterior wing margin; and unenlarged femora. Cell r4 + 5 is hyaline only at or near its base. RECOGNITION.

The two species of Mylogymnocarena are restricted to relatively high altitudes in the western mountain chains of the North American continent (see Map 31). The species are relatively rare and are poorly represented in collections. In addition to Gymnocarena mexicana Aczel, it is quite likely that species occur in Mexico and elsewhere south of the United States. Virtually nothing is known about the biology of the two known species. discussion.

Key to Known Species of Mylogymnocarena Dark pattern of wing confined to its apical third except for a slight yellow¬ ing of pterostigma; cell r2 + 3 with 2 hyaline spots apicad of terminus of vein R2+3; at least 3 hyaline spots present in cell m (fig. 255, a) . .apicata (Thomas) Apical 2A of wing, including part of pterostigma, mostly dark; cell r2+3 completely dark apicad of terminus of vein R2+3; cell m with 1 large hyaline spot sometimes containing a small dark area at or near its center (fig. 256, a) .bicolor (Foote)

Mylogymnocarena apicata (Thomas) (Fig. 255; Map 31) Urellia apicata Thomas 1914: 428 (female holotype, MCZ; Colo.). Mylogymnocarena apicata: Foote 1960h: 111 (review). - Foote 1965a: 669 (in catalog). — Blanc and Foote 1987: 431 (revision).

a Figures 255, 256. Right wings, Mylogymnocarena spp. 255, apicata (Thomas); 256, bicolor (Foote).

216

Systematic Treatment of the Genera recognition.

At first glance, the wing pattern of apicata roughly resembles that

of several species of Trupanea in being hyaline except for the pterostigma and a large apical dark area, but the body size and color, the shape of the head in profile, the number of scutellar setae, and the location of the dorsocentral bristles readily distin¬ guish these two genera. M. apicata is distinguished from bicolor by its distinctive wing pattern (compare figs. 255 and 256). In addition to the three hyaline spots shown in fig. 255, a very small fourth one also may be present. Map 31. M. apicata has been collected in Durango, but its dis¬

distribution.

tribution in Mexico is otherwise uncertain. host.

Not

known.

Mylogymnocarena bicolor (Foote) (Fig. 256; Map 31) Gymnocarena bicolor Foote 1960h: 113 (holotype male, NMNH; Indian Creek Canyon, Chiricahua Mts., Ariz.). — Foote 1965a: 676 (in catalog). Mylogymnocarena bicolor: Blanc and Foote 1987: 431 (revision). recognition.

In contrast to the wing pattern of apicata, in which the brown area

occupies only the apical third of an otherwise hyaline wing (fig. 255), that of bicolor comprises a largely dark wing with contrasting hyaline areas incised into it (fig. 256). Cell m sometimes lacks the small dark spot in the center of the large hyaline area (fig. 256, a). distribution.

Map 31. M. bicolor also has been collected from Sinaloa, but its

distribution in Mexico is otherwise unknown. host.

Not

known.

Genus Myoleja Rondani Myoleja Rondani 1856: 112 (type species, Tephritis lucida Fallen 1826: 13, by original designation). — Coquillett 1910: 573 (type data). — F. H. Benjamin 1934: 9 (review, Fla.). — Curran (part) 1934: 289 (in key). — Blanc and Foote 1961: 75 (key to species). — Foote and Blanc 1963: 32 (review, Calif.). — Foote 1965a: 669 (in catalog). — Steyskal 1972b: 207 (key to species, taxonomy). — Wasbauer 1972: 118 (hosts). — Maddison and Bartlett 1989: 32 (zoogeography). Aciura Robineau-Desvoidy: Coquillett (part) 1899c: 263 (taxonomy). — V. T. Phil¬ lips (part) 1923: 121, 130 (in key, review). Aciura (Eucosmoptera): V. T. Phillips (part) 1923: 131 (key to subgenera, e. U.S.; see Xantbaciura).

217

My olej a Eucosmoptera Phillips: Curran (part) 1932b: 12 (in key, key to species). - Curran (part) 1934: 297 (in key). Euleia Walker: Bates 1933a: 50 (review). Pseudeuleia Hering 1940b: 16 (type species, Aciura limata Coquillett 1899c: 263, by original designation). Small to moderate-sized flies with heavily marked wings, yellow to dark brown bodies, and, in some species, greatly enlarged frontal bristles in the male. The face is receding and without a projecting oral margin; the genae lack elongated setae in addition to the genal bristle; all of the postocular bristles are brown to black; there are three pairs of frontal bristles; two pairs of orbitals, the posterior pair reclinate; two pairs of scutellar bristles on a flat, usually yellowish scutellum; and one pair of dorsocentral bristles situated at or behind a transverse line through the supra-alars, sometimes closer to the acrostichals than to the supra-alars. Vein r-m is close to vein dm-cu, distinctly apicad of the middle of cell dm, and cell cup is drawn out to an elongated point. The characters given in the key and the heavily marked wing (figs. 257-260) distinguish this genus from all others occurring in the United States and Canada. recognition.

Myoleja, a large, nearly cosmopolitan genus, is poorly delimited from a number of related genera in the Trypetinae. Fourteen Palearctic and more than 30 Oriental species of Myoleja have been described. In the New World, Myoleja occurs in the West Indies and Brazil, suggesting that it is distributed throughout much of the Neotropical Region. Although only limata has been reared (from the fruit of several Ilex species), the shape of the aculeus tip suggests that at least nigricornis and rhino are also frugivorous. discussion.

d

c

c

b

Figures 257—260. Right wings, Myoleja spp. 257, unifasciata Blanc and Foote; 258, nigricornis (Doane); 259, limata (Coq.); 260, rhino Steysk.

Systematic Treatment of the Genera

Figures 261-264. Lateral view, head, Myoleja spp. 261, unifasciata Blanc and Foote; 262, nigricornis (Doane), male; 263, limata (Coq.); 264, rhino Steysk., male.

Key to U.S. Species of Myoleja 1. One hyaline triangle in cell r, immediately distad of pterostigma (fig. 257, b); cell c with dark brown medial crossband (fig. 257, a); cell dm wholly dark brown (fig. 257, c); frontal bristles of male not enlarged (fig. 261, a)

.unifasciata Blanc and Foote Two hyaline incisions in cell r? distad of pterostigma (figs. 258, b; 259, a; 260, a); cell c medially hyaline, brown basally and apically (fig. 258, a); cell dm with preapical hyaline spot or crossband (figs. 258, e; 259, c; 260, b); frontal bristles of males sometimes greatly enlarged (figs. 262, a; ^64, a) ..

219

Myoleja 2. Apical fourth of wing dark brown, with 1 hyaline triangle in cell m (fig. 258, d); cell r4+5 dark brown except for round hyaline spot near vein r-m (fig. 258, c); cell cuaj with 2 hyaline areas (fig. 258, f); abdomen largely blackish; male with 2 or 3 greatly enlarged frontal bristles (fig. 262, a) ...nigricornis (Doane) Apical fourth of wing with 2 hyaline marks, at least 1 of which crosses cell r4+5 (fig- 2-59, b); cell cua, with only 1 hyaline subbasal area (figs. 259, d; 260, c); abdomen blackish or yellowish; male with frontal bristles en¬ larged or not .3 3. Second hyaline mark crossing cell r, distad of pterostigma separate from or (rarely) weakly connected to hyaline spot(s) in apex of cell dm (fig. 259, c); ocellar bristles weak, shorter than postocular setae; abdomen with at least tergites 4 and 5 usually brown; male with 3 frontal bristles, not enlarged (fig. 263, a) .limata (Coquillett) Second hyaline mark crossing cell rl distad of pterostigma broadly crossing cell dm subapically as an uninterrupted band (fig. 260, b); ocellar bristles longer than postocular setae; abdominal tergites yellowish (if dark, from internal dark material); male with 2 frontal bristles, greatly enlarged (fig. 264, a) .rhino Steyskal

Myoleja limata (Coquillett) (Figs. 66, 259, 263; Map 32) Aciura limata Coquillett 1899c: 263 (female holotype, NMNH; New Bedford, Mass.). — Aldrich 1905: 607 (in catalog). — Britton 1920: 204 (Conn.). — Bates 1934a: 295 (taxonomy). — Hennig 1952: 220 (in catalog immature stages). — Fischer 1964: 272 (parasite). Aciura (Eucosmoptera) limata: V. T. Phillips 1923: 132 (review). — Johnson 1925b: 261 (N. Engl.). Eucosmoptera limata: Curran 1932b: 12 (in key). Euleia limata: Bates 1933a: 51 (taxonomy). Myoleja limata: F. H. Benjamin 1934: 21 (review, Fla.). - V. T. Phillips 1946: 50, 115 (description, biology larva; hosts). — Foote 1965a: 669 (in catalog). — Steyskal 1972b: 209 (taxonomy). — Wasbauer 1972: 118 (hosts). — Wharton and Marsh 1978: 149 (parasite). Pseudeuleia limata: Hering 1940b: 16 (type data). The dorsocentral bristles of limata are situated quite far behind a transverse line through the supra-alars; approximately the anterior third of the tergum is yellowish, darkening to reddish brown in the posterior two-thirds; and the frontal bristles of the male are not at all enlarged. Cell c in limata is largely hyaline, without a dark median transverse bar; two hyaline triangles are present immediately distad of the pterostigma in cells rt and r2+3; two slender hyaline marks also are present in the apical third of the wing disk (fig. 259, b); and the apical half of the pattern, except for the margins, is lighter than the basal half. This combination of characters distinguishes limata from all the other North American species of Myolerecognition.

220

Systematic Treatment of the Genera

Map 32. Distribution of Myoleja limata, M. nigricornis, M. rhino, and M. unifasciata.

ja. M. limata is the only Nearctic Myoleja with short, weak ocellar setae. The aculeus tip is slender, but it is not flattened in the sagittal plane. distribution.

Map 32.

A number of species of Ilex have been recorded as hosts. The larvae breed in the fruit. HOSTS.

Myoleja nigricornis (Doane) (Figs. 258, 262; Map 32) Aciura nigricornis Doane 1899: 183 (holotype (sex not known), WSU; Pa.). Johannsen 1903: 16 (description). - Aldrich 1905: 607 (in catalog). - Johnson 1909: 113 (distribution). - Johannsen 1926: 159 (N.Y.). - Johnson 1927: 217

221

Myoleja (Mt. Desert). - Leonard 1928: 851 (N.Y.). - Bates 1934a: 295 (taxonomy). Hering 1940b: 16 (taxonomy). - Foote 1966b: 123 (type data). - Zack 1984: 31 (type data). Aciura (Eucosmoptera) nigricornis: V. T. Phillips 1923: 131 (review). - Johnson 1925b: 261 (N. Engl.). Eucosmoptera nigricornis: Curran 1932b: 12 (in key). Euleia nigricornis: Bates 1933a: 51 (taxonomy). Myoleja nigricornis: Ouellet 1941: 130 (Que.). - Foote 1965a: 669 (in catalog). Steyskal 1972b: 209 (taxonomy). M. nigricornis is the only species of Myoleja found in the United States and Canada having the following combination of characters: the frontal bris. ties of the male are very black and enlarged, two hyaline triangles are present in cells r1 and r2+3 directly distad of the pterostigma, a rounded hyaline spot is present in cell dm posterior to vein r-m, cell r4+5 is dark except for a hyaline spot more or less in line with vein dm-cu, and cell c is dark only at its proximal and distal ends. The aculeus tip is flattened in the sagittal plane as in rhino. recognition.

distribution.

host.

Not

Map 32.

known.

Myoleja rhino Steyskal (Figs. 260, 264; Map 32) Myoleja rhino Steyskal 1972b: 207 (male holotype, NMNH; Fla.). A most distinctive species in wing pattern (fig. 260), rhino has a single hyaline triangle immediately distad of the pterostigma; in addition, a long, narrow, even-sided hyaline stripe extends from the costa transversely across the wing into cell dm between veins r-m and dm-cu (fig. 260, b). A relatively large hyaline spot is present in cell br directly behind the pterostigma. These characters, together with the strongly receding face and extremely large frontal bristles of the male (fig. 264, a), distinguish this species from all others occurring in the United States and Canada. The aculeus tip is flattened in the sagittal plane as in nigricornis. recognition.

distribution.

host.

Not

Map 32.

known.

Myoleja unifasciata Blanc and Foote (Figs. 257, 261; Map 32) Myoleja unifasciata Blanc and Foote 1961: 73 (male holotype, NMNH; Calif.). — Foote and Blanc 1963: 32 (review, Calif.). - Foote 1965a: 669 (in catalog). Steyskal 1972b: 209 (taxonomy). — Arnaud 1979: 330 (type data).

222

Systematic Treatment of the Genera recognition.

In unifasciata, the dorsocentral bristles lie only slightly behind a

transverse line between the supra-alars, and the frontal bristles of the male are not enlarged. The largely hyaline cell c is divided at the center by a transverse brown band, only one hyaline triangle is present in cell r1 immediately distad of the pterostigma, and cell dm is entirely dark without any hyaline markings. This com¬ bination of characters distinguishes unifasciata from all other Myoleja species dis¬ cussed in this handbook. The female is still unknown. distribution.

host.

Not

Map 32.

known.

Genus Neaspilota Osten Sacken Aspilota Loew 1873: 286 (type species, Trypeta alba Loew 1861a: 345, by subse¬ quent designation of Coquillett 1910: 511) (preoccupied Forster, 1862). - Osten Sacken 1878: 260 (nomenclature). — V. T. Phillips 1923: 139 (type data). Neaspilota Osten Sacken 1878: 192 (new name for Aspilota; type species Trypeta alba Loew, automatic). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 610 (in catalog). - Williston 1908: 287 (in key). - Coquillett 1910: 574 (type data). Cresson 1914a: 276 (type data). - V. T. Phillips 1923: 122, 139 (in key; review, e. U S.). - Johnson 1925b: 262 (N. Engl.). - Johnson 1927: 218 (Mt. Desert). Curran 1932a: 3 (key to species). - Curran 1932b: 3 (in key). - F. H. Benjamin 1934: 34 (review, Fla.). - Curran 1934: 289 (in key). - Malloch 1942: 18 (key to species). - Quisenberry 1949c: 83 (key to species). - McFadden and Foote 1960: 254 (taxonomy). - Foote and Blanc 1963: 33 (review, Calif.). - Foote 1965a: 672 (in catalog). - Stegmaier 1968b: 46 (hosts). - Wasbauer 1972: 118 (hosts). - Steck 1981: 6 (hosts). - Freidberg and Mathis 1986: 1 (revision). - Goeden 1989a: 164 (hosts). Neospilota [error]: Williston 1896: 122 (in key). - Williston 1908: 287 (in key). Flendel 1914b: 92 (in key). Trypeta Meigen [misidentification]: Coquillett (part) 1899c: 262 (taxonomy). Aspilomyia Hendel 1907: 98 (new name for Aspilota; type species, Trypeta alba Loew, automatic). Terellia Robineau-Desvoidy [misidentification]: V. T. Phillips (part) 1923: 122, 138 (in key; review, e. N.A.). - Johnson 1925b: 262 (N. Engl.). Flies of the genus Neaspilota are small and yellowish, having con¬ vergent posterior orbital bristles (fig. 27, a); dorsocentral bristles approximately in recognition.

line with the supra-alars (fig. 23, a, b); and a dark lyre-shaped pattern, usually heavily tomentose, covering the otherwise yellow scutum. In these characters, the genus most closely resembles the North American species of Orellia, from which it may be distinguished by the generally smaller body size, the usual absence of setae on

222

Systematic Treatment of the Genera recognition.

In unifasciata, the dorsocentral bristles lie only slightly behind a

transverse line between the supra-alars, and the frontal bristles of the male are not enlarged. The largely hyaline cell c is divided at the center by a transverse brown band, only one hyaline triangle is present in cell r1 immediately distad of the pterostigma, and cell dm is entirely dark without any hyaline markings. This com¬ bination of characters distinguishes unifasciata from all other Myoleja species dis¬ cussed in this handbook. The female is still unknown. distribution.

host.

Not

Map 32.

known.

Genus Neaspilota Osten Sacken Aspilota Loew 1873: 286 (type species, Trypeta alba Loew 1861a: 345, by subse¬ quent designation of Coquillett 1910: 511) (preoccupied Forster, 1862). - Osten Sacken 1878: 260 (nomenclature). — V. T. Phillips 1923: 139 (type data). Neaspilota Osten Sacken 1878: 192 (new name for Aspilota; type species Trypeta alba Loew, automatic). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 610 (in catalog). - Williston 1908: 287 (in key). - Coquillett 1910: 574 (type data). Cresson 1914a: 276 (type data). - V. T. Phillips 1923: 122, 139 (in key; review, e. U S.). - Johnson 1925b: 262 (N. Engl.). - Johnson 1927: 218 (Mt. Desert). Curran 1932a: 3 (key to species). - Curran 1932b: 3 (in key). - F. H. Benjamin 1934: 34 (review, Fla.). - Curran 1934: 289 (in key). - Malloch 1942: 18 (key to species). - Quisenberry 1949c: 83 (key to species). - McFadden and Foote 1960: 254 (taxonomy). - Foote and Blanc 1963: 33 (review, Calif.). - Foote 1965a: 672 (in catalog). - Stegmaier 1968b: 46 (hosts). - Wasbauer 1972: 118 (hosts). - Steck 1981: 6 (hosts). - Freidberg and Mathis 1986: 1 (revision). - Goeden 1989a: 164 (hosts). Neospilota [error]: Williston 1896: 122 (in key). - Williston 1908: 287 (in key). Flendel 1914b: 92 (in key). Trypeta Meigen [misidentification]: Coquillett (part) 1899c: 262 (taxonomy). Aspilomyia Hendel 1907: 98 (new name for Aspilota; type species, Trypeta alba Loew, automatic). Terellia Robineau-Desvoidy [misidentification]: V. T. Phillips (part) 1923: 122, 138 (in key; review, e. N.A.). - Johnson 1925b: 262 (N. Engl.). Flies of the genus Neaspilota are small and yellowish, having con¬ vergent posterior orbital bristles (fig. 27, a); dorsocentral bristles approximately in recognition.

line with the supra-alars (fig. 23, a, b); and a dark lyre-shaped pattern, usually heavily tomentose, covering the otherwise yellow scutum. In these characters, the genus most closely resembles the North American species of Orellia, from which it may be distinguished by the generally smaller body size, the usual absence of setae on

223

Neaspilota

the node of vein Rs (fig. 47, a), and in most species the asymmetric fifth tarsomere of the male fore leg (see key and figs. 275-282). In contrast to all North American Ore Ilia species, cell r4+5 is hyaline or only very slightly clouded apically. Neaspilota also is very closely allied to Cbaetostomella (see discussion below), from which it may be distinguished by the smaller body size and by the absence of a group of strong dark setae on the anterior corner of the gena. Species of Paraterellia, some species of Procecidocharoides, Rbagoletis acuticornis, and Paroxyna p alii dip ennis, all of which also have convergent posterior upper fronto-orbital setae, might possibly be con¬ fused with Neaspilota, but these can be distinguished easily from all three New World terelliine genera by the lack of a lyre-shaped scutal pattern. The genera Neaspilota, Orellia, and Cbaetostomella are the North American representatives of the tribe Terelliini, a tephritid group that is mostly Holarctic. Freidberg and Mathis (1986) revised Neaspilota, an entirely New World genus, and divided it into two groups: the subgenus Neaspilota containing four species, and the subgenus Neorellia Freidberg and Mathis containing the remainder. The taxonomic characters of the two groups are set forth in the accompanying key. This division is supported by host data; almost all of the hosts of the four species of the subgenus Neaspilota belong to the tribe Vernoniaeae, whereas those in the sub¬ genus Neorellia infest some 20 plant genera in several other tribes of the Asteraceae. We are deeply indebted to Amnon Freidberg and Wayne N. Mathis for most of the information presented here. We have borrowed heavily from their revision of the genus (Freidberg and Mathis 1986) so that our key, descriptions, and additional data conform to that excellent review of a taxonomically difficult group of fruit flies. Many additional details are available from that publication. The genus occurs in northern Mexico, but its distribution south of the United discussion.

States remains unstudied.

Key to Continental U.S. and Canadian Species of Neaspilota 1. Frons setulose; in profile, frontofacial angle rounded, barely projecting ante¬ rior to eye, making head appear somewhat oval (fig. 268); anterior spine of tarsal comb of 5th tarsomere of male foreleg enlarged (fig. 275, a); anepisternum usually with 2 large, dark bristles; infesting species of Vernonia in eastern and central U.S. (subgenus Neaspilota).2 Frons usually bare, occasionally with 1-3 setulae; in profile, frontofacial angle distinctly projecting (fig. 269), making head appear squarish; tarsal comb with anterior spine not enlarged (fig. 276, a); anepisternum usually with only 1 large, dark bristle; hosts various, but not species of Vernonia; Alaska to Mexico and throughout U.S. (subgenus Neorellia).5 2. Wing with dark markings in addition to dark pterostigma (fig. 265); frons and face distinctly silvery tomentose.vernoniae (Loew) Wing entirely hyaline or with only pterostigma darkened; head nearly lack¬ ing silvery tomentum .3

Figures 265-267. Right wings, Neaspilota spp. 265, vernoniae (Lw.).; 266, achilleae Johns.; 267, viridescens Quis. Figures 268-271. Lateral view, head, Neaspilota spp. 268, alba (Lw.); 269, signifera (Coq.); 270, pubescens F.& M.; 271, aenigma F. & M. Figures 272-274. Dorsal view, abdomen, Neaspilota spp. 272, alba (Lw.).; 273, floridana Ibr.; 274, wilsoni Blanc and Foote.

224

a

Figures 275-282. Fifth tarsomere, male foreleg, Neaspilota spp. (redrawn from Freidberg and Mathis 1986). 275, alba (Lw.); 276, pubescens F. & M.; 277, callistigma F. & M.; 278, albiseta F. & M.; 279, dolosa Benj.; 280, footei F. & M.; 281, viridescens Quis.; 282, achilleae Johns. Figures 283, 284. Hind tibia, Neaspilota spp. 283, punctistigma Benj.; 284, appendiculata F. & M. Figure 285. Mid femur, male, Neaspilota isochela F. & M.

225

Systematic Treatment of the Genera

3. Abdominal setulae mostly brown, fine; body usually predominantly black; pterostigma uniformly dark, usually brown .albidipennis (Loew) Abdominal setulae mostly whitish, coarse; body predominantly yellow; pterostigma hyaline to yellowish .4 4. Oviscape longer than its width at base (fig. 272, a); abdominal tergites usually yellow, sometimes with dark areas .alba (Loew) Oviscape as long as, or shorter than, its width at base, equal to the length of 2 tergites (fig. 273, a); tergites predominantly black. .floridana Ibrahim 5. Abdominal setulae slender, brown; wing with distinct pattern, including a more or less complete transverse band aligned with vein dm-cu (fig. 266, a).achilleae Johnson Abdominal setulae whitish, coarse; wing without pattern as above, at most pterostigma brownish or apex infuscated.6 6. Head distinctly longer than high in profile (fig. 269); haustellum elongate, about 2 times as long as antenna (fig. 269, a); labellum spatulate (fig. 269, b); supra-alar bristle lacking; basal half of pterostigma brown ... .signifera (Coquillett) Head as high as, or higher than, long (fig. 270); haustellum slightly longer or shorter than antenna, labellum capitate (fig. 270, a); supra-alar bristle present; pterostigma variable .7 7. Scutum with whitish setulae arranged in 5 distinct longitudinal rows, the areas between them bare; abdomen shining; 5th tarsomere of male fore¬ leg with anterobasal spinose projection (fig. 276, b); oviscape equal to 2.5 tergites; V3 to 1h> of pterostigma brown. .pubescens Freidberg and Mathis Whitish setulae of scutum more or less uniformly distributed; abdomen subshining; 5th tarsomere of male foreleg not as above; oviscape and pterostigma variable.8 8. Abdominal tergal pattern with 4 triangular black areas on each tergite, the lateral triangular areas broader, areas sometimes fusing to form 2 spots on each tergite (fig. 274); pterostigma yellow, brownish on basal V3 to V2 .wilsoni Blanc and Foote Abdominal tergal pattern with black areas not as above, or tergites entirely yellow; pterostigma variable.9 9. Distal 1-3 tarsomeres darker than others, dark brown to black in male, brown in female; oviscape long, equal to 4-5 tergites; pterostigma yel¬ low, increasingly brown toward base; large species . .brunneostigmata Doane Tarsomeres unicolorous; oviscape shorter, or if as long, then body shorter and pterostigma hyaline or yellowish .10 10. Pterostigma with blackish brown spot on basal 2A to 3A; 5th tarsomere of male foreleg with a tranverse row of spines (fig. 277, a); oviscape equal to 2 tergites.callistigma Freidberg and Mathis Pterostigma hyaline, yellow, or with paler or smaller dark spot; 5th tar¬ somere of male foreleg not as above, usally with longitudinal comb; oviscape usually distinctly longer .11 11. Hind tibia with 1 or 2 semierect, short, distinct setae preapically and posteroventrally (fig. 283, a); pterostigma with squarish or rounded brown

227

Neaspilota

spot at basal V3, usually not quite reaching vein Rj; abdominal tergites mostly blackish.punctistigma Benjamin Hind tibia lacking setae as above; pterostigma and usually abdominal tergites not as above .12 12. Head distinctly higher than long (as in fig. 270), with head bristles usually concolorous; pterostigma usually opaque, yellow; 4th and 5th tarsomeres of male foreleg symmetrical, the 4th tarsomere with anteroapical spinose projection (fig. 278, a); oviscape equal to 3 tergites . .albiseta Freidberg and Mathis Head lower, with some of its bristles darker than others; 4th and sometimes also 5th tarsomeres of male foreleg symmetrical: oviscape variable .... .13 13. Fifth tarsomere of male foreleg, including claws and pulvilli, symmetrical; middle femur of male with longer, sparse, erect setae ventrally at basal half (fig. 285, a); comb of hind tibia indistinct; oviscape equal in length to 4.5 or 5 tergites; pterostigma and veins yellowish . .isochela Freidberg and Mathis Fifth tarsomere of male foreleg distinctly asymmetrical; comb of middle femur variable; comb of hind tibia variable; oviscape considerably short¬ er; pterostigma and veins variable.14 14. Comb on hind tibia indistinct; pterostigma usually hyaline or light yellow; abdominal tergites usually yellow, sometimes with narrow blackish bands; middle and hind femur of male with row of long, erect setae ventrally (as in fig. 285, a); 5th tarsomere of male foreleg, including claws and pulvilli, distinctly asymmetrical (fig. 279).dolosa Benjamin Comb on hind tibia distinct; pterostigma variable; abdomen with or with¬ out blackish bands; middle and hind femur of male with short, indistinct setulae ventrally; 5th tarsomere of male foreleg distinctly asymmetrical, or almost symmetrical, with symmetrical claws and pulvilli .15 15. Fifth tarsomere of male foreleg almost symmetrical, with symmetrical claws and pulvilli (fig. 280); pterostigma usually hyaline or light yellow; abdo¬ men usually with tergites more or less blackish basally, banded. .footei Freidberg and Mathis Fifth tarsomere of male foreleg, including claws and pulvilli, distinctly asymmetrical; pterostigma usually distinctly brown basally; abdomen banded or completely yellow .16 16. Dorsocentral bristles even with or slightly anterior to supra-alar bristles; wing (fig. 267) usually distinctly darkened, with dark pterostigma, cross¬ veins, longitudinal veins (especially at distal half), and microtrichiae (es¬ pecially on anterior half), but sometimes only the pterostigma and apices of longitudinal veins dark .viridescens Quisenberry Dorsocentral bristles in line with or slightly behind supra-alar bristles; wing not darkened, at most pterostigma and veins dark .17 17. Tarsal comb of male with only 2 rows of 2-3 spines each; pterostigma with a distinct brown spot basally; abdomen usually completely yellow, some¬ times with very narrow brown bands basally on tergites 3-5; oviscape about as long as 2 tergites .stecki Freidberg and Mathis Tarsal comb of male with more spines; pterostigma brown at base merged with the yellow at apex; abdomen completely yellow or banded; oviscape usually about as long as 3 tergites .18

228

Systematic Treatment of the Genera

18. Pterostigma mainly yellow, light brownish at basal third to half, without distinct border line between brown and yellow; larger species, wing length averaging 3.5 mm .appendiculata Freidberg and Mathis Pterostigma brown at basal half to two-thirds, yellow at apex; proximal border of brown clear-cut, distal border not so; smaller species, wing length averaging 2.5 mm .aenigma Freidberg and Mathis

Neaspilota achilleae Johnson (Figs. 266, 282; Map 33) Neaspilota achilleae Johnson 1900a: 328 (male lectotype, MCZ; N.J.). - Aldrich 1905: 610 (in catalog). - Johnson 1909: 113 (taxonomy). - Johnson 1910: 802

229

Neaspilota

(N.J.).-Johnson 1913b: 84 (Fla.). - Sturtevant 1918: 36 (Ala.).-Johnson 1925b: 263 (N. Engl.). — Sturtevant 1925: 215 (seminal receptacles). - Johnson 1930: 51 (Nantucket). - Curran 1932a: 3 (in key). - F. H. Benjamin 1934: 36, 37 (in key; review, Fla.). — Brimley 1938: 384 (N. Car.). — Malloch 1942: 19 (in key). — V. T. Phillips 1946: 51, 116 (description, biology larva; hosts). - Quisenberry 1949c: 84 (in key). — Muesebeck and Walkley 1951: 166 (parasite). — Hennig 1952: 214 (in catalog, immature stages). - Foote 1965a: 672 (in catalog). - Stegmaier 1968b: 47 (hosts). - Wasbauer 1972: 118 (hosts). — Freidberg and Mathis 1986: 28 (revision, synonymy, lectotype designation). - Goeden 1989a: 164 (host). Trypeta acbilleae: Johnson 1900b: 687 (N.J.). — C. F. Baker 1904: 30 (Calif.). Trypeta acbillae [error]: Woodworth 1913: 137 (Calif.). Neaspilota ocbilleae [error]: V. T. Phillips 1923: 140 (review). N. acbilleae superficially resembles small specimens of vernoniae, which also has a patterned wing in addition to a basally dark pterostigma (fig. 266). In contrast to acbilleae, vernoniae possesses dark-colored abdominal setae, the head recognition.

is oval rather than square in profile, the frons is haired, and two anepisternal bristles are present. These two species are the only Neaspilota with any discrete wing mark¬ ings other than on the pterostigma. Map 33. It is not certain whether the smaller western U.S. speci¬ mens may represent a disjunct population (Freidberg and Mathis 1986). distribution.

Freidberg and Mathis (1986) reported species in the genera Aster, Cbrysopsis, Erigeron, Hieracium, Prenantbes, Sericocarpus, and Trilisa as hosts of acbilleae. Goeden (1989a) reared acbilleae from Stepbanomeria virgata in southern California. hosts.

Neaspilota aenigma Freidberg and Mathis (Fig. 271; Map 34) Neaspilota aenigma Freidberg and Mathis 1986: 32 (male holotype, NMNH; Death Valley Junction, Inyo County, Calif.). — Goeden 1989a: 165 (hosts). N. aenigma is recognized by its head profile (fig. 271); bare frons; one anepisternal bristle; uniformly distributed scutal setulae; yellow or narrowly black-banded abdominal tergites; pterostigma basally dark brown and apically yellow; and asymmetrical male fifth fore tarsomere, claws, and pulvilli. It is distin¬ guished from stecki by its pterostigmal marking, which intergrades smoothly from its brown base to a yellow apex, and from viridescens by the more posterior position recognition.

of the dorsocentral bristles, which are behind a line through the supra-alars. It is distinguished from appendiculata by its smaller size and darker brown pterostigma. distribution.

Map 34.

230

Systematic Treatment of the Genera

Haplopappus Goodingii is a confirmed host, and H. hartwigi is a sus¬ pected host (Freidberg and Mathis 1986). Goeden (1989a) reared aenigma from one species each of the genera Acamptopappus and Machaerantbera in California. HOSTS.

Neaspilota alba (Loew) (Figs. 268, 272, 275; Map 34) Trypeta alba Loew 1861a: 345 (male lectotype, MCZ; Pa.). - Loew 1861b: 39 (see

Loew 1861a). — Loew 1862c: 64 (in key). — Osten Sacken 1862: 100 (description adult). - Loew 1864: 39 (taxonomy). - Loew 1873: 285 (review). - Doane 1899: 187 (taxonomy). - Coquillett 1910: 511 (type species designation). - Freidberg and Mathis 1986: 15 (lectotype designation). Aspilota alba: Loew 1873: 286, 336 (taxonomy). - Osten Sacken 1880: 53 (host). Trypeta (Aspilota) alba: Osten Sacken 1877: 345 (taxonomy). Trypeta (Neaspilota) alba: Osten Sacken 1878: 192, 260 (in catalog). Neaspilota alba: Johnson 1900b: 688 (N.J.). - F. H. Snow 1903: 219 (Kans.). Aldrich 1905: 610 (in catalog). - Cresson 1907: 101 (description adult). - Tucker 1907: 104 (Kans., Colo.). — Coquillett 1910: 574 (type data). — Johnson 1910: 802 (N. J.). - V. T. Phillips 1923: 139 (review). - Johnson 1925b: 262 (N. Engl.). Leonard 1928: 852 (N.Y.). - Curran 1932a: 3 (in key). - Janes and Thomas 1932: 103 (Utah). - Brimley 1938: 384 (N. Car.). - Malloch 1942: 19 (in key). Brandhorst 1943: 168 (seed damage). - Schwitzgebel and Wilbur 1943: 4 (biolo-

231

Neaspilota

gy). — V. T. Phillips 1946: 116 (hosts). — Quisenberry 1949c: 84 (in key). — Steyskal 1957: 94 (eye color). — Foote 1965a: 672 (in catalog). - Cole and Schlinger 1969: 355 (in list). — Wasbauer 1972: 119 (hosts). — Ibrahim 1982: 297 (taxonomy). — Freidberg and Mathis 1986: 12 (revision). Neoaspilota [error] alba: Hendel 1914b: 92 (type data). recognition.

N. alba, like floridana, has an oval head, haired frons, predomi¬

nantly yellow body, two anepisternal bristles, and a yellowish pterostigma. It may be distinguished from floridana by the almost completely yellow abdomen with occa¬ sional irregular black spots that never form distinct black bands. The oviscape in alba (fig. 272) is longer than its width at base and is usually about as long as the last three abdominal tergites; that of floridana (fig. 273) is shorter than its basal width. distribution.

hosts.

Map 34.

Vernonia Baldwini, V. interior, and V. sp. were reported as hosts by Freid¬

berg and Mathis (1986). Most of the collection records of alba that are dated earlier than Freidberg and Mathis (1986) (from Kansas to Pennsylvania, south to Texas and Florida) actually refer to floridana. See the discussion of that species. discussion.

Neaspilota albidipennis (Loew) (Map 35) Trypeta albidipennis Loew 1861a: 345 (male lectotype, MCZ: Pa.). — Loew 1861b:

39 (see Loew 1861a). — Osten Sacken 1862: 100 (description). — Loew 1864: 39 (taxonomy). — Loew 1873: 286 (review). — Doane 1899: 188 (taxonomy). — Freidberg and Mathis 1986: 17 (lectotype designation). Aspilota albidipennis: Loew 1873: 286, 330 (taxonomy). Trypeta (Neaspilota) albidipennis: Osten Sacken 1878: 192 (in catalog). Neaspilota albidipennis: Johnson 1900b: 688 (N.J.). - Aldrich 1905: 610 (in catalog). — Johnson 1910: 802 (N.J.). — V. T. Phillips 1923: 140 (review). — Curran 1932a: 3 (in key). - F. H. Benjamin 1934: 36 (in key). — Curran 1934: fig. 34 (figure head). — Malloch 1942: 19 (in key). — Schwitzgebel and Wilbur 1943: 4 (biology). — V. T. Phillips 1946: 53, 116 (description, biology larva; hosts). — Quisenberry 1949c: 84 (in key). — Hennig 1952: 214 (in catalog, immature stages). — Steyskal 1957: 94 (eye color). — Foote 1965a: 672 (in catalog). — Wasbauer 1972: 119 (hosts). — Freidberg and Mathis 1986: 16 (revision). Neaspilota albipennis [error]: Johnson 1925b: 263 (N. Engl.). N. albidipennis closely resembles alba, floridana, and vernoniae in having an oval head, a haired frons, and two anepisternal bristles. It is distin¬ guished from vernoniae by the lack of dark markings on the wing other than in the pterostigma, and from alba and floridana by the fact that the pterostigma is quite recognition.

232

Systematic Treatment of the Genera

Map 35. Distribution of Neaspilota albidipennis, N. albiseta, and N. appendiculata.

dark rather than hyaline or pale yellow and by the predominantly dark body color. Occasional specimens from Michigan and Kansas are reported to be generally paler. distribution.

hosts.

Map 35.

N. albidipennis is restricted to several species of Vernonia.

Neaspilota albiseta Freidberg and Mathis (Fig. 278; Map 35) Neaspilota albiseta Freidberg and Mathis 1986: 35 (male holotype, NMNH; Valley

Junction, Inyo County, Calif.). - Goeden 1989a: 165 (host). Unusual among Neaspilota species, the head of albiseta in profile, although squarish, is distinctly higher than long (as in figs. 268, 270) with most head setae the same color and white rather than yellow as in most other species. The pterostigma is opaque but uniformly yellow, and veins r-m, dm-cu, and the distal parts of the longitudinal veins are darkened. The oviscape is about as long as the last two abdominal tergites. The fourth tarsomere of the male fore tarsus has a spinose projection (fig. 278, a) similar to that on the fifth tarsomere of pubescens (fig. 276, b), to which albiseta is closely related. In the former species, the scutal setae are arrayed in longitudinal rows, whereas in the latter they are more or less uniformly distributed. recognition.

233

Neaspilota

distribution.

Map 35.

Goeden (1989a) reared albiseta from mature heads of Machaeranthera tortifolia in Inyo County, Calif. host.

Neaspilota appendiculata Freidberg and Mathis (Fig. 284; Map 35) Neaspilota appendiculata Freidberg and Mathis 1986: 37 (male holotype, NMNH;

Williams, Ariz.). — Goeden 1989a: 166 (hosts). recognition.

N. appendiculata has genitalia very similar to those of wilsoni,

from which it can be distinguished by the presence of narrow black bands on several of the abdominal tergites (2—4 black triangular markings are characteristic of wil¬ soni). Additional characters helping to distinguish it from other Neaspilota species are its relatively large size and wing length and the coloration of the pterostigma. The dorsocentral bristles are situated slightly behind a line through the supra-alar bris¬ tles, the claws and pulvilli of the male fifth fore tarsomeres are asymmetrical, and a distinct comb of setae is present ventrally on the hind tibia (fig. 284). distribution.

host.

Map 35.

In California, appendiculata was reared by Goeden (1989a) from Ma-

chaeranthera canescens.

Neaspilota brunneostigmata Doane (Map 36) Neaspilota brunneostigmata Doane 1899: 187 (female lectotype, WSU; Pullman,

Wash.). — Cole and Lovett 1921: 326 (Oreg.). — Curran 1932a: 3 (in key). — F. H. Benjamin 1934: 35 (in key). — Foote and Blanc 1963: 33 (review, Calif.). — Foote 1965a: 672 (in catalog). - Cole and Schlinger 1969: 355 (review). — Wasbauer 1972: 119 (hosts). — Zack 1984: 32 (type data). — Freidberg and Mathis 1986: 39 (revision). Neaspilota hrunneistigmata [error]: Aldrich 1905: 610 (in catalog). Neaspilota brunneistigma [error]: Malloch 1942: 19 (in key). — Quisenberry 1949c: 84 (in key). Neaspilota brunneostigma [error]: Goeden 1989a: 166 (hosts). N. brunneostigmata is one of the largest Neaspilota species and usually may be recognized, along with isochela, by its oviscape, which is nearly as recognition.

long as the abdomen. The pterostigma is yellow, becoming gradually browner to¬ ward its base. The apical one to three tarsomeres of the male are usually brown to black and darker than the basal ones, although that color is pale and more difficult to distinguish in females. All the tarsomeres of isochela are unicolorous.

Systematic Treatment of the Genera V

o

Q

r -'-'rL

1

o

O /

°

/7

o

° °L

oW

jp

J> o

/

1

1—•l •• ••

•

i

• J,

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• Neaspilota brunneostigmata o Neaspilota vernoniae

Map 36. Distribution of Neaspilota brunneostigmata and N. vernoniae.

distribution. Map 36. hosts. Freidberg and Mathis (1986) listed species in the genera Aster, Cbrysotbamnus, Coretbrogyne, Erigeron, Haplopappus, Heterotbeca, and Macbaerantbera as hosts of brunneostigmata. Goeden (1989a) reared this species from Lessingia lemmonii in California, having reported it originally as aenigma. discussion. N. brunneostigmata has often been misidentified in the literature, and all the records cited in the foregoing synonymy should be re-examined in every case where specimens are available.

235

Neaspilota

Map 37. Distribution of Neaspilota callistigma, N. pubescens, and N. wilsoni.

Neaspilota callistigma Freidberg and Mathis (Fig. 277; Map 37) Neaspilota signifera [misidentification]: Foote and Blanc 1963: 34 (in part). Neaspilota callistigma Freidberg and Mathis 1986: 43 (male holotype, CAS; San Diego, Calif.). — Goeden 1989a: 166 (hosts). Neaspilota sp.: Goeden 1986: 327 (host). recognition. The wing of callistigma is mainly hyaline with some of the veins slightly darkened; the pterostigma is unique among known Neaspilota in containing a striking blackish brown spot in its basal two-thirds to three-fourths, with the apex yellow. The oviscape is relatively short, only as long as the last two abdominal tergites. The fifth tarsomere of the male foreleg possesses a transverse rather than a longitudinal comb (fig. 277, a), and the claws are about equally long. The species may be distinguished from signifera, with which it has apparently been confused in the past, by the head, which is higher than long in profile. distribution. Map 37. N. callistigma also occurs in Baja California. hosts. Haplopappus venetus var. vernonioides was reported as a host by Freid¬ berg and Mathis (1986). Goeden (1989a) reared callistigma from that plant as well as from Haplopappus acradenius in California.

Neaspilota dolosa Benjamin (Fig. 279; Map 38) Neaspilota dolosa Benjamin 1934: 39 (male holotype, NMNH; Orlando, Fla.). Malloch 1942: 20 (review). — V. T. Phillips 1946: 116 (host). — Quisenberry

236

Systematic Treatment of the Genera

Map 38. Distribution of Neaspilota dolosa and N. viridescens.

1949c: 84 (in key). — Hennig 1952: 214 (in catalog, immature stages). — Foote 1965a: 672 (in catalog). - Stegmaier 1968b: 47 (hosts). - Wasbauer 1972: 119 (hosts). - Freidberg and Mathis 1986: 43 (revision). - W. A. Palmer 1987: 192 (hosts). recognition. N. dolosa is often difficult to distinguish from other Neaspilota species by color or wing character alone. As in isochela, long yellow ventral setae are present near the bases of the middle and hind femora of the male (as in fig. 285, a), and both sexes lack a tibial comb. N. dolosa may be distinguished from isochela by

237

Neaspilota the distinctly asymmetrical fifth tarsomere of the male foreleg (fig. 279) and the shorter oviscape. The pterostigma of dolosa is opaque, yellowish, and sometimes darker yellow or pale brown at its base. distribution.

Map 38. N. dolosa also occurs in northern Mexico.

Freidberg and Mathis (1986) listed species in the plant genera Aster, Chrysopsis, Erigeron, Heterotheca, Pluchea, and Sideranthus as hosts of dolosa. hosts.

Neaspilota floridana Ibrahim (Fig. 273; Map 34) Neaspilota alba (Loew) [misidentification]: F. FI. Benjamin 1934: 36, 37 (in key; review, Fla.). Also probably many other authors. Neaspilota floridana Ibrahim 1982: 297 (female holotype, NMNH; Orlando, Fla.). — Freidberg and Mathis 1986: 20 (revision). Among the species of Neaspilota with an oval head, a haired frons, two anepisternal bristles, and a predominantly yellow body, floridana is distin¬ guished by the presence of distinct wide black bands on the abdominal tergites. In this respect it differs from the closely similar alba, which only occasionally has dark spots that never form distinct bands. The oviscape in floridana is shorter than, or equal to, its width at the base and is only about as long as the last two abdominal tergites (fig. 273). In alba, the oviscape is longer than its basal width (fig. 272). recognition.

distribution.

Map 34.

Several species of Vernonia are confirmed hosts; floridana also was repor¬ tedly reared from Pluchea and Chrysopsis, but Freidberg and Mathis (1986) doubted hosts.

the validity of those records. Before its description in 1982, floridana was misidentified as alba, especially as detailed by Benjamin (1934). It does not appear possible to sort out and discussion.

correct these earlier misidentifications.

Neaspilota footei Freidberg and Mathis (Fig. 280; Map 33) Neaspilota footei Freidberg and Mathis 1986: 49 (male holotype, NMNH; White Hall, Frederick County, Md.). recognition.

The claws and pulvilli of males of footei are symmetrical, but the

fifth tarsomere of the male foreleg possesses one or two distinct spines (fig. 280, a) anteroapically. The comb on the hind tibia is distinct (as in fig. 285, a) in contrast to that of dolosa, which is absent or indistinct. The middle and hind femora have short,

238

Systematic Treatment of the Genera

indistinct setulae ventrally, and the hind tibia lacks semierect setae. The abdominal tergites usually are blackened basally, broadly in males and less so in females. Map 33. According to Freidberg and Mathis (1986), the distribu¬ tional pattern may be an artifact because of insufficient collecting in the midwestern and south central states. distribution.

Erigeron canadensis and Chrysothamnus sp. were listed by Freidberg and Mathis (1986). hosts.

Neaspilota isochela Freidberg and Mathis (Fig. 285; Map 33) Neaspilota isochela Freidberg and Mathis 1986: 52 (male holotype, NMNH; Mexia, Tex.). recognition.

One of the few species of Neaspilota with the fore fifth tarsomere

of the male entirely symmetrical, and only rarely with black markings on the abdom¬ inal tergites. As in brunneostigmata, the oviscape is one of the longest in the genus, about equal to the length of the abdomen, but isochela is distinguished from that species by the entirely yellow pterostigma, sometimes slightly tinged with brown basally, in both sexes. The hind tibia lacks a distinct comb, but erect ventral setae are present on the middle and hind femora of the male (fig. 285, a). distribution.

Map 33.

Chrysopsis sp., Grindelia sp., and Sideranthus rubiginosus were listed by Freidberg and Mathis (1986). hosts.

Neaspilota pubescens Freidberg and Mathis (Figs. 270, 276; Map 37) Neaspilota pubescens Freidberg and Mathis 1986: 55 (male holotype, NMNF1; Barton Flat, So. Fork Camp, Calif.). N. pubescens is unique among members of this genus in having five longitudinal patches or rows of setulae on the scutum. The scutal tomentum is recognition.

more yellow between the rows of setulae; thus, this species is still recognizable if the setulae have been rubbed off or otherwise damaged. The fore fifth tarsomere of the male possesses a spinose projection (fig. 276, b), also unique among Neaspilota species. In other respects, this species somewhat resembles punctistigma, from which it may be separated by the characters given in the key to species. distribution.

Map 37.

239

Neaspilota Goeden (1989a) reared pubescens (reported as appendiculata) from Corethrogyne filaginifolia in Fresno County, California. host.

Neaspilota punctistigma Benjamin (Fig. 283; Map 39) Neaspilota punctistigma Benjamin 1934: 38 (male holotype, NMNFi; 4 mi. e. Ft. Christmas, Fla.). — Malloch 1942: 19 (review). — V. T. Phillips 1946: 116 (hosts). — Quisenberry 1949c: 84 (in key). — Hennig 1952: 214 (in catalog, immature stages). — Foote 1965a: 672 (in catalog). — Wasbauer 1972: 119 (hosts). — Freidberg and Mathis 1986: 57 (revision). A relatively small species, punctistigma may be recognized by the shape of the head, which is almost square in profile; by the haustellum, which is about as long as the antenna; and by the mostly blackish abdominal tergites. It is the only Neaspilota species in which one or two small preapical posteroventral setae are present on the hind tibia (fig. 283, a), a character sometimes difficult to see in pinned specimens if the legs are folded. It is the only Neaspilota species in which the dark brown area in the basal one-third of the pterostigma has the appearance of a squarish spot. recognition.

distribution.

Map 39.

Map 39. Distribution of Neaspilota punctistigma, N. signifera, and N. stecki.

240

Systematic Treatment of the Genera hosts.

Chrysopsis and Pluchea are the principal plant genera infested.

discussion.

Benjamin (1934) figured the larva and pupa of punctistigma.

Neaspilota signifera (Coquillett) (Fig. 269; Map 39) Trypeta (Neaspilota) signifera Coquillett 1894: 73 (male lectotype, NMNH; Los Angeles, Calif.). - Freidberg and Mathis 1986: 62 (lectotype designation). Trypeta signifer [error]: Doane 1899: 187 (taxonomy). - Johnson 1913b: 84 (Fla.). Neaspilota signifera: Aldrich 1905: 610 (in catalog). - Curran 1932a: 3 (in key). F. H. Benjamin 1934: 36 (in key). — Malloch 1942: 19 (in key). — Quisenberry 1949c: 84 (in key). - Foote and Blanc (part) 1963: 34 (review, Calif.). - Foote 1965a: 672 (in catalog). - Cole and Schlinger 1969: 355 (review). - Wasbauer 1972: 120 (hosts). — Freidberg and Mathis 1986: 60 (revision). Neospilota [error] signifera: Woodworth 1913: 137 (Calif.). N. signifera is unique among species of Neaspilota in that the supra-alar bristles are lacking; the head, squarish in profile, is distinctly longer than high and with a bare frons; and the haustellum and labellum are elongated (fig. 269, recognition.

a, b). The hind tibia lacks semierect setae, the femur lacks well-developed ventral setae, the comb of the hind tibia is distinct, and the fifth tarsomere of the male foreleg is elongated. The wing is mainly hyaline but has a pale brown spot on the humeral crossvein, and the pterostigma is brown in its basal half. distribution.

Map 39. This species also occurs in northern Mexico.

Baeria Fremontii and Hemizonia pungens were the only two hosts listed by Freidberg and Mathis (1986). hosts.

According to Freidberg and Mathis (1986), some of the specimens identified as signifera by Foote and Blanc (1963) are actually callistigma. The host data reported by them actually apply to callistigma. discussion.

Neaspilota stecki Freidberg and Mathis (Map 39) Neaspilota stecki Freidberg and Mathis 1986: 63 (male holotype, NMNH; Willard, Torrence County, N. Mex.). - Goeden 1989a: 166 (host). In many respects, stecki is similar to aenigma, from which it can be separated by the basal brown area in the pterostigma, which is darker, more conspicuous, and more strongly contrasting with the apical yellow color than in RECOGNITION.

241

Neaspilota

aenigma. The oviscape is much shorter, and genitalic characters provide further means of identification. N. stecki differs from aenigma and most other Neaspilota in the nature of the tarsal comb, which consists of two rows of only two or three spines each. distribution.

Map 39.

In California, Goeden (1989a) reared stecki from Acamptopappus spkaerocepbalus, the only known host. host.

Neaspilota vernoniae (Loew) (Figs. 49, 265; Map 36) Trypeta vernoniae Loew 1861a: 346 (male lectotype, MCZ; Pa.). - Loew 1861b: 40 (see Loew 1861a). - Osten Sacken 1862: 101 (description, adult). - Loew 1864: 40 (taxonomy). - Loew 1873: 286 (review). — Coquillett 1899c: 262 (taxonomy). -Johnson 1900b: 687 (N.J.). - Freidberg and Mathis 1986: 23 (lectotype designa¬ tion). Aspilota vernoniae: Loew 1873: 287, 330 (taxonomy). Trypeta (Aspilota) vernoniae: Osten Sacken 1878: 192 (in catalog). Neospilota [error] vernoniae: Johnson 1895b: 337 (Fla.). Neaspilota vernoniae: Coquillett 1899c: 262 (taxonomy). - Aldrich 1905: 610 (in catalog). — Johnson 1910: 802 (N.J.). - Johnson 1913b: 84 (Fla.). — F. H. Benjamin 1934: 36 (in key). — Malloch 1942: 18 (in key). — Schwitzgebel and Wilbur 1943: 4 (note, biology). - Quisenberry 1949c: 83 (in key). - Steyskal 1957: 94 (eye color). - Foote 1965a: 672 (in catalog). - Wasbauer 1972: 120 (host). - Freidberg and Mathis 1986: 23 (revision). Terellia vernoniae: V. T. Phillips 1923: 139 (review). - Johnson 1925b: 262 (N. Engl.). — Brimley 1938: 384 (N. Car.). Among the species with an oval head, setose frons, and two anepisternal bristles, vernoniae is distinctive in that, in addition to the pterostigma, the wing is patterned (fig. 265). In this respect it superficially resembles acbilleae, but the large yellow body, the yellow abdominal setulae, and the characters given above distinguish vernoniae from that species. N. vernoniae and acbilleae are the only known Neaspilota species occurring north of Mexico with extensively patterned recognition.

wings. distribution.

hosts.

Map 36. Johnson’s record from Florida may represent acbilleae.

N. vernoniae is known only from Vernonia interior and Vernonia sp.

Almost nothing is known about the biology of this species. See Schwitzgebel and Wilbur (1943) for the minimal details given there. discussion.

Systematic Treatment of the Genera

Neaspilota viridescens Quisenberry (Figs. 267, 281; Map 38) Neaspilota viridescens Quisenberry 1949c: 82 (female holotype, NMNH; Rabbit Ears Pass, Colo.). — Foote 1965a: 672 (in catalog). — Freidberg and Mathis 1986: 65 (revision). - Goeden 1989a: 166 (hosts). - Goeden and Fleadrick 1992: 59 (life history, immature stages, hosts).

recognition.

The wing of viridescens (fig. 267) lacks a distinct pattern, but

many specimens have faint brownish areas, especially near the distal portions of the radial veins and veins r-m and dm-cu. The veins are brownish in these areas, and the pterostigma is entirely light brown or yellowish at its apex. The dorsocentral bristles are in line with or slightly anterior to the supra-alars; the hind tibia lacks setae preapically and posteroventrally; and the fifth tarsomere of the male foreleg (fig. 281) is asymmetrical, with the anterior claw somewhat longer than the posterior one. This species is often difficult to distinguish from aenigma on the basis of external characters.

distribution.

HOSTS.

Map 38.

Aster spinosus was the only host recorded by Freidberg and Mathis

(1986). Since then, Goeden (1989a) reared viridescens from an additional species of Aster and from one species of Chrysothamnus, two species of Erigeron, one species of Gutierrezia, six species of Haplopappus, one species of Lepidospartum, and two species of Mach aer anther a in California. Goeden and Headrick (1992) reared vir¬ idescens from two additional species of Aster and Chrysopsis villosa.

discussion.

Freidberg and Mathis (1986) suggested that viridescens very proba¬

bly represents a complex of species that requires further study. The life history of viridescens was reported in detail by Goeden (1992).

Neaspilota wilsoni Blanc and Foote (Fig. 274; Map 37) Neaspilota wilsoni Blanc and Foote 1961: 78 (male holotype, CAS; Jacolitas Can¬ yon, Fresno County, Calif.). - Foote and Blanc 1963: 35 (review, Calif.) - Foote 1965a: 672 (in catalog). - Wasbauer 1970: 50 (type data). - Arnaud 1979: 330 (type data). - Freidberg and Mathis 1986: 69 (revision). - Goeden 1989a: 167 (hosts).

N. wilsoni may be distinguished from most other Neaspilota spe¬ cies by the presence of triangular black spots on each of tergites 3-5 of males and 36 of females (fig. 274). The two spots on either side are usually fused at the anterior recognition.

243

Neotephritis margin in males, but in females they are usually separate and are generally lighter in color. The dorsocentral bristles are even with or slightly posterior to the supra-alars, the hind tibia lacks semierect setae preapically and posteroventrally, and the fifth tarsomere of the male foreleg is asymmetrical. The pterostigma is brown basally, graduating to yellow apically, with veins r-m and dm-cu usually brownish. distribution.

Map 37.

Haplopappus squarrosus ssp. grindelioides, the only known host, was re¬ corded by Freidberg and Mathis (1986). Goeden (1989a) indicated that a doubtful record of Coreopsis calliosidea is probably not valid. host.

Genus Neotephritis Hendel Tepbritis (part): Loew 1873: 295 (taxonomy); 328 (in key). — Aldrich 1905: 64 (in catalog). Trypeta (Tepbritis) (part): Osten Sacken 1878: 193 (in catalog). Neotephritis Hendel 1935: 54 (type species, Trypeta finalis Loew 1862a: 222 (1864: 92), by original designation). - Foote 1960e: 72 (taxonomy). - Foote 1960f: 145 (revision). — Foote and Blanc 1963: 35 (review, Calif.). - Foote 1965a: 666 (in catalog). — Foote 1967b: 31 (in catalog). — Steyskal 1972c: 414 (key to known species). — Wasbauer 1972: 120 (hosts).

recognition.

Small flies with black bristles and gray to yellowish gray body

tomentum. There are three pairs of frontal bristles, two pairs of orbital bristles, two pairs of scutellar bristles, and the single pair of dorsocentral bristles is situated very close behind the transverse suture. Neotephritis is easy to distinguish from all other Nearctic genera with hyaline-spotted wings, even with the unaided eye, by the promi¬ nent inverted triangle at the apex of the pterostigma composed of several partially fused hyaline spots, and similar fairly prominent hyaline areas near the apex of cell r2+3 and in cell m. Neotephritis (at least the two species north of Mexico) and Acinia are the only two genera in the United States and Canada with three pairs of frontal and two pairs of scutellar bristles in combination with hyaline-spotted wings in which the apex of cell r4+5 is entirely dark. Neotephritis may be separated from Acinia by the characters given in the key and the discussion of the latter genus. N. finalis is one of the most commonly encountered tephritids in the United States and Canada. At least eight other species have been described from various locations in Mexico, the West Indies, and Central and South America, and discussion.

two species are known from Hawaii.

Systematic Treatment of the Genera

Figures 286-288. Right wings, Neotephritis spp. 286, finalis (Lw.) (typical); 287, finalis (Lw.) (variant); 288, rava Foote. Figures 289, 290. Dorsal view, oviscape, Neotephritis spp. 289, finalis (Lw.); 290, rava Foote.

Key to U.S. and Canadian Species of Neotephritis Scutum and abdominal tergum gray; oviscape (viewed from above) about 1.5 times as long as wide at base (fig. 289); wing grayish, the hyaline spots with distinct margins; wing pattern as in figs. 286 and 287 .finalis (Loew) Scutum and abdominal tergum yellow or amber; oviscape (when seen from above) somewhat longer than wide at base (fig. 290, a); wing yellowish brown, the hyaline spots with indistinct margins; wing pattern as in fig. 288 .rava Foote

Neotephritis finalis (Loew) (Figs. 100, 286, 287, 289; Map 40) Trypeta finalis Loew 1862a: 222 (1864: 92) (male and female syntypes, MCZ; Calif.). - Loew 1873: 296 (review, taxonomy).

Neotepbritis

• •

245

•

Neotepbritis finalis

★ Neotepbritis finalis (state only record) °

Neotepbritis rava

Map 40. Distribution of Neotepbritis finalis and N. rava.

Tepbritis finalis: Loew 1873: 297, 330 (taxonomy). - W. A. Snow 1894: 172 (taxon¬ omy). - Coquillett 1899c: 264 (synonymy). - Doane 1899: 188 (distribution). Wulp 1900: 419 (in catalog). - Johnson 1903b: 106 (Beulah, N. Mex.). - F. H. Snow 1903: 219 (Kans.). - F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 611 (in catalog). - Tucker 1907: 105 (Colo.). - Hyslop 1912: 101 (biology). — Cockerell 1914: 195 (host, distribution). - Cole and Lovett 1921: 326 (Oreg.). Flendrickson 1930: 143 (Iowa prairies). - Janes and Thomas 1932: 104 (Utah). Bissell 1936: 233 (oviposition damage). — Walker 1936: 21 (biology). - Breland 1938: 204, 205 (parasites). - Essig 1938: 605 (note). - Breland 1939: 719 (biolo¬ gy, parasites). - V. T. Phillips 1946: 79, 123 (description, biology larva; host). Essig 1958: 605 (note).

Systematic Treatment of the Genera Trypeta (Tephritis) finalis: Osten Sacken 1877: 346 (in catalog). — Osten Sacken 1878: 193 (in catalog). Tephritis affinis Snow 1894: 172 (male and female syntypes, UKL; Mont., Calif.). Coquillett 1899c: 264 (synonymy). - Aldrich 1905: 611 (in catalog). Quisenberry 1951: 59 (taxonomy). Tephritis inornata Coquillett 1902b: 181 (male and female syntypes, NMNH; Las Vegas Hot Springs, N. Mex.). — Aldrich 1905: 612 (in catalog). — Cresson 1907: 104 (description adult). — Tucker 1908: 303 (Colo.). — Goeden et al. 1987: 552 (synonym). Straussia affinis: Woodworth 1913: 137 (Calif.). Euribia finalis: Hendel 1914c: 67 (review). — Knowlton and Harmston 1937: 145 (Utah). — Brimley 1938: 385 (N. Car.). — Aczel 1949: 184 (in catalog). Spathulina finalis: Cockerell 1917: 17 (Boulder County, Colo.). Neotephritis finalis: Hendel 1935: 54 (type data). — Quisenberry 1951: 59 (taxon¬ omy). — Foote 1960f: 146 (review). — Foote 1962: 172 (taxonomy). — Foote and Blanc 1963: 35 (review, Calif.). — Foote 1965a: 669 (in catalog). — Novak et al. 1967: 148 (hosts, biology). — Schulz and Lipp 1969: 99 (biology, damage). — Lipp and Schulz 1970: 27 (damage). — Beckham and Tippins 1972: 865 (biology). Steyskal 1972c: 415 (in key, known species). - Wasbauer 1972: 120 (hosts). Hardy and Delfinado 1980: 48 (Hawaii). - Hilgendorf and Goeden 1981: 105 (host). — Hilgendorf and Goeden 1982: 152 (host). — Goeden et al. 1987: 552 (biology, taxonomy). — W. A. Palmer 1987: 192 (hosts). Neotephritis inornata: Quisenberry 1951: 59 (taxonomy). - Foote 1960f: 149 (revi¬ sion). — Foote and Blanc 1963: 37 (review, Calif.). - Foote 1965a: 669 (in catalog). - Steyskal 1972c: 415 (in key). — Goeden et al. 1987: 552 (synonymy). Thephritis [error] finalis: Hennig 1952: 212 (in catalog immature stages). Euarestoides finalis: Beirne 1971: 50 (economic importance). Neotephritis electellum [error]: Hilgendorf and Goeden 1981: 105 (host).

recognition.

Individuals of finalis are larger and more robust than rava. Usu¬

ally only two rounded hyaline spots are present in cell r4+5 immediately anterior to vein M (fig. 286, b); in rava, three or more spots are present in that position. However, a small percentage of finalis specimens show three spots (fig. 287, a), but those flies can easily be distinguished from rava by other characters given in the key. In finalis, a hyaline mark is almost always present close to the apex of cell r1 (fig. 286, a); this spot is never present in rava. All of the hyaline spots in the wing of finalis are more sharply bordered by the dark background color, and less fusion of these spots is characteristic of finalis. The oviscape of finalis is usually darkened at its apex and is about 1.5 times as long as its width at the base (fig. 289).

Map 40. In addition to its distribution in the United States and Canada, finalis has been reported from the Mexican states of Durango, Jalisco, Guerrero, Veracruz, and Guadalajara, and it may well occur elsewhere in Mexico. distribution.

247

Oedicarena Goeden (1987) reported that finalis has been reared from the flower heads of many genera of the subtribe Verbesininae of the tribe Heliantheae, family Asteraceae. hosts.

N. finalis is one of the most commonly encountered tephritids in North America, in part because of its wide host range. It has been reported by a number of authors as being of possible economic importance in several different plants, including dahlia and sunflower. discussion.

Neotephritis rava Foote (Figs. 288, 290; Map 40) Neotephritis rava Foote 1960f: 150 (female holotype, NMNH; Rustler Park, Portal, Ariz.). - Foote 1965a: 669 (in catalog). - Steyskal 1972c: 415 (in key, known species of Neotephritis). - Arnaud 1979: 330 (type data). N. rava differs from finalis in that the spots in the wing fuse to form more extensive hyaline areas at the apex of the pterostigma, at the apex of cell r2+3, and in cell m. The presence of many small hyaline areas in cells r2+3, r4+5, and cell dm, as well as a somewhat shorter oviscape (fig. 290) and a yellowish gray cast in the body color, also distinguish this species from finalis. recognition.

Map 40. We have seen specimens of Neotephritis resembling rava from Mexico but are uncertain whether they are conspecific. distribution.

host.

Not

known.

Genus Oedicarena Loew Oedicarena Loew 1873: 247 (type species, Trypeta tetanops Loew 1873: 245, by monotypy). - W. A. Snow 1894: 162 (taxonomy). - Williston 1896: 121 (in key). - Coquillett 1899c: 267 (in key). - F. H. Snow (part) 1903: 219 (Kans.). - Aldrich 1905: 604 (in catalog). - Williston 1908: 284 (in key). - Coquillett 1910: 578 (type data). - Curran 1932b: 3 (in key). - Curran 1934: 289 (in key). - Tiering 1940a: 4 (taxonomy). - Foote 1960h: 112, 114 (taxonomy, in key). - Foote 1965a: 675 (in catalog). - Steyskal and Foote 1977: 152, 153 (taxonomy, in key). - Berlocher and Bush 1982: 136 (taxonomy). - Norrbom et al. 1988: 98 (revi¬ sion). Trypeta (Oedicarena): Osten Sacken (part) 1878: 190 (in catalog). Spilographa Loew: Wulp (part) 1899: 406 (taxonomy). -Cresson 1907: 100 (taxon¬ omy). Phorellia Robineau-Desvoidy: Hendel (part) 1914c: 28 (review).

247

Oedicarena Goeden (1987) reported that finalis has been reared from the flower heads of many genera of the subtribe Verbesininae of the tribe Heliantheae, family Asteraceae. hosts.

N. finalis is one of the most commonly encountered tephritids in North America, in part because of its wide host range. It has been reported by a number of authors as being of possible economic importance in several different plants, including dahlia and sunflower. discussion.

Neotephritis rava Foote (Figs. 288, 290; Map 40) Neotephritis rava Foote 1960f: 150 (female holotype, NMNH; Rustler Park, Portal, Ariz.). - Foote 1965a: 669 (in catalog). - Steyskal 1972c: 415 (in key, known species of Neotephritis). - Arnaud 1979: 330 (type data). N. rava differs from finalis in that the spots in the wing fuse to form more extensive hyaline areas at the apex of the pterostigma, at the apex of cell r2+3, and in cell m. The presence of many small hyaline areas in cells r2+3, r4+5, and cell dm, as well as a somewhat shorter oviscape (fig. 290) and a yellowish gray cast in the body color, also distinguish this species from finalis. recognition.

Map 40. We have seen specimens of Neotephritis resembling rava from Mexico but are uncertain whether they are conspecific. distribution.

host.

Not

known.

Genus Oedicarena Loew Oedicarena Loew 1873: 247 (type species, Trypeta tetanops Loew 1873: 245, by monotypy). - W. A. Snow 1894: 162 (taxonomy). - Williston 1896: 121 (in key). - Coquillett 1899c: 267 (in key). - F. H. Snow (part) 1903: 219 (Kans.). - Aldrich 1905: 604 (in catalog). - Williston 1908: 284 (in key). - Coquillett 1910: 578 (type data). - Curran 1932b: 3 (in key). - Curran 1934: 289 (in key). - Tiering 1940a: 4 (taxonomy). - Foote 1960h: 112, 114 (taxonomy, in key). - Foote 1965a: 675 (in catalog). - Steyskal and Foote 1977: 152, 153 (taxonomy, in key). - Berlocher and Bush 1982: 136 (taxonomy). - Norrbom et al. 1988: 98 (revi¬ sion). Trypeta (Oedicarena): Osten Sacken (part) 1878: 190 (in catalog). Spilographa Loew: Wulp (part) 1899: 406 (taxonomy). -Cresson 1907: 100 (taxon¬ omy). Phorellia Robineau-Desvoidy: Hendel (part) 1914c: 28 (review).

Systematic Treatment of the Genera Rhagoletoides Foote 1960d: 145 (type species, Spilographa latifrons Wulp 1899: 407, by monotypy). — Foote 1960h: 114 (taxonomy). — Foote 1965a: 675 (in catalog). - Foote 1965b: 238 (type data). - Foote 1967b: 41 (in catalog, Neotropi¬ cal Region). — Steyskal and Foote 1977: 152 (synonymy). The body color of these medium-sized flies is orange to dark brown ventrally, sometimes darker brown dorsally. A whitish anepisternal stripe is recognition.

present, although it is weakly differentiated in light-colored specimens. The scutum is densely tomentose with bare spots at the insertions of the dorsocentral bristles and at the mesal ends of the transverse suture, and the scutellum sometimes has a diffuse apical spot. The dorsocentral bristles are situated behind the midpoint between the acrostichal and supra-alar bristles; the apical third of the middle and hind femora have short, sometimes stout, ventral bristles; the apex of the 1st flagellomere is rounded; the apex of cell cup is narrowly but distinctly extended; and an intercalary band is present in the wing pattern (see discussion of Rbagoletis wing bands; also fig. 291, a). The bands lying on veins r-m and dm-cu are often fully connected at the posterior margin of the wing as in fig. 293, c, unlike the condition shown in figs. 291, d and 292, c. The intercalary band, the location of the dorsocentral bristles, and the pattern of the scutal pruinosity distinguish this genus from Trypeta, Euleia, and Chetostoma, genera which closely resemble Oedicarena, and from all other genera occurring in the United States and Canada. Norrbom et al. (1988) pointed out that Oedicarena is the only genus of Tephritidae known to have four spermathecae. discussion.

The genus Oedicarena was recently revised by Norrbom et al.

(1988) to include five species from the western United States and the highlands of Mexico. O. latifrons, the type and sole species included in Rhagoletoides by Foote (1960d), has stronger femoral setae and a narrower gena than tetanops, the type species of Oedicarena. Steyskal and Foote (1977) concluded that these two species do not otherwise differ significantly and are properly congeneric. Norrbom et al. (1988) discussed details of the taxonomic characters, including the genitalia, and the rela¬ tionships of the genus at considerable length. Specimens of this genus are rarely encountered. One species has been bred from fruits of a species of Solanum and two others have been collected on Solanum species.

Key to U.S. Species of Oedicarena 1. Gena wide, at least half as high as eye (fig. 294, b); parafacial at narrowest point comparatively wide (fig. 294, a); wing with apical costal band about as wide in cell r2+3 as in cell r4+5 (fig. 291, b, c); apex of scutellum concolorous with base; aculeus short, less than two-thirds length of oviscape, tip broad, with small acute median elongation (fig. 297) . .persuasa (Osten Sacken) Gena narrow, one-third or less as high as eye (fig. 295, b); parafacial at narrowest point narrower (fig. 295, a); wing with apical costal band much broader in cell r4+5 than in cell r2 + 3 (fig. 293, b) or with separate

Figures 291—293. Right wings, Oedicarena spp. 291, persuasa (O.S.); 292, latifrons (Wulp); 293, beameri Norrbom and Ming. Figures 294, 295. Lateral view of head, Oedicarena spp. 294, persuasa (O.S.); 295, latifrons (Wulp). Figure 296. Dorsal view, scutellum, Oedicarena latifrons (Wulp). Figures 297—299. Aculeus tip, Oedicarena spp. 297, persuasa (O.S.); 298, latifrons (Wulp); 299, beameri Norrbom and Ming.

249

250

Systematic Treatment of the Genera

apical spot (fig. 292, b); apex of scutellum sometimes with diffuse dark spot (fig. 296, a); aculeus more than 9/io length of oviscape, tip more gradually tapering (figs. 298, 299) .2 2. Apical spot in cell r4+5 not connected to dark mark at apex of vein R2 + 3 (fig. 292, a); knob of halter yellow to light brown; aculeus tip gradually tapering and with a pair of small subapical steps, nonserrate (fig. 298) .latifrons (Wulp) Narrow costal band in cell r2 + 3 connecting dark marks in cell r4 + 5 and at apex of vein R2+3 (fig. 293, a); knob of halter brown; aculeus tip parallel¬ sided, then abruptly tapering, with fine serrations (fig. 299). .beameri Norrbom and Ming

Oedicarena beameri Norrbom and Ming (Figs. 293, 299; Map 41) Oedicarena beameri Norrbom and Ming in Norrbom et al. 1988: 107 (female holotype, UKL; Santa Rita Mts., Ariz.). Among the species of Oedicarena that occur in the United States, beameri most closely resembles latifrons in having a large compound eye, a narrow recognition.

gena which is distinctly less than half the height of the eye (fig. 295, b), a relatively narrow parafacial (fig. 295, a), often a dark spot at the apex of the scutellum (fig. 296), and well-differentiated short, stout anteroventral and posteroventral bristles on the middle and hind femora. O. beameri is distinguished from latifrons most easily by its wing pattern; in the latter species, the spot at the apex of cell r4+5 is never connected along the costa to the band lying upon vein dm-cu (fig. 292, a), whereas such a connection is always present in beameri (fig. 293, a). O. persuasa differs from beameri and latifrons in having a smaller compound eye, a gena wider than half the eye height (fig. 294, b), and weakly differentiated femoral bristles. The nature of the dark spot in the apex of cell r4+5 and its connection with the band lying upon vein dm-cu also distinguishes these two species. The aculeus tips of the three U.S. species (figs. 297-299) are distinctly different, as are the genitalia of the males. distribution.

Map 41. O. beameri is known from only two specimens collected

in Arizona. host.

Not

known.

Oedicarena latifrons (Wulp) (Figs. 31, 292, 295, 296, 298; Map 41) Spilographa obfuscata Wulp 1899: 406 (male holotype, BMNH; Omilteme, Guer¬ rero, Mex., 8,000 ft.). - Foote 1960d: 146 (taxonomy). - Foote 1965b: 238 (taxonomy, synonymy). Spilographa latifrons Wulp 1899: 407 (female holotype, BMNH; Ciudad, Durango,

Oedicarena

A

Oedicarena beameri

▲ Oedicarena latifrons o

Oedicarena persuasa

•

Orellia occidentalis

Map 41. Distribution of Oedicarena beameri, O. latifrons, O. persuasa, and Orellia occidentalis.

Mex., 8,100 ft.). — Aldrich 1905: 604 (in catalog). — Foote 1965b: 238 (tax¬ onomy). Phorellia latifrons: Hendel 1914c: 28 (taxonomy). — Aczel 1949: 250 (in catalog). Phorellia obfuscata: Hendel 1914c: 28 (taxonomy). — Aczel 1949: 250 (in catalog). Rbagoletoides latifrons: Foote 1960d: 146 (review). — Foote 1965a: 675 (in catalog). — Foote 1967b: 41 (in catalog, Neotropical Region). Oedicarena latifrons: Steyskal and Foote 1977: 153 (taxonomy, in key). — Foote 1980: 38 (taxonomy). — Norrbom et al. 1988: 104 (revision). O. latifrons most closely resembles beameri in a number of characters, including a large compound eye, a comparatively narrow gena (fig. 295, recognition.

Systematic Treatment of the Genera

b), a narrow parafacial (fig. 295, a), well-differentiated short, stout femoral spines, and often a dark spot at the apex of the scutellum (fig. 296, a). The most obvious character separating latifrons from beameri and persuasa is the nature of the wing pattern, in which the spot occupying the apex of cell r4+5 in latifrons is always separated from the band lying on vein dm-cu by a hyaline area at the costa (fig. 292, a). The slender, elongated aculeus tip (fig. 298) and the male genitalia also distin¬ guish latifrons from the other two U.S. species. Map 41. O. latifrons was originally described from Mexico. It is widespread in the Mexican highlands. distribution.

This species has been reared from the fruit of a species of Solanum (Norrbom et al. 1988). host.

Oedicarena persuasa (Osten Sacken) (Figs. 291, 294, 297; Map 41) Trypeta (Oedicarena) persuasa Osten Sacken 1877: 344 (male lectotype, MCZ; Denver, Colo.). - Osten Sacken 1878: 190 (in catalog). - Norrbom et al. 1988: 113 (lectotype designation). Spilographa (Oedicarena) persuasa: Wulp 1899: 406 (taxonomy). Oedicarena persuasa: F. H. Snow 1903: 219 (Kans.). - Foote 1960h: 114 (in key). Foote 1965a: 675 (in catalog). — Steyskal and Foote 1977: 153 (in key). Norrbom et al. 1988: 113 (revision). Spilographa persuasa: Aldrich 1905: 604 (in catalog). recognition.

O. persuasa is most closely related to tetanops, the type species of

Oedicarena, a Mexican species (Norrbom et al. 1988). In both of these species, the gena is comparatively wide (fig. 294, b), as is the parafacial at its narrowest point (fig. 294, a; see key to species for details), and the scutellum is concolorous without any apical darkening. O. persuasa differs from tetanops in that the apical dark mark of the wing is crescentic in shape and connected to the band lying on vein dm-cu by a dark area along the costa of about the same width (fig. 291, b, c). These characters, in addition to the distinctive aculeus tip, distinguish persuasa from the other two Oedicarena species occurring in the United States. The body vestiture and the apically rounded 1st flagellomere separate this species from Rhagoletis basiola and both American species of Zonosemata, all three of which it resembles closely in wing pattern. DISTRIBUTION.

hosts.

Not known, but probably Solanum spp.

discussion.

the field.

Map 41.

O. persuasa is a poorly known species that is rarely encountered in

253

Ore Ilia

Genus Orellia Robineau-Desvoidy

Sitarea Robineau-Desvoidy 1830: 763 (type species, Sitarea scorzonerae RobineauDesvoidy, by designation of Coquillett 1910: 606). - White 1986: 149 (syn¬ onymy). Orellia Robineau-Desvoidy 1830: 765 (type species, Orellia flavicans RobineauDesvoidy, by monotypy) (= punctata (Schrank)). - Coquillett 1910: 578 (type data). - Cresson 1914b: 323 (taxonomy). - Curran 1934: 291 (in key). McFadden and Foote 1960: 253 (revision). - Foote and Blanc 1963: 38 (review, Calif.). - Foote 1965a: 671 (in catalog). - Wasbauer 1972: 121 (hosts). - Foote and Freidberg 1980: 31 (type data). - Foote 1984: 104 (in catalog). Trypeta Meigen [misidentification]: Loew (part) 1873: 254, 256 (taxonomy). Williston (part) 1896: 121 (in key). - Coquillett (part) 1899c: 262 (taxonomy). Aldrich (part) 1905: 604 (in catalog). - Williston (part) 1908: 286 (in key). Tryreta [error]: Loew (part) 1873: 329 (taxonomy). Trypeta (Trypeta) [misidentification]: Osten Sacken 1878: 190 (in catalog). Terellia Robineau-Desvoidy [misidentification]: V. T. Phillips (part) 1923: 122, 138 (in key, revision). — Curran (part) 1932b: 3 (in key). - Curran (part) 1934: 289 (in key).

Medium-sized yellowish flies, the body marked with black and the wing with light to dark brown bands or spots. The frons is bare; there are two or three pairs of frontal bristles and two pairs of orbital bristles, the posterior pair of which are well developed and strongly convergent (fig. 27, a); two pairs of scutellar bristles; and the single pair of dorsocentral bristles is situated on a transverse line connecting the supra-alar bristles. The genus is similar to Neaspilota in general recognition.

habitus and in having convergent posterior orbital bristles, but individuals of the latter are smaller, have almost entirely unmarked wings except in the vicinity of the pterostigma, and usually a haired frons. Cbaetostomella also is very similar to Orellia, but in the former genus, a group of rather well-developed setae is present on the anterior oral margin of the head.

Orellia, Cbaetostomella, and Neaspilota, quite similar in many re¬ spects, are the only three U.S. and Canadian genera belonging to the tribe Terelliini, which is characterized by the posterior orbital bristles being strongly convergent and the scutum with a dark, typically lyre-shaped pattern (Freidberg 1985). Only one species, palposa, has ever been seen from south of the United States; it was collected discussion.

near Guadalajara. Korneyev (1985, 1987) revised the classification of the Terelliini of the USSR and transferred most species previously in Orellia, including the three New World species, to the subgenera Terellia Robineau-Desvoidy and Cerajocera Rondani of Terellia. These changes came to our attention too late to modify the taxonomic arrangement of this handbook.

Systematic Treatment of the Genera

Key to U.S. and Canadian Species of Orellia 1. Wing with apical costal band and preapical band (covering vein dm-cu) never connected, each usually also interrupted by various hyaline areas in cells rls r2+3, and r4+5 (fig. 300, a) .ruficauda (Fabricius) Apical and preapical bands complete and usually united with each other in cell r1 (fig. 301, a) .2 2. A dark band lying upon vein CuA} extending from vein bm-cu to past mid¬ point of vein CuAj and usually joining posterior end of preapical band (fig. 301, b); .palposa (Loew) No dark band lying upon vein CuAj in the position described above; wing pattern as in fig. 302 .occidentals (Snow)

Orellia occidentalis (Snow) (Figs. 47, 302; Map 41) Trypeta occidentalis Snow 1894: 163 (female lectotype, UKL; Colo.). - Coquillett 1899c: 262 (synonymy). - Doane 1899: 179 (taxonomy). - Doane 1900: 47 (taxonomy). - Johnson 1903b: 106 (N. Mex.). - C. F. Baker 1904: 30 (Calif., Nev.). - Aldrich 1905: 605 (in catalog). - Janes and Thomas 1932: 103 (Utah). Strickland 1938: 204 (Alta.).-V. T. Phillips 1946: 127 (host). - Foote 1960b: 254 (taxonomy). — McFadden and Foote 1960: 259 (lectotype designation). — Byers et al. 1962: 181 (type data). Trypeta straminea Doane 1899: 179 (male lectotype, WSU; Pullman, Wash.). Coquillett 1899c: 262 (synonymy). - Doane 1900: 47 (taxonomy). - Aldrich 1905: 605 (in catalog). - Grossbeck 1912: 378 (type data). - McFadden and Foote 1960: 254 (synonymy). - Byers et al. 1962: 181 (type data). - Foote 1962: 177 (lectotype designation). - Foote 1966b: 125 (type data). - Zack 1984: 33 (type data).

302 Figures 300-302. Right wings, Orellia spp. 300, ruficauda (F.); 301, palposa (Lw.); 302, occiden¬ talis (Snow).

255

Ore Ilia Orellia occidentalis: Foote 1960b: 254 (taxonomy). — McFadden and Foote 1961: 258 (review). - Foote 1962: 177 (taxonomy). — Foote and Blanc 1963: 38 (review, Calif.). — Foote 1965a: 671 (in catalog). — Wasbauer 1972: 121 (hosts). — Lamp and McCarty 1979: 117 (biology, hosts). - Steck 1981: 124 (biology). - Lamp and McCarty 1982: 847 (seed predation). — Pemberton et al. 1985: 972 (hosts). Goeden and Ricker 1986b: 951 (host). - Goeden and Ricker 1986c: 957, 958 (hosts). 0. occidentalis and palposa have similar wing patterns, with dark bands rather than discrete spots as in ruficauda. Unlike palposa, the discal band (fig. 302, a) in occidentalis proceeds directly and uninterrupted across the wing into cell cual5 and vein CuAj is never covered by an elongated dark band or stripe. Further, the anterior connection between the apical and subapical bands is sometimes inter¬ recognition.

rupted by a hyaline area at the costa, and the wing bands generally are usually a paler brown. 0. occidentalis most closely resembles Cbaetostomella undosa, but unlike occidentalis, the wing of undosa is more darkly marked, a black spot is present on the scutellum, some of the postocular setae are black, and distinct dark setae are present laterally on the gena close to the oral margin. DISTRIBUTION.

Map 41.

O. occidentalis was recorded from about 10 species of Cirsium by Pember¬ ton et al. (1985). Foote and Blanc (1963) reported Cnicus parryi as a host. hosts.

O. occidentalis is encountered much more commonly than any oth¬ er species of Orellia in the United States. discussion.

Orellia palposa (Loew) (Figs. 44, 48, 301; Map 42) Trypeta palposa Loew 1862c: 74 (male holotype, MCZ; n. Wis.). — Loew 1873: 253 (review). — W. A. Snow 1894: 163 (taxonomy). — Doane 1899: 179 (distribution). — Johnson 1900b: 687 (N.J.). — Aldrich 1905: 605 (in catalog). — Johnson 1909: 113 (distribution). — Johnson 1910: 801 (N.J.). — Rosewall 1922: 176 (host). — V. T. Phillips 1946: 92, 127 (description, biology larva; hosts). — Foote 1960b: 254 (taxonomy). Trypeta sp.: Osten Sacken 1877: 345 (taxonomy). Trypeta (Trypeta) palposa: Osten Sacken 1878: 190 (in catalog). Terellia palposa: V. T. Phillips 1923: 138 (review). — Johnson 1925b: 262 (N. Engl.). — Sturtevant 1925: 215 (seminal receptacles).—Johnson 1930: 151 (Nantucket).Brimley 1938: 384 (N. Car.). - Strickland 1938: 204 (Alta.). - Ouellett 1941: 130 (Que.). — Hennig 1952: 218 (in catalog immature stages). — Steyskal 1957: 94 (eye color). - Foote 1960b: 254 (taxonomy). Orellia palposa: McFadden and Foote 1960: 260 (revision). — Foote 1965a: 671 (in catalog). — Wasbauer 1972: 121 (hosts). — Steck 1981: 36 (biology).

Systematic Treatment of the Genera

Orellia palposa Orellia palposa (state record only)

Map 42. Distribution of Orellia palposa.

O. palposa closely resembles occidentals in that the dark mark¬ ings on the wing form bands rather than separated spots as in ruficauda. 0. palposa RECOGNITION.

can always be readily recognized by the presence of an elongated dark band lying upon vein CuA1 (fig. 301, b), which usually connects with the posterior end of the subapical band. In many specimens, this band is lightly or entirely separated from the subapical band near the apex of cell cual5 and it seldom connects with the discal band (the band covering vein r-m). The bands in palposa are usually darker than in occidentalis. Map 42. O. palposa is the only species of Orellia we have seen from south (Mexico) of the United States. distribution.

Ore Ilia

hosts.

This species is apparently restricted to Cirsium.

Judging from the number of specimens in museum collections, palposa is a much less commonly encountered species than either ruficauda or occidendiscussion.

talis.

Orellia ruficauda (Fabricius) (Fig. 300; Map 43) Musca ruficauda Fabricius 1794: 353 (type lost, sex not known; France). Trypeta florescentiae [misidentification]: Loew 1873: 254 (review). — Flarrington 1893: 85 (damage, parasite). — W. A. Snow 1894: 159 (distribution). — Aldrich

Map 43. Distribution of Orellia ruficauda.

258

Systematic Treatment of the Genera

1905: 605 (in catalog). — Detmers 1927: 26 (biology, parasite). — V. T. Phillips 1946: 90, 127 (description, biology larva; hosts). — Hennig 1952: 218 (in catalog immature stages). Trypeta (Trypeta) florescentiae [misidentification]: Osten Sacken 1878: 191 (in catalog). Dipterous larva: Ashmead 1892: 309 (parasite). Trypeta ruficauda: Aldrich 1905: 605 (in catalog). — Johnson 1909: 113 (distribu¬ tion). — Winn and Beaulieu 1915: 153 (Que.). — Marcovitch 1916: 150 (biology, description larva). - V. T. Phillips 1946: 127 (hosts). — Foote 1960b: 254 (tax¬ onomy). Terellia florescentiae [misidentification]: V. T. Phillips 1923: 139 (review). — Johnson 1925b: 262 (N. Engl.). — Johnson 1927: 218 (Mt. Desert). — Leonard 1928: 852 (N.Y.). - Brimley 1938: 384 (N. Car.). - Tonks 1953: 28 (B.C.). - Muesebeck 1958: 34 (parasite). Terellia ruficauda: V. T. Phillips 1923: 139 (synonymy). — Petch and Maltaise 1932: 56 (Que.). Orellia ruficauda: Hendel 1927: 135 (review). - Hennig 1952: 214 (in catalog immature stages). - Foote 1960b: 254 (taxonomy). - McFadden and Foote 1960: 255 (revision). - Foote and Blanc 1963: 39 (review, Calif.). - Foote 1965a: 671 (in catalog). - Peschken 1971: 79 (host, economic importance). - Wasbauer 1972: 121 (hosts). — Steck 1981: 6 (hosts). Terellia floriscentiae [error, misidentification]: Curran 1934: 288 (figure of wing). Orella [error] ruficauda: Forsell 1947: 167 (Puget Sound).

recognition.

Of the three species of Orellia treated herein, ruficauda is the most

distinctive in that its wing pattern, unlike those of the other two species, comprises extensive dark spots rather than bands. The dark spot covering vein dm-cu (fig. 300, b) is often darker than shown; in some specimens, it is connected to the dark mark descending from the costa near the center of cell rt to form a subapical band as in occidentalis or palposa. The discal band never attains vein r-m, and the subbasal band is indistinct in all specimens. The species may be further recognized by faint clouding, separated from the dark markings by lighter hyaline areas, in the hyaline areas of the wing.

Map 43. O. ruficauda occurs throughout the northern parts of the Palearctic Region from western Europe to the easternmost parts of the former USSR. distribution.

Cirsium arvense is the only host recorded by Wasbauer (1972) in North America for this species. host.

O. ruficauda was probably accidentally introduced into North America; its only reported host here is an introduced Palearctic thistle. discussion.

259

Oxyna

Genus Oxyna Robineau-Desvoidy Oxyna Robineau-Desvoidy 1830: 755: (type species, Oxyna flavescens RobineauDesvoidy 1830: 756, by designation of Hendel 1914b: 96) (= flavipennis (Loew)). -Aldrich 1905: 609 (in catalog). - Hendel 1914b: 96 (type designation, in key). Quisenberry 1949a: 71 (revision). — Foote 1960f: 146 (taxonomy). - Foote and Blanc 1963: 40 (review, Calif.). — Foote 1965a: 664 (in catalog). — Wasbauer 1972: 121 (hosts). - Foote and Freidberg 1980: 32 (type data). - Freidberg 1984: 133 (galls). recognition. The wing patterns in Oxyna most closely resemble those of

Tepbritis, Dioxyna, and Paroxyna in exhibiting whitish spots distributed in a dark field. The genus is always recognizable by the presence of only a single pair of black frontal bristles, a character not present in any other North American fruit fly genus. These dark frontals are always accompanied by a small group of slightly more robust, light-colored setae situated on a level with the dark bristle or anterior to it, or one light-colored seta anterior to and one posterior to the dark bristle; at least one of these light bristles is usually almost as long as the dark one (fig. 87, a, b). The labellum is elongated and reflexed as in Gonioxyna, Dioxyna, and Paroxyna, but the head is distinctly higher than long and differently shaped in lateral view in contrast to the condition in those genera. Two pairs of scutellar bristles are present, and the dorsocentral bristles are situated very close to the suture. discussion. Oxyna is known in Mexico by only a single specimen; apparently it

does not occur farther south in the New World. Twenty species are known from the Palearctic Region, where the genus is widespread. Korneyev (1989) recently mono¬ graphed the eastern Palearctic species. The three known species in the United States and Canada, which are rarely encountered, fit Korneyev’s diagnosis for the genus except that in utahensis, the frons is sometimes setulose. Many of the Palearctic species, like palpalis and utahensis, breed in Artemisia.

Key to U.S. and Canadian Species of Oxyna 1. Wing with 2 dark, distinct, broad, oblique bands not connected by dark markings (fig. 303, b); no dark markings present in cell be (fig. 303, a); scutellum pale yellow to nearly white, at least ventrally. .palpalis (Coquillett) Wing indistinctly banded (figs. 304, 305), the hyaline area between the bands comprising coalesced hyaline spots (fig. 304, a); dark markings present in cell be (fig. 305, a); scutellum dark yellow to brown .2 2. Femora black except narrowly at apices; wing pattern as in fig. 304 . .utahensis Quisenberry Femora yellowish with a black spot or stripe at base ventrally and some¬ times some black dusting dorsally; wing pattern as in fig. 305 . .aterrima (Doane)

Systematic Treatment of the Genera

a

Figures 303-305. Right wings, Oxyna spp. 303, palpalis (Coq.); 304, utahensis Quis.; 305, ater¬ rima (Doane).

Oxyna aterrima (Doane) (Fig. 305; Map 44) Eurosta aterrima Doane 1899: 187 (female holotype, WSU; Colo.). - Aldrich 1905: 609 (in catalog). - Foote 1966b: 122 (type data). - Zack 1984: 32 (type data). Tephritis aterrima: Knowlton and Harmston 1937: 145 (Utah). Oxyna aterrima: Quisenberry 1949a: 73 (revision). - Foote and Blanc 1963: 40 (review, Calif.). - Foote 1965a: 665 (in catalog). The wing pattern of aterrima most closely resembles that of utah¬ ensis in that the transverse hyaline area distad of vein dm-cu extending from the costa recognition.

to the posterior margin comprises about eight partially merged hyaline spots with some dark areas remaining between them. O. aterrima differs from utahensis in having the femora yellow, except for the black spots ventrally and occasionally black dusting dorsally rather than black, except at the apices, as in utahensis. distribution.

host.

Not

Map 44.

known.

Oxyna palpalis (Coquillett) (Figs. 87, 303; Map 44) Tephritis palpalis Coquillett in Baker 1904: 30 (male holotype, NMNH; Ormsby County, Nev.). Oxyna palpalis: Quisenberry 1949a: 76 (revision). - Quisenberry 1951: 59 (taxon¬ omy). - Foote and Blanc 1963: 41 (review, Calif.). - Foote 1965a: 665 (in catalog). - Novak et al. 1967: 147 (host, gall). - Wasbauer 1972: 121 (host).

261

Oxyna

•

Oxyna aterrima

•

, - -,

•

Oxyna palpalis

•

Oxyna utahensis

Map 44. Distribution of Oxyna aterrima, O. palpalis, and O. utahensis.

O. palpalis may be distinguished from aterrima and utahensis by the wing pattern. In palpalis, the light area from the costa to the posterior wing margin distad of vein dm-cu has no dark markings within its borders, in contrast to that stripe in the other two species, as described for aterrima. recognition.

distribution.

host.

Map 44.

Novak et al. (1967) described a gall on Artemisia tridentata made by this

species.

Oxyna utahensis Quisenberry (Fig. 304; Map 44) Oxyna utahensis Quisenberry 1949a: 73 (male holotype, AMNH; Temple Fork, Logan Canyon, Utah). - Foote and Blanc 1963: 41 (review, Calif). - Foote 1965a: 665 (in catalog).

recognition.

See recognition characters of aterrima.

distribution.

Map 44.

A female has been reared from Artemisia tridentata var. vaseyana. One female (NMNFI) was reared from a gall by R. G. Jones in Idaho. host.

262

Systematic Treatment of the Genera

Genus Paracantha Coquillett Carphotricha Loew 1873: 279, 328 (taxonomy). - Williston 1896: 122 (in key). — Aldrich 1905: 608 (in catalog). - Williston 1908: 287 (in key). - Cresson 1914a: 277 (type designation). - Cresson 1914b: 323 (nomenclature). - Rosewall 1922: 176 (host). — F. H. Benjamin 1934: 30 (taxonomy). Carpotricha [error]: Loew (part) 1873: 279, 328 (taxonomy). - V. T. Phillips 1923: 141 (synonymy). Trypeta (Carphotricha): Osten Sacken 1878: 192 (in catalog). Scriptotricha [error]: Cockerell 1889a: 1 (biology). - Cockerell 1889b: 106 (biology, Colo.). — Cockerell 1900b: 400 (nomenclature). Paracantha Coquillett 1899c: 264 (type species, Trypeta culta Wiedemann 1830: 486, 680, by original designation). - Aldrich 1905: 608 (in catalog). - Coquillett 1910: 583 (type data). - Cresson 1914a: 277 (type data). - Cresson 1914b: 323 (type data, taxonomy). - Hendel 1914b: 93 (in key). - V. T. Phillips 1923: 122, 141 (review, in key). - Curran 1932b: 4 (in key). - F. H. Benjamin 1934: 29 (review, Fla.). - Curran 1934: 291 (in key). - Hering 1940d: 53 (key to species). Malloch 1941: 33 (revision). - Aczel 1952b: 199 (revision, Neotropical). - Foote and Blanc 1963: 41 (review, Calif.). - Foote 1965a: 662 (in catalog). - Wasbauer 1972: 121 (hosts).

A genus of comparatively large yellow flies with a brownish wing extensively margined with hyaline incisions, with small hyaline spots, and with an recognition.

extremely strong bulla in cell r4+^. The head is unusual in having three pairs of white to yellowish orbital bristles situated well inside the frontals and the face usually with a black spot in the center (best visible from front view) and with a pair of black spots on its ventrolateral corners (fig. 312, b). There is a distinct black spot on the parafacial beside the antenna (fig. 312, c) and another on the occiput behind the eye (fig. 312, d). The frons is broad with three pairs of frontal bristles, the anterior pair short and pale. The surface of the scutum bears white to yellowish setae in patches, leaving darker brown, heavily tomentose areas. There is one pair of dorsocentral bristles situated closer to the transverse suture than to a line between the supra-alar bristles, and two pairs of scutellar bristles, the apical pair inserted in a pair of discrete black subapical spots. The middle and hind femora each usually bear two posterior black marks. With this combination of characters and the nature of the wing pattern (figs. 306—310), this genus cannot be confused with any other occurring north of Mexico.

Paracantha occurs throughout most of the New World but is not found elsewhere. Its closest relatives appear to be Neotropical, although in Africa and the Oriental Region there are other genera that may be related. The North discussion.

American species breed in flowers of Cirsium or in several genera of Heliantheae, including sunflowers. Headrick and Goeden (1989, 1990a,b,c) studied in great detail the biology of gentilis, one of the thistle-breeding species.

Figures 306—310. Right wings, Paracantha spp. 306, cultaris (Coq.); 307, genalis Mall.; 308, forficula Benj.; 309, culta (Wied.); 310, gentilis Her. Figures 311-313. Lateral view of head, Paracantha spp. 311, genalis Mall.; 312, culta (Wied.); 313, gentilis Her. Figures 314, 315. Posteri¬ or surface of fore femur, Paracantha spp. 314, forficula Benj.; 315, gentilis Her.

263

264

Systematic Treatment of the Genera

Key to U.S. and Canadian Species of Paracantha 1. Posterior dark ray at apex of cell r4+5 terminating at apex of vein M (fig. 306, b); apex of cell r2+3 with 1-2 dark rays (fig. 306, a) . .cultaris (Coquillett) Posterior dark ray at apex of cell r4+5 terminating anterior to apex of vein M (fig. 307, b); apex of cell r2+3 with 2-3 dark rays terminating anterior to apex of vein M in cell r4 + 5 (fig. 307, a) .2 2. A group of 6—10 stout, black bristles situated immediately anterior to genal bristle (fig. 311, a); dark spot on lower corner of face, if present, large, bordering the group of black bristles; wing pattern as in fig. 307 . .genalis Malloch At most 2 or 3 such black bristles in this position, usually none; dark spot on lower corner of face, if present, smaller, distinctly separated from any black bristles .3 3. Apex of cell r4+5 with 3 dark rays, all 3 equally dark (fig. 308, a); 2 black spots on posterior surface of fore femur (fig. 314); oviscape less than 0.75 times as long as mesonotum.forficula Benjamin Apex of cell r4 + 5 with only 2 dark rays or, if a third ray is present, it is much weaker than the other 2 (compare figs. 308-310); usually only 1 black spot, or none, on posterior surface of fore femur (fig. 315); oviscape more than 0.75 times as long as mesonotum.4 4. Black parafacial spot between base of antenna and eye large, elongate, or triangular (fig. 312, c); wing pattern as in fig. 309 . .culta (Wiedemann) Black parafacial spot small, subcircular (fig. 313, a); wing pattern as in fig. 310 .gentilis Hering

Paracantha culta (Wiedemann) (Figs. 88, 309, 312; Map 45) Trypeta cutta [error]: Wiedemann 1830: 486 (female syntypes, prob. NMW; Ga.). Osten Sacken 1858: 79 (in catalog). Trypeta culta Wiedemann 1830: 680 (error correction). — Loew 1862c: 94 (review). - Loew 1873: 276 (review). - Coquillett 1910: 583 (type data). - Cresson 1914a: 277 (taxonomy). Tephritis marginepunctata Macquart 1835: 464 (female syntypes, MNHN, proba¬ bly lost; Pa.). New synonymy. Acinia fimbriata Macquart 1843a: 385 (1843b: 228) (male syntype(s), poss. MHNL; “Carolina”). — Loew 1862c: 58 (synonymy). — Osten Sacken 1878: 192 (in catalog). - Aldrich 1905: 608 (in catalog). Trypeta sarnia Walker 1849: 1029 (female syntypes, BMNH; probably U.S.). — Hardy 1959: 223, 235 (synonymy). - Foote 1964d: 323 (synonymy, taxonomy). Trypeta marginepunctata: Osten Sacken 1858: 77 (in catalog). - Osten Sacken 1862: 97 (description of adult). - Loew 1873: 337 (taxonomy). - Osten Sacken 1878: 195 (in catalog).

265

Paracantba

•

Paracantba culta

★

Paracantba culta (state record only)

o

Paracantba cultaris

☆ Paracantba cultaris (state record only)

Map 45. Distribution of Paracantba culta and P. cultaris.

Trypeta fimbriata [misidentification]: Osten Sacken 1858: 79 (in catalog). - Loew 1862c: 58 (synonymy). - Loew 1873: 336 (synonymy). Carpotricba [error] culta: Loew 1873: 279 (distribution). Carpbotricba culta: Loew 1873: 329, 336 (taxonomy). - W. A. Snow 1894: 169 (distribution). - Johnson 1895b: 337 (Fla.). - Doane (part) 1899: 185 (taxon¬ omy). - F. H. Snow 1904: 219 (Ariz.). - Aldrich 1905: 608 (in catalog).-Johnson 1913b: 85 (Fla.). - Cole and Lovett 1921: 326 (Oreg.). Trypeta (Carpbotricba) culta: Osten Sacken 1878: 192 (in catalog). — Cockerell 1898: 155 (N. Mex.). Scriptotricba culta: Cockerell 1889a: 1 (biology). — Cockerell 1893b: 369 (Colo.). — Cockerell 1899: 91 (gall, Colo.).

Systematic Treatment of the Genera

Carphotriche [error] culta: Coquillett 1899c: 264 (taxonomy). Paracantha culta: Coquillett 1899c: 264 (taxonomy). — Johnson 1903b: 106 (N. Mex.). — Hendel 1914b: 93 (type data). — Hendel 1914c: 50 (review, Neotropical). — V. T. Phillips (part) 1923: 142 (review). — F. H. Benjamin 1934: 30 (review, Fla.). - Knowlton and Harmston 1937: 145 (Utah). — Essig 1938: 605 (taxonomy). Strickland 1938: 204 (Alta.). — Hering 1940d: 53 (in key). — Malloch 1941: 35 (review). - V. T. Phillips 1946: 55, 116 (description, biology larva; hosts). Ryckman 1951: 127 (host, damage). — Aczel 1952b: 217 (review). — Hennig 1952: 210 (in catalog immature stages). - Hardy 1959: 223 (synonymy). - Foote 1965a: 662 (in catalog). — Wasbauer 1972: 121 (hosts.). — Morihara and Balsbaugh 1976: 692 (S. Dak., host). - Lamp and McCarty 1979: 117 (biology, host). - Batra et al. 1981: 75 (host). — Steck 1981: 19 (hosts). — Lamp and McCarty 1982: 847 (seed predation). - Elzinga and Broce 1986: 208 (labellar morphology). Carphotricha (Paracantha) culta: Washburn 1905: 118 (Minn.). Carphotricba marginepunctata: Aldrich 1905: 609 (in catalog). Paracantha marginepunctata: V. T. Phillips 1923: 142 (taxonomy). - Foote 1965a: 662 (in catalog). Acinia fimbrata [error]: Malloch 1941: 35 (synonymy). recognition.

P. culta is similar to gentilis in head and body markings, wing

pattern, and in the possession of a comparatively long oviscape. Both species often possess a dark spot on the lower frons between the antennal bases, a smaller dark spot on either side of the latter at the base of each antenna, and usually only one (fig. 315) or no black spot on the posterior surface of the fore femur. These two species are distinguished from each other by the size of the black spot on the parafacial between the base of the antenna and the eye; that of culta is quite extensive (fig. 312, c), whereas that of gentilis is distinctly smaller (fig. 313, a). The characters presented in the key to species are adequate to distinguish culta from the other Paracantha species discussed in this handbook. distribution.

Map 45. Records from states on the West Coast are errors.

Wasbauer (1972) reported culta from Helianthus biennis (possibly a misidentification of cultaris) and several species of Cirsium. He also indicated it has been found in Cynara scolymus in California, but that record is in error. hosts.

We have included references to Macquart’s species marginepunctata in the synonymy of culta because that species was described from a specimen ori¬ ginating in Pennsylvania and because culta is the only American species of the genus known to occur in the northeastern United States. discussion.

Paracantha cultaris (Coquillett) (Figs. 89, 306; Map 45) Trypeta (Carphotricha) cultaris Coquillett 1894: 72 (male and female syntypes, NMNH; so. Calif.).

267

Faracantba

Trypeta culta [misidentification]: Doane 1899: 185 (taxonomy). Carphotricha culta [misidentification]: Wulp 1900: 422 (synonymy, distribution). Carpbotricba cultaris: Aldrich 1905: 608 (in catalog). Carpbotricbia [error] culta [misidentification]: Woodworth 1913: 135 (Calif.). Carpbotricba culta [misidentification]: Woodworth 1913: 137 (Calif.). Faracantba cultaris: F. H. Benjamin 1934: 30 (taxonomy). - Hering 1940d: 50 (in key). - Malloch 1941: 36 (review). - Aczel 1949: 260 (in catalog). - Aczel 1952b: 221 (revision). — Foote and Blanc 1963: 42 (review, Calif.). — Foote 1965a: 662 (in catalog). - Wasbauer 1972: 122 (hosts). - Flilgendorf and Goeden 1981: 106 (host). - Cavender and Goeden 1984: 213 (biology, host). As indicated in our key to species, cultaris is distinguished from all Faracantba species occurring north of Mexico by the position of the ray bounding the posterior margin of the hyaline triangle in the apex of cell r4+5; it terminates on vein M rather than within the preceding cell (fig. 306, b). The wing usually has only two recognition.

hyaline incisions in the apex of cell r2+3, but occasionally a dark ray partially or completely divides the apical incision (fig. 306, a). P. cultaris often has a black spot on the lower frons between the antennal bases and only one black spot on the posterior surface of the fore femur near its apex, which in some specimens is mir¬ rored by a black spot on the anterior surface in a similar position. Map 45. P. cultaris occurs through Mexico south to Costa Rica and questionably in Fiaiti. distribution.

P. cultaris was recorded by Wasbauer (1972) from Heliantbus; his records of Borricbia and Cirsium are in error. Cavender and Goeden (1984) reared this fly from flower heads of Heliantbus annuus ssp. lenticularis and H. annuus var. mac¬ hosts.

rocarpus.

Paracantha forficula Benjamin (Figs. 308, 314) Faracantba culta [misidentification]: V. T. Phillips (part) 1923: 142 (review). Faracantba forficula Benjamin 1934: 31 (male holotype, NMNH; Cocoa Beach, Fla.). — Hering 1940b: 54 (in key). — Malloch 1941: 37 (revision). — V. T. Phillips 1946: 116 (host). — Aczel 1952b: 210 (revision). - Hennig 1952: 210 (in catalog immature stages). - Foote 1965a: 662 (in catalog). - Stegmaier 1967b: 157 (biolo¬ gy, hosts). — Bush and Huettel 1970: 91 (host). — Wasbauer 1972: 122 (host). Among the species of Faracantba occurring north of Mexico, for¬ ficula is distinguished primarily by the nature of the hyaline triangle in the apex of cell r4+5, which is divided longitudinally by a dark, narrow ray through its center (fig. 308, b); except in some specimens of genalis and in the other U.S. and Canadian species, this triangle is entirely hyaline or has only a faint triangular medial mark much lighter than the usual two rays. This species has three or four hyaline incisions recognition.

268

Systematic Treatment of the Genera

•

•

Paracantha genalis

Map 46. Distribution of Paracantha genalis and P. gentilis.

in the apex of cell r2+3, and it lacks the group of black genal setae characteristic of genalis. In all specimens we have seen, two dark spots are present on the posterior surface of the fore femur (fig. 314). The oviscape is much shorter than in cult a, genalis, and gentilis. distribution.

Not mapped. P. forficula has been found in at least five Florida

locations. P. forficula was reared from flower heads of Borrichia frutescens collected at the type locality of Cecidocharella borrichia (Bush and Huettel 1970). host.

Paracantha genalis Malloch (Figs. 96, 102, 307, 311; Map 46) Paracantha culta [misidentification]: Lange 1941: 56 (host). Paracantha genalis Malloch 1941: 40 (male holotype, NMNH; Halfmoon Bay, Calif.). - Aczel 1952b: 230 (revision). - Foote and Blanc 1963: 42 (review, Calif.). - Foote 1965a: 662 (in catalog). - Pemberton et al. 1985: 792 (hosts). - Goeden and Ricker 1986c: 957 (host). The wing of genalis is generally darker than that of the other species of Paracantha treated herein. The wing pattern differs from those of culta and recognition.

gentilis, which it most closely resembles, by the presence in many specimens of a longitudinal division of the distalmost hyaline wedge in the apex of cell r2+3 (fig.

Paracantha 307); this incision also is sometimes divided in gentilis. P. genalis may invariably be distinguished from all other Paracantha occurring north of Mexico by the presence of a group of stout black setae between the black spot on the lower corner of the face and the genal bristle (fig. 311, a). There is a variable black spot on the lunule between the antennal bases, and the blackish spot on the lower corner of the face is larger than in the other species. The base of the fore femur may be darkened in some specimens, giving the appearance of two black spots as shown in fig. 314. distribution.

Map 46.

P. genalis was reared from three species of Cirsium by Pemberton et al. (1985). Goeden and Ricker (1986c, 1987a) reported it from Cirsium californica and C. congdonii. hosts.

Paracantha gentilis Hering (Figs. 310, 313, 315; Map 46) Paracantha gentilis Hering 1940d: 54 (male and female syntypes, BMNH; Hunters Cr., Wyo.). - Aczel 1952b: 233 (revision). - Foote and Blanc 1963: 43 (review, Calif.). — Foote 1965a: 662 (in catalog). — Wasbauer 1972: 122 (hosts). - Steck 1981: 114 (biology). - Pemberton et al. 1985: 793 (hosts). - Goeden and Ricker 1986b: 948, 951 (hosts). — Goeden and Ricker 1986c: 957, 958 (hosts). Headrick and Goeden 1989: 594 (parasite, hosts). - Headrick and Goeden 1990a: 220 (description immature stages). - Headrick and Goeden 1990b: 776 (life histo¬ ry). - Headrick and Goeden 1990c: 512 (resource utilization). Paracantha culta [misidentification]: Lange 1941: 56, 67 (damage, parasite, host, Calif.). Paracantha mexicana Malloch 1941: 41 (male holotype, NMNH; Guanajuato, Mexico). — Aczel 1952b: 233 (synonymy). Paracantha mimetica Malloch 1941: 41 (male holotype, NMNH; Alsea Mt., Oreg.). — Aczel 1952b: 233 (synonymy). Paracantha mimetica var. elongata Malloch 1941: 41 (male holotype, NMNH; Koebler, N. Mex.). — Aczel 1952b: 233 (synonymy). P. gentilis most closely resembles culta in head and body mark¬ ings, wing pattern, and in the possession of a long oviscape, which is about three times as long as the terminal abdominal tergite. This species is distinguished from culta by the smaller size of the parafacial spot (fig. 313, a); from cultaris by the longer oviscape and the location of the posterior dark ray in the apex of cell r4+5 (as in fig. 307, b); from genalis by the absence of the group of short, black, stout genal setae (as in fig. 313, b); and from forficula by the longer oviscape and the lack of a longitudinal division of the hyaline triangle in the apex of cell r4+5. recognition.

Map 46. P. gentilis also has been recorded from the states of Veracruz and Guanajuato in Mexico. distribution.

270

Systematic Treatment of the Genera

Pemberton et al. (1985) and Goeden and Ricker (1986b, 1986c; 1987a, 1987b) recorded gentilis from 10 species of Cirsium. This is probably the species, hosts.

reported as culta by Lange (1941), that causes minor damage to artichoke. discussion.

Headrick and Goeden (1990a) described the larva in detail, and

they (1990b) presented a detailed account of the life history of gentilis.

Genus Paraterellia Foote Trypeta (part): Aldrich 1905: 604 (in catalog). Acidia (part): Curran 1934: 287 (in key). Paraterellia Foote 1960a: 121 (type species, Trypeta varipennis Coquillett 1902b: 180, by original designation). — McFadden and Foote 1960: 254 (taxonomy). Foote and Blanc 1963: 44 (review, Calif.). - Foote 1965a: 671 (in catalog). Wasbauer 1972: 122 (hosts). — Steck 1981: 6 (hosts). recognition.

Medium-sized flies having a yellow to brown thorax, abdomen,

and legs; the abdomen patterned with dark markings; and prominently banded wings. These flies possess three pairs of frontal bristles, 2 pairs of orbitals, the posterior pair convergent; two pairs of well-developed scutellar bristles on a flat scutellum; one pair of dorsocentral bristles situated close to a transverse line through the acrostichal bristles; and slender, black, sharply pointed postocular bristles. In structure and color, Paraterellia resembles several other North American genera having essentially an orange or light brown body with somewhat darker bristles; slender, sharply pointed postoculars; and the dorsocentral bristles situated at or behind a transverse line through the supra-alar bristles. However, the convergent posterior orbital bristles invariably distinguish it from those genera. Paraterellia is separable from Chaetostomella, Orellia, and Neaspilota, the only three North Amer¬ ican genera of the tribe Terelliini (all of which have convergent orbital bristles), by the location of the dorsocentral bristles, by the wing pattern, and by the lack of a dark pattern and moderately stout white setulae on the scutum. Rbagoletis acuticornis and several species of Procecidocbaroides also have convergent orbital bristles. These species are easily distinguished from Paraterellia, the former by its mostly black body color and the extremely short apical extension of cell cup, and the latter by the presence of a pair of presutural dorsocentral bristles. Foote (1960a) presented a detailed discussion of this endemic west¬ ern North American genus and a revision of three of the four included species. He discussion.

also described the presence on the labellum of two species of slender capitate setae never seen in any other tephritid genus. Paraterellia is not commonly encountered and is of no known economic importance. It is not known to occur south of the United States. Although originally placed in the Terelliini because of its convergent posterior orbitals (Foote 1960a), Paraterellia belongs in the Trypetinae (Freidberg

271

Paraterellia 1985); it has three spermathecae, distinct scapular setae, and is biochemically more similar to other Trypetinae (Steck 1981). Its feeding habits also support this classifi¬ cation; all the Terelliini breed in species of Asteraceae (Freidberg 1985), whereas the two species of Paraterellia whose biology is known breed in the fruit of Juniperus. The shape of the surstyli in the male suggests its relationship with the subtribe Carpomyina.

Key to Known Species of Paraterellia 1. Brown or yellowish costal band as wide as cell rj distad of pterostigma (fig. 316, b); transverse dark band distad of pterostigma incomplete or inter¬ rupted in cell dm .2 Brown costal band narrower than cell (fig. 318, a); transverse dark band distad of pterostigma entire .3 2. Cell rj entirely yellowish and brown, no hyaline area immediately distad of pterostigma (fig. 316, a); subscutellum with pair of dark brown marks .varipennis (Coquillett) Cell r1 with a hyaline area, sometimes not reaching costa, immediately distad of pterostigma (fig. 317, a); subscutellum entirely orange or brown .immaculata Blanc 3. Band covering vein r-m separate from dark area on pterostigma, not Y-shaped, connected to band covering vein dm-cu along posterior wing margin (fig. 318, c) and to costal apical band anteriorly (fig. 318, a); band covering vein dm-cu not connected to apical costal band anteriorly; bristles orange; subscutellum entirely orange-brown .... superba Foote Band covering vein r-m Y-shaped, connected to dark area on pterostigma (fig. 319, a) and separate from band covering vein dm-cu and from apical costal band; the latter 2 bands connected anteriorly; bristles black; sub¬ scutellum with pair of dark brown marks .ypsilon Foote a

b

a

b

Figures 316-319. Right wings, Paraterellia spp. 316, varipennis (Coq.); 317, immaculata Blanc; 318, superba Foote; 319, ypsilon Foote.

Systematic Treatment of the Genera

Map 47. Distribution of Paraterellia immaculata, P. superba, P. varipennis, and P. ypsilon.

Paraterellia immaculata Blanc (Figs. 27, 317; Map 47) Paraterellia immaculata Blanc in Foote and Blanc 1979: 170 (male holotype, NMNH; 10 mi. w. Portal, Ariz.). The wing pattern of immaculata closely resembles that of varipen¬ nis in that a wide yellowish, brown-spotted band is present from the extreme base of recognition.

the wing to the apex of vein M bordering the costa (figs. 316, c; 317, c). This band is much wider in immaculata and varipennis than in the other two Paraterellia species. In immaculata, the band is broken immediately distad of the pterostigma (fig. 317, a), and the costal cells and the pterostigma are lighter in color than the remainder of the band, whereas in varipennis there is no such break (fig. 316, a), and the basal part of the band is darker. The inner margin of the dark area in the apex of cell r4+5 lies more perpendicular to vein M in immaculata than in varipennis (compare figs. 316, c and 317, c). It further differs from varipennis and ypsilon in lacking black spots on the sides of the subscutellum, and those on the sides of the scutum below the postalar bristle are smaller. Neither immaculata nor varipennis bear the capitate labellar bristles described for the genus. distribution.

Map 47.

Type specimens were swept from, and observed copulating on, Juniperus decipiens (Foote and Blanc [1979] and unpublished data). Additional specimens in HOSTS.

273

Paraterellia the National Museum of Natural History are labeled as reared from berries of Juniperus deppiana.

Paraterellia superba Foote (Fig. 318; Map 47) Paraterellia superba Foote 1960a: 123 (female holotype, NMNH; Calif.). — Foote and Blanc 1963: 44 (review, Calif.). - Foote 1965a: 671 (in catalog). - Foote and Blanc 1979: 170 (in key). P. superba differs from all other Paraterellia in that the bands covering veins r-m and dm-cu unite at the posterior wing margin (fig. 318, c), and the bristles are orange rather than black. Both superba and ypsilon differ from the other two Paraterellia species in having a comparatively narrow costal apical wing band (fig. 318, a) and two hyaline areas in cell r1? one immediately distad of the pterostigma and one in the apical third. Both superba and ypsilon bear capitate labellar recognition.

bristles. distribution.

host.

Map 47.

Not known.

Paraterellia varipennis (Coquillett) (Fig. 316; Map 47) Trypeta varipennis Coquillett 1902b: 180 (female holotype, NMNH; Ariz.). — Aldrich 1905: 605 (in catalog). - Foote 1960b: 254 (taxonomy). Trypeta versatilis Curran 1932b: 13 (female holotype, AMNH; Antelope Mt., Harney County, Oreg.). - Foote 1960a: 122 (synonymy). - Foote 1965a: 671 (in catalog). — Arnaud and Owen 1981: 152 (type data). Acidia versatilis: Curran 1934: 286, 287 (figure of wing, taxonomy). Paraterellia varipennis: Foote 1960a: 122 (revision). - Foote and Blanc 1963: 45 (review, Calif.). - Foote 1965a: 671 (in catalog). - Wasbauer 1972: 122 (hosts). Foote and Blanc 1979: 170 (in key). The wing patterns of varipennis and immaculata are similar in having a wide light brown band on the anterior margin (figs. 316, b; 317, b) extending from the base of the wing to the apex of vein M. The width of this band distinguishes both species from the other two Paraterellia species. In varipennis, this band is unbroken and is rather even in color along its entire length (fig. 316, b), but in immaculata a definite break of varying extent is present immediately distad of the pterostigma (Fig. 317, a) and there are more contrasting yellowish and dark brown areas. The inner margin of this band in the apex of cell r4+5 (fig. 316, c) in varipennis lies at an angle of less than 45° to vein M, but in immaculata this brown margin lies recognition.

274

Systematic Treatment of the Genera

more at a right angle to the long axis of the wing. In varipennis, a pair of black spots is present laterally on the subscutellum, the black spot on the side of the scutum below the postalar bristle is large, and there is sometimes a dark spot on the pleuron anterior to the haltere. Except for a small spot on the side of the scutum, these spots are absent in immaculata. Neither varipennis nor immaculata have capitate labellar bristles. distribution.

host.

Map 47.

P. varipennis has been reared from berries of Juniperus.

Paraterellia ypsilon Foote (Fig. 319; Map 47) Paraterellia superba [error]: Foote 1960a: 124 (name corrected in reprint). Paraterellia ypsilon Foote 1960a: 122 (in key); 124 (female holotype, NMNH; Calif.). - Foote and Blanc 1963: 45 (review, Calif.). - Foote 1965a: 671 (in catalog). - Foote and Blanc 1979: 170 (in key); 172 (taxonomy). P. ypsilon and superba are distinguished from the other two spe¬ cies of Paraterellia by their rather similar wing patterns, in which the apical costal recognition.

band is narrow, not filling the apical third of cell

P. ypsilon is the only Paraterellia

species with a distinctive Y-shaped mark associated with the pterostigma (fig. 319, a), and it is unique in the genus in having the band covering crossvein dm-cu and the apical costal band united anteriorly (fig. 319, b). P. ypsilon also differs from superba and immaculata in having a pair of blackish spots on the subscutellum. Both ypsilon and superba bear capitate labellar bristles. distribution.

HOST.

Not

Map 47.

known.

Genus Paroxyna Hendel Tephritis Latreille [misidentification]: Loew (part) 1873: 295, 328 (taxonomy). Aldrich (part) 1905: 611 (in catalog). Trypeta (Tephritis): Osten Sacken 1878: 193 (in catalog). Ensina Robineau-Desvoidy [misidentification]: Williston 1896: 122 (in key). Aldrich (part) 1905: 610 (in catalog). - Williston 1908: 287 (in key). - Hendel 1914b: 96 (in key). Tephritis (Ensina) [misidentification]: Cresson (part) 1907: 99 (taxonomy). Eunbia Hendel [misidentification]: V. T. Phillips (part) 1923: 123, 149 (in key; review, e. U.S.).

275

Paroxyna Paroxyna Hendel 1927: 146 (type species, Trypeta tessellata Loew 1844: 396, by original designation). — F. H. Benjamin (part) 1934: 40 (review, Fla.). — Curran 1934: 293 (in key). — Hering 1944a: 17 (taxonomy). — Foote 1959a: 16 (taxon¬ omy). — Foote 1960e: 72 (taxonomy). - Foote 1960f: 146 (taxonomy). — Foote and Blanc 1963: 46 (review, Calif.). — Foote 1965a: 665 (in catalog). - Wasbauer 1972: 122 (hosts). — Novak 1974b: 9 (revision). - Foote and Blanc 1979: 172 (syn¬ onymy, nomenclature). - White 1986: 150 (type species). Paroxyma [error]: Palmer and Bennett 1988: 222 (host). Flies belonging to Paroxyna range from quite small to medium¬ sized and in color from gray or black body and wings to amber body and light gray wings. The wings of all but one species have a characteristic speckled appearance, are void of definite stripes or bands, and the hyaline spots are relatively small and rounded. The two characters that are most quickly observed and are used as the primary criteria for recognizing Paroxyna are the geniculate proboscis with a rela¬ tively long labellum and the presence in most specimens of a round hyaline spot centered in the pterostigma; this spot is lacking in a few species and in the males of a few others. Other features that distinguish the adults of the Nearctic species are given in the key to genera. The two species of Dioxyna known to occur in America north of Mexico, very similar in many respects to those of Paroxyna, can be separated from Nearctic species of that genus by the head, which is distinctly longer than high, and recognition.

by the lack of predistiphallic setulae. We consider Dioxyna as a genus separate from Paroxyna, although in the past, some workers grouped them together under the latter name. The name Stylia is a synonym of Paroxyna, as shown by Foote and Blanc (1979). We use a broader concept for the genus than some recent workers (e.g., Korneyev 1989). In this sense, Paroxyna (s. lat.) incudes about 160 species worldwide, with 68 in the Palearctic, 28 in the Afrotropical, 23 in the Oriental, and 21 in the Neotropical Regions. Novak (1974b) has provided the only comprehensive revision of Paroxyna occurring in America north of Mexico. In that work, he described nine new species and provided wing photographs, genitalia drawings, distributional maps, and a key for the identification of 22 species, most of which are discussed herein. Knowledge of the taxonomy of immatures in Paroxyna is almost totally nonexistent. The larvae of all species that have been reared feed in flowers, mainly of species of Astereae, Senecioneae, and Lactuceae. Also lacking is an understanding of the importance of these flies in the balance of nature. Because they are all phytophagous as larvae on their composite hosts, and many have been found and reared from developing seed heads, their restraining effect on the competitiveness of certain host shrubs could possibly be significant. Species of Paroxyna tend to range farther into the northern latitudes than do most other tephritids and are represented in collections from Alaska and such Canadian provinces as Newfoundland, Northwest Territories, and Yukon. The distribution patterns of many species follow the major mountain ranges of western Canada and the United States. A few species range into coastal and discussion.

lowland desert habitats.

Systematic Treatment of the Genera

a

Figures 320-326. Right wings, Paroxyna spp. 320, opacipennis (Foote); 321, dupla (Cress.); 322, variabilis (Doane); 323, coloradensis Quis.; 324, snowi Her.; 325, footeorum Novak; 326, pallidipennis (Cress.).

Key to U.S. and Canadian Species of Paroxyna 1. Cells br, r2+3, r4+5, and dm almost completely dark except for several small, marginal hyaline spots (fig. 320) .opacipennis (Foote) Not with this combination of characters .2 2. More than half of cell c dark, usually with 2 small subhyaline spots, one at each end of the cell (fig. 321, a); at least basal halves of all femora with black or bluish ground color .3 Cell c with large subhyaline spots (fig. 327, a), the central dark spot no more than half the length of cell; leg color variable.7 3. Scutum with brownish stripes .4 Scutum without stripes .5

Ill

Paroxyna

4. Scutum largely brown-pollinose with faint brown stripes; hyaline spots of wing with indistinct borders; pterostigma completely dark in male (fig. 321, b) but with a subhyaline spot in female; all femora and generally hind tibia with grayish black ground color .dupla (Cresson) Scutum grayish blue-pollinose with distinct brown stripes; hyaline wing spots with distinct borders; apex of cell r4+5 usually with 2 hyaline spots (fig. 322, b) .variabilis (Doane) 5. Scutal setulae almost completely black; thorax and abdomen with black ground color, dark brown tomentose; wings very dark; pterostigma gen¬ erally completely dark in male (fig. 323, b) and with small subhyaline spot in female .coloradensis Quisenberry Scutal setulae almost completely white or yellow; other characters various .6 6. Scutum brown-tomentose with yellow setulae; wing spots generally with indistinct borders (fig. 324).snoxvi Hering Scutum faint brownish gray-tomentose with white setulae; wing spots with distinct borders (fig. 325).footeorum Novak 7. Legs yellow, parts of some of the femora with dark tomentum .8 At least basal half of all femora with black or bluish ground color . 15 8. Abdominal tergites with 2 longitudinal rows of black or brownish spots ..9 Abdominal tergites without spots, or at times with very faint spots .13 9. Wing almost subhyaline in male, darker in female; spots in cells r2+^ and r4+5 tending to be broadly coalesced (fig. 326, a). .pallidipennis (Cresson) Wing pattern not as above .10 10. Basal half of wing more than half subhyaline; many scattered subhyaline areas present (figs. 327, 328). 11 Basal half of wing less than half subhyaline; if greater, subhyaline areas present adjacent to broader dark areas.12 11. Apex of cell r4 + 5 with a subhyaline spot (fig. 327, b); scutellum yellow only at apex .pygmaea Novak Apex of cell r4+5 without a subhyaline spot (fig. 328, a); scutellum almost 2/3 yellow.occidentalis Novak 12. Apices of cells r2+3 and r4+5 dark (fig. 329, b, c); pterostigma almost entirely dark (fig. 329, a).distincta Quisenberry Apex of cell r2 + 3 with 2 rounded hyaline spots (fig. 330, b); apex of cell r4+5 with rounded marginal hyaline spot (fig. 330, c); pterostigma with a prominent round hyaline spot (fig. 330, a).albiceps (Loew) 13. Wing dark, with less than basal half subhyaline (fig. 331); abdominal tergites without spots.murina (Doane) Basal half or more of wing subhyaline (fig. 332); abdominal tergites some¬ times with faint spots .14 14. Wing extensively subhyaline, with darker area distally (fig. 332); body length less than 3.25 mm .steyskali Novak Wing less subhyaline, distal xh> more extensively dark (fig. 333); body length more than 3.5 mm .jamesi Novak

278

Systematic Treatment of the Genera

a

Figures 327-333. Right wings, Paroxyna spp. 327, pygmaea Novak; 328, occidentalis Novak; 329, distincta Quis.; 330, albiceps (Lw.); 331, murina (Doane); 332, steyskali Novak; 333, jamesi Novak.

15. Abdominal tergites without spots, or tergites 1-5 each with a pair of faint spots; wing mostly subhyaline, darkest at pterostigma (fig. 334, a) and distal portion of cell r2+3; scutum brownish tomentose. .. ..californica Novak Abdominal tergites 1-5 each with a pair of distinct dark spots .16 16. Scutum with narrow brown stripes on bluish white to yellowish back¬ ground; if brownish stripes wholly lacking, body length less than 2.5 mm .17 Scutum either without stripes or with broad brown stripes; total body length usually exceeding 2.5 mm.18 17. Bristles of fore femur thick and white (2 or 3 at distal end of ventral comb may be darker); wing as in fig. 335 .clathrata (Loew) All bristles of fore femur slender and amber brown; wing as in fig. 336 . . .farinata Novak

279

Paroxyna

a

b

340 Figures 334-340. Right wings, Paroxyna spp. 334, californica Novak; 335, clathrata (Lw.); 336, farinata Novak; 337, tenebrosa (Coq.); 338, dreisbachorum Novak; 339, genalis (Thoms.); 340, sabroskyi Novak.

18. Distal portions of cells rl5 r2+3, and r4+5 very dark and without hyaline areas (fig. 337, c) .tenebrosa (Coquillett) Distal portions of cells rl5 r2+3, and r4+5 each with hyaline areas (figs. 338340) . 19 19. Scutum with gray stripes on a brown background; hyaline spots in distal portion of cells r2+3 and r4+5 small, round, and separated (fig. 338, b) .dreisbachorum Novak Scutum with brown stripes on a gray background; some of the hyaline spots in the distal portion of cells r2+3 and r4+5 ovate and contiguous (fig. 339, c) .20 20. Scutellum all brown, or brown with amber apex; the 3 large hyaline spots distad of pterostigma in both sexes broad and close together (fig. 339, b);

Systematic Treatment of the Genera

hyaline spot in pterostigma well developed in both sexes (fig. 339, a) .. .genalis (Thomson) Scutellum grayish tomentose; the 1st of the 3 spots distad of the pterostigma of the male somewhat isolated from the 2nd and with its terminal point directed posterolaterally (fig. 340, b); hyaline spot in pterostigma usually lacking or diminutive in the male and subhyaline in the female (fig. 340, a).sabroskyi Novak

Paroxyna albiceps (Loew) (Figs. 98, 330; Map 48) Trypeta albiceps Loew 1873: 302 (male and female syntypes, MCZ; Canada, Me.). Tepbritis albiceps: Loew 1873: 304, 330 (taxonomy). — W. A. Snow 1894: 173 (distribution). — Doane 1899: 189 (distribution). — Johnson 1900b: 688 (N.J.). — McGillivray and Houghton 1903: 13 (N.Y.). - Hine 1904: 92 (B.C.). — Aldrich 1905: 611 (in catalog). — Johnson 1910: 803 (N.J.). — Winn and Beaulieu 1915: 154 (Que.). — Johnson 1925b: 264 (N. Engl.). — Johannson 1926: 159 (N.Y.). — Johnson 1927: 218 (Mt. Desert). — Leonard 1928: 853 (N.Y.). - Johnson 1930: 151 (Nantucket). - Petch and Maltaise 1932: 57 (Que.). - Petch 1935: 90 (Que.). — Strickland 1938: 204 (Alta.). Trypeta euryptera Loew 1873: 304, 305, 330 (female holotype, MCZ; West Point, N.Y.). - Aldrich 1905: 611 (in catalog). - V. T. Phillips 1923: 150 (synonymy). — Leonard 1928: 853 (N.J.). Trypeta (Tepbritis) albiceps: Osten Sacken 1878: 193 (in catalog). Trypeta (Tepbritis) euryptera: Osten Sacken 1878: 193 (in catalog). Euribia albiceps: V. T. Phillips 1923: 150 (review). — Brimley 1938: 385 (N. Car.). Paroxyna albiceps: Quisenberry 1951: 59 (taxonomy). - Foote 1965a: 665 (in catalog). - Novak et al. 1967: 147 (host). - Novak and Foote 1968: 108 (descrip¬ tion of immature stages, biology). - Wasbauer 1972: 122 (hosts). - Novak 1974b: 12 (revision).

recognition.

The wing pattern of albiceps is distinctive within much of its

geographical range. The eastern population displays large, very round hyaline spots in the center of the wing, each large spot having smaller angular spots wedged around it at 45, 135, 225, and 315°, giving the wing a lacelike appearance. The wing of specimens from the western part of its range often lacks the small angular wedgeshaped spots and resembles the pattern of murina. The presence of pairs of large black spots on the abdominal tergites in albiceps separates the western specimens from murina. P. albiceps is one of the eight Nearctic species having yellow legs. It can be separated from the other seven species by the two characters mentioned above and by the differences given in the key.

Map 48. According to Novak (1974b), albiceps is distributed mainly in the Canadian boreal forest but also inhabits temperate forests of the northeastern United States and Pacific coast. distribution.

281

Paroxyna

Map 48. Distribution of Paroxyna albiceps.

hosts.

Novak (1974b) listed six species of Aster as hosts.

discussion. P. albiceps

is one of the most widely distributed and easily recog¬

nized Paroxyna species.

Paroxyna californica Novak (Fig. 334; Map 49) Paroxyna californica Novak 1974b: 16 (female holotype, UCD; Calif.). - Goeden

1983: 399 (host).

282

Systematic Treatment of the Genera

Map 49. Distribution of Paroxyna californica and P. dupla.

a rather rare species, is distinguishable by its com¬ bination of black femora and the absence of paired spots on the abdominal tergites recognition. P. californica,

(or sometimes with very faint spots). The wing markings (fig. 334) are weak, espe¬ cially in the males, except for a markedly black pterostigma (fig. 334, a) and the extension of black posteriorly from the pterostigma across cell r^ The male usually lacks the hyaline spot in the pterostigma. distribution.

host.

Map 49.

Goeden (1983) reared adults from flower heads of Arnica sororia.

Paroxyna

Paroxyna clathrata (Loew) (Fig. 335; Map 50) Trypeta clathrata Loew 1862c: 80 (female syntypes, MCZ; “Middle States”)- —

Loew 1873: 297 (review). Tephritis clathrata: Loew 1873: 298, 330 (taxonomy). - Johnson 1904b: 162 (N.J.).

-Aldrich 1905: 611 (in catalog).-Johnson 1910: 803 (N.J.). - Aldrich 1913: 215 (Great Basin). — Cole and Lovett 1921: 326 (Oreg.). — Janes and Thomas 1932: 104 (Utah). - Knowlton and Harmston 1937: 145 (Utah). - Strickland 1938: 204 (Alta.).

Map 50. Distribution of Paroxyna clathrata.

Systematic Treatment of the Genera Trypeta (Tephritis) clathrata: Osten Sacken 1878: 193 (in catalog). Tephritis despecta [misidentification]: Cresson 1907: 105 (N. Mex.). Euribia clathrata: V. T. Phillips 1923: 151 (review). Paroxyna (Paroxyna) clathrata: Hendel 1935: 54 (taxonomy). Paroxyna clathrata: Quisenberry 1951: 59 (taxonomy). — Foote and Blanc 1963: 47 (review, Calif.). - Foote 1965a: 666 (in catalog). - Novak 1974b: 16 (revision). — Goeden 1983: 400 (hosts). recognition. P. clathrata, a common western species, is characterized by its

small size (less than 2.5 mm), narrow faint brown or tan longitudinal stripes on a

o Paroxyna coloradensis • Paroxyna distincta ▲ Paroxyna dreisbachorum

Map 51. Distribution of Paroxyna coloradensis, P. distincta, P. dreisbachorum, P. opacipennis, P. pallidipennis, and P. tenebrosa.

285

Paroxyna bluish white scutum, and black femora. The scutal stripes are lacking in some speci¬ mens. P. clathrata most closely resembles farinata but can be distinguished from that species by the color and thickness of the bristles on the fore femur; in clathrata these bristles are white and thickened, but in farinata they are amber to dark brown and thin. In clathrata there are sometimes two or three darker bristles at the distal end of the ventral comb of the fore femur, but the shorter bristles on the remainder of the femur are thick and white in contrast to those of farinata, in which they are thin and amber or brown. The hyaline spot in the pterostigma (fig. 335, a) is sometimes lacking, especially in the males. Map 50. P. clathrata is found at relatively high elevations through¬ out the greater part of its range. distribution.

Goeden (1983) reared clathrata from Haplopappus pinifolius, Senecio douglasii vars. douglasii and monoensis, and S. integerrimus var. exaltatus. hosts.

P. clathrata is commonly encountered throughout its range. Novak (1974b) included a large number of specimens of this species in his revisionary study. discussion.

Paroxyna coloradensis Quisenberry (Fig. 323; Map 51) Paroxyna coloradensis Quisenberry 1949c: 85 (male holotype, originally CSUFC, transferred to NMNH; Colo.). — Foote 1965a: 666 (in catalog). - Novak 1974b: 20 (revision). P. coloradensis is a dark, high-mountain species and is distin¬ guished by its black scutal setulae, by the more than half-darkened cell c (fig. 323, a) and by the lack of longitudinal scutal stripes. The wings are very dark. The pterostig¬ ma of the male is usually completely black (fig. 323, b), whereas that of the female displays a subhyaline spot. P. coloradensis is similar to dupla but can be distin¬ guished from that species by its lack of scutal stripes. recognition.

Map 51. P. coloradensis is confined to high Rocky Mountain elevations (up to 14,000 ft. in Colorado) and Mt. Rainier in the state of Washington. distribution.

host.

Not

known.

Paroxyna distincta Quisenberry (Fig. 329; Map 51) Paroxyna distincta Quisenberry 1949c: 85 (female holotype, originally CSUFC, transferred to NMNH; between Cerro Summit and Cimarron, Montrose County, Colo.). - Foote 1965a: 666 (in catalog). - Novak 1974b: 22 (revision).

Systematic Treatment of the Genera P. distincta is a yellow-legged Paroxyna species similar to albiceps in having abdominal tergal spots, which are sometimes faint, and a dark wing, the recognition.

basal half of which is less than half hyaline. P. distincta differs from albiceps in having a diminutive hyaline pterostigmal spot, totally lacking in some specimens (fig. 329, a), and in having the apex of cell r2+3 dark (fig. 329, b). P. albiceps has a large hyaline pterostigmal spot, and two prominent round hyaline spots bordering on the costa are present near the apex of cell r2+3. DISTRIBUTION.

host.

Not

Map 51.

known.

Paroxyna dreisbachorum Novak (Fig. 338; Map 51) Paroxyna dreisbachorum Novak 1974b: 23 (female holotype, UMSP; Colo.). recognition.

P. dreisbachorum is unique among Paroxyna in having a brown

scutum with longitudinal gray-tomentose stripes. Other distinguishing characters include grayish black femora, a totally brown scutellum, and a small, centrally located subhyaline pterostigmal spot (fig. 338, a) which is lacking in the male. The hyaline spots in the distal portions of cells r2+3 and r4+5 are small, round, and separated (fig. 338, b). DISTRIBUTION.

host.

Not

Map 51.

known.

P. dreisbachorum is a rather rare species that occurs in high moun¬ tain areas up to 12,500 ft. discussion.

Paroxyna dupla (Cresson) (Fig. 321; Map 49) Tephritis dupla Cresson 1907: 102 (male lectotype, ANSP; Beulah, N. Mex.). - Byers et al. 1962: 181 (type data). - Novak 1974b: 23 (type data). Paroxyna dupla: Foote 1962: 174 (lectotype designation). - Foote 1965a: 666 (in catalog). - Novak 1974b: 23 (revision). - Lavigne 1982: 49 (biology, host). P. dupla is a rather small, dark fly which is distinguished by bluish black femora and by the coloration of cell c, which is black over more than half of its surface (fig. 321, a). The scutum is largely brown tomentose with rather faint brown recognition.

stripes, a character separating dupla from the similar coloradensis, which lacks scutal stripes. The wing of dupla gives the appearance of being almost totally infuscated, with a prominent rectangular black area (fig. 321, b) covering the pterostigma

287

Paroxyna and extending posteriorly across cell r1 and becoming slightly lighter as it crosses cell r2+3- The pterostigmal hyaline spot is lacking in the male and is faint in the female. distribution.

Map 49. P. dupla occurs at relatively high elevations throughout

most of its range. host.

Lavigne (1982) reared dupla from Erigeron peregrinus.

Paroxyna farinata Novak (Fig. 336; Map 52) Paroxyna farinata Novak 1974b: 24 (male holotype, CAS; Ida.). - Arnaud 1979: 330 (type data). P. farinata is similar to clatbrata in its size, its bluish white scutum with faint, narrow light brown longitudinal stripes, and its blackened femora. As in clatbrata, the hyaline pterostigmal spot is usually present in females and usually absent in males (fig. 336). P. farinata can be separated from clatbrata by the color and thickness of the bristles on the fore femur, as discussed in the Recognition section recognition.

for clatbrata. distribution.

Map 52. P. farinata has been found farther north than almost any

other Paroxyna species included in this handbook. Although farinata has been found associated with many different plants, most of which were listed by Novak (1974b), none has been confirmed as a host. The host record of Elymus condensatus (Gramineae) reported by Novak (1947b) seems host.

doubtful.

Paroxyna footeorum Novak (Fig. 325; Map 53) Paroxyna footeorum Novak 1974b: 27 (male holotype, WSU; Mont.). - Arnaud 1979: 331 (type data). RECOGNITION.

In footeorum, a medium-sized, dark-winged fly, cell c is darkened

over half or more of its surface with a hyaline spot near each end. The wing color is dark brown. Hyaline spots are for the most part small, well separated throughout the wing, and with distinct borders. A row of small round, evenly spaced hyaline spots is present around the posterior periphery of the wing from cell r4+5 to the anal lobe (fig. 325, a). The brownish gray tomentose scutum is without stripes and bears short, whitish yellow setulae. The apical half of the scutellum is yellow. P. footeorum resembles snowi, but in the latter species, yellow setulae are present on the scutum, and the hyaline wing spots have indistinct borders.

Systematic Treatment of the Genera

Map 52. Distribution of Paroxyna farinata.

distribution.

Map 53. P. footeorum is primarily a high-altitude species.

Arnica cordifolia and A. latifolia were recorded by Novak (1974b). Our unpublished records also include A. rydbergi. hosts.

Paroxyna genalis (Thomson) (Fig. 339; Map 54) Trypeta genalis Thomson 1869: 585 (male lectotype, NRS; Calif.). - Loew 1873: 344 (taxonomy). - Cockerell 1898: 155 (N. Mex.). - Coquillett 1899c: 264 (synonymy). - Novak 1974b: 28 (lectotype designation).

289

Paroxyna

Map 53. Distribution of Paroxyna footeorum and P. jamesi.

Tephritis genalis: Loew 1873: 336 (taxonomy). - Coquillett 1899c: 264 (synonymy). - Cockerell 1902: 346 (N. Mex.). - Johnson 1903b: 106 (N. Mex.). - C. F. Baker 1904: 30 (Nev.). - Aldrich 1905: 612 (in catalog). - Tucker 1908: 303 (Colo.). Trypeta (Tephritis?) genalis: Osten Sacken 1878: 195 (in catalog). Tephritis corpulenta Cresson 1907: 103 (male holotype, AMNH; N. Mex.). Goeden and Blanc 1986: 88 (synonymy). Straussia genalis: Woodworth 1913: 137 (Calif.). Paroxyna difficilis americana Hering 1944a: 11 (male holotype, BMNH; So. Fork, Rio Grande R. [Ariz.?]). - Blanc and Foote 1987: 432 (synonymy). Paroxyna genalis: Hering 1944a: 11 (taxonomy). - Quisenberry 1951: 59 (taxon¬ omy). - Foote and Blanc 1963: 48 (review, Calif.). - Frick 1964: 709 (biology,

Systematic Treatment of the Genera

Map 54. Distribution of Paroxyna genalis.

hosts). - Foote 1965a: 666 (in catalog). - Wasbauer 1972: 123 (hosts). - Novak 1974b: 28 (revision). - Goeden 1983: 400 (host). - Goeden and Blanc 1986: 88 (hosts, synonymy). Paroxyna franciscana Hering 1947a: 5 (female holotype, BMNH; San Francisco, Calif.). — Foote 1965a: 665 (in catalog, synonymy). Paroxyna corpulenta: Quisenberry 1951: 59 (taxonomy). - Foote and Blanc 1963: 48 (review, Calif.). - Foote 1965a: 666 (in catalog). - Novak 1974b: 21 (revision). — Goeden and Blanc 1986: 88 (taxonomy). Paroxyna americana: Foote and Blanc 1963: 46 (review, Calif.). - Foote 1965a: 665 (in catalog). — Novak 1974b: 15 (revision). — Goeden and Benjamin 1985: 392 (fungal parasite).

291

Paroxyna P. genalis a variable species characterized by an all-brown scutellum, sometimes with an amber apex; the three large hyaline spots in cell r1 distad of the pterostigma (fig. 339, b) are broad and closely adjacent in both sexes; and the pterostigmal hyaline spot is present and well developed in both sexes. P. tenebrosa differs from genalis by the all-dark anterodistal portion of its wing and the distal portions of cells rl5 r2+3, and r4+5 (fig. 337). P. dreisbachorum differs from genalis by possessing gray stripes on a brown scutum, whereas in genalis the scutum recognition.

has brown stripes on a gray background. The closely similar sabroskyi differs in having a grayish tomentose scutellum. The male of sabroskyi usually lacks a pterostigmal spot, and the first of the three hyaline spots in cell rt distad of the stigma is somewhat isolated from the second, is wedge-shaped, and its sharp terminal point is directed posterolaterally. distribution.

Map 54. P. genalis possibly extends southward into Mexico.

Goeden and Blanc (1986) recorded genalis from five species of Haplopappus, five species of Senecio, and from plants belonging to the genera Eriophyllum, Layia, Madia, and Venegasia. Novak (1974b) also recorded a Microseris species (fly as corpulenta). hosts.

As a result of extensive rearing in California, Goeden and Blanc (1986) concluded that corpulenta, a fairly widely distributed species, is conspecific with genalis. discussion.

Paroxyna jamesi Novak (Fig. 333; Map 53) Paroxyna jamesi Novak 1974b: 29 (male holotype, WSU; Wash.). P. jamesi is one of the eight species of Paroxyna discussed in this handbook having yellow legs. Pairs of dark spots on the abdominal tergites are faint or lacking. The basal half of the wing is subhyaline (fig. 333), but the distal third is dark, differing from the wing of steyskali, which is dark only at its apex. P. steyskali is a smaller fly with a body length of usually under 3.25 mm, whereas that of jamesi is usually longer than 3.5 mm. The apical half of the scutellum is yellow in both of these species. Because of the variability in the abdominal tergal spots, jamesi can be confused with occidentalis. The latter species lacks a round hyaline spot at the tip of cell r4+5, whereas that spot is usually present in jamesi (fig. 333, a). The oviscape of jamesi is as long as the preceding five abdominal segments, but it is only as long as the preceding three in occidentalis. In general, jamesi inhabits lower elevations, whereas the closely similar steyskali is found at higher altitudes. recognition.

distribution.

host.

nana.

Map 53.

P. jamesi was recorded by Novak (1974b) only from Grindelia nana var.

292

Systematic Treatment of the Genera

•

Paroxyna murina

Map 55. Distribution of Paroxyna murina.

Paroxyna murina (Doane) (Fig. 331; Map 55) Tephritis murina Doane 1899: 189 (male lectotype, WSU; Whidby I., Wash.). — Coquillett 1900: 459 (Alaska). — Aldrich 1905: 612 (in catalog). — Cresson 1907: 102 (taxonomy). — Tucker 1908: 303 (Colo.). — Cole and Lovett 1921: 326 (Oreg.). — Strickland 1938: 204 (Alta.). — Foote 1966b: 123 (lectotype designa¬ tion). — Novak 1974b: 30 (type data). — Zack 1984: 32 (type data). Paroxyna maculifemorata Hering 1947a: 6 (female holotype, BMNH; Wash.). — Foote 1965a: 666 (in catalog). — Novak 1974a: 44 (synonymy). Paroxyna murina: Quisenberry 1951: 59 (taxonomy). - Foote and Blanc 1963: 49

293

Paroxyna (review, Calif.). - Foote 1965a: 666 (in catalog). - Foote 1966b: 123 (taxonomy). — Wasbauer 1972: 123 (host). — Novak 1974b: 30 (revision). Paroxyna maculifermorta [error]: Novak 1974b: 9 (synonymy). P. murina is fairly large compared with other members of the genus. It has yellow legs and a dark wing with less than its basal half hyaline. The light blue abdominal tergites lack spots and are densely covered by long, coarse, white setae. P. murina can be confused with western populations of albiceps, but the latter species has a pair of large, dark spots on each abdominal tergite. Both steyskali and jamesi resemble murina, but the wings of those two species have more than the basal half subhyaline, whereas the wing of murina (fig. 331) has less than the basal half subhyaline, giving it a blackish appearance. P. steyskali and jamesi often have faint, large, paired brown or gray spots on the abdominal tergites. recognition.

distribution.

host.

Map 55.

According to Novak (1974b), the only known host is Haplopappus

bloomeri.

Paroxyna occidentalis Novak (Fig. 328; Map 56) Paroxyna occidentalis Novak 1974b: 32 (male holotype, NMNH; Oreg.). - Arnaud 1979: 331 (type data). recognition.

Yellow legs, distinct pairs of black spots on the abdominal tergites,

and a wing with its basal half mostly subhyaline are the characteristics of this medium-sized Paroxyna species. P. pygmaea has these features but differs in having a hyaline spot near the distal tip of cell r4+5 and in having the scutellum yellow only at its apex, whereas occidentalis lacks the hyaline spot at the apex of cell r4+5 (fig. 328, a), and its scutellum is almost two-thirds yellow. P. occidentalis can be confused with jamesi or steyskali when the abdominal tergal spots of the latter two species are somewhat prominent but, as in pygmaea, both of those species usually have a round hyaline spot in the distal portion of cell r4+5. The oviscape of occidentalis is approxi¬ mately as long as the three preceding tergites, whereas that of steyskali and jamesi is as long as the preceding five. distribution. host.

Not

Map 56.

known.

Paroxyna opacipennis (Foote) (Fig. 320; Map 51) Tephritis opacipennis Foote 1960e: 74 (male holotype, NMNH; Cedar Pass, Modoc County, Calif.). — Foote and Blanc 1963: 69 (review, Calif.). — Foote 1965a: 668 (in catalog). Paroxyna opacipennis: Jenkins 1985: 76 (new combination).

294

Map 56. Distribution of Paroxyna occidentalis and P. pygmaea.

Paroxyna Because its wing is extensively blackened almost to its base (fig. 320), opacipennis is easily distinguished from all other Paroxyna species in America north of Mexico. The wing pattern somewhat resembles that of Trupanea conjuncta (fig. 446), but the latter has only one pair of scutellar bristles, the basal half of the wing is extensively hyaline, and the proboscis is not geniculate. The dark wing pattern of opacipennis also resembles that of Gonioxyna fuscata (fig. 244), but in that species, as well as in all species of Gonioxyna, the costa is strongly bowed and cell r1 is extremely broad. Additional characters that distinguish Paroxyna from Gonioxyna are given in the key to genera. The scutellum in most specimens of opacipennis bears a central yellowish spot seen best from above and behind. recognition.

Map 51.

DISTRIBUTION.

host.

Not

known.

Jenkins’ (1985) suggested transfer of opacipennis from Tephritis to Paroxyna was based mainly on the presence of a geniculate proboscis and the pres¬ ence in the male genitalia of predistiphallic setae. This nomenclatural combination discussion.

has not heretofore been used in the literature.

Paroxyna pallidipennis (Cresson) (Fig. 326; Map 51) Tephritis pallidipennis Cresson 1907: 104 (male lectotype, ANSP; Colo.). - Byers et al. 1962: 118 (type data). - Novak 1974b: 9, 33 (lectotype designation). Paroxyna pallidipennis: Quisenberry 1951: 59 (taxonomy). - Foote 1965a: 666 (in catalog). - Novak 1974b: 33 (revision). Paroxyna pallidipennis Foote [error for Cresson]: Harper 1962: 85 (Calif.). recognition.

This yellow-legged Paroxyna is readily recognized by its wing pat¬

tern (fig. 326). Except for its darker costal border, the entire wing is areolate, lightcolored, and with the hyaline spots either coalesced or very faintly outlined. The pterostigma is dark and has a subhyaline spot. The abdominal tergites each bear a pair of dark spots. distribution.

Map 51. Novak (1974b) indicated pallidipennis eventually may

be found in Mexico. host.

Not

known.

Paroxyna pygmaea Novak (Fig. 327; Map 56) Paroxyna pygmaea Novak 1974b: 33 (male holotype, CAS; Ida.). — Arnaud 1979: 331 (type data).

Systematic Treatment of the Genera

recognition.

P. pygmaea, a small fly, has yellow legs, prominent abdominal

tergal spots, and the basal half of the wing disk is more than half subhyaline. P. pygmaea differs from occidentalis by the presence of a round hyaline spot near the apex of cell r4+5 (fig. 327, b) and by the yellow color occupying only the apex of the scutellum. In occidentalis, the hyaline spot near the apex of cell r4+5 is lacking and the scutellum is about two-thirds yellow. Although the legs of pygmaea are predomi¬ nantly yellow, each of the femora often bears a large gray area dorsally.

distribution.

Map 56. P. pygmaea occurs mainly at high altitudes throughout

its range.

host.

Not

known.

discussion.

P. pygmaea is a commonly collected species in the western United

States.

Paroxyna sabroskyi Novak (Fig. 340; Map 57) Paroxyna sabroskyi Novak 1974b: 35 (male holotype, NMNH; Colo.). - Arnaud 1979: 331 (type data).

recognition.

P. sabroskyi has bluish black femora and large, prominent, dark

brown to black abdominal tergal spots. It differs from dreisbachorum in having brown longitudinal scutal stripes on a gray background; dreisbachorum has gray stripes on a brown scutum. In dreisbachorum, the hyaline spots in the distal portions of cells r2+3 and r4+5 are small, round, and well separated; those spots in sabroskyi (fig. 340, c) are large and sometimes close together or contiguous. P. sabroskyi resembles genalis, but in the former the scutellum is gray tomentose; the scutellum of genalis is totally dark brown or dark brown with an amber apex. In the male of sabroskyi, the first of the three hyaline spots in cell r1 distad of the pterostigma is somewhat isolated from the second, its sharp terminal point directed posterolaterally (fig. 340, b); and the pterostigmal hyaline spot is diminutive or lacking (fig. 340, a). In both sexes of genalis, the first three spots distad of the pterostigma are broad and close together, and the pterostigmal hyaline spot is well developed. Males of sabro¬ skyi have a prominent, wide black band covering the pterostigmal area and extending posteriorly, at least to vein M.

distribution.

Map 57.

Stephanomeria virgata was the only host recorded for sabroskyi (Goeden and Blanc 1986). host.

297

Map 57. Distribution of Paroxyna sabroskyi and P. snowi.

Systematic Treatment of the Genera Paroxyna snowi Hering (Fig. 324; Map 57) Tephritis obscuripennis Snow 1894: 174 (male syntype, UKL; Mt. Hood, Oreg.) (preoccupied by obscuripennis Loew 1850). — Coquillett 1899c: 264 (synonymy). — Aldrich 1905: 612 (in catalog). — Byers et al. 1962: 181 (type data). — Foote 1962: 176 (type data). — Foote 1965a: 666 (in catalog). — Novak 1974b: 36 (type data). Paroxyna snowi Hering

1944a:

8

(new

name

for obscuripennis

Snow). —

Quisenberry 1949c: 86 (taxonomy). — Foote and Blanc 1963: 49 (review, Calif.). — Novak 1974b: 36 (revision). Paroxyna obscuripennis: Quisenberry 1951: 59 (taxonomy, synonymy). recognition.

The size of snowi, a dark brown fly, is medium to large compared

with other species in the genus. Cell c is mostly dark with hyaline areas near each end (fig. 324, a). The legs are dark brown with a grayish black infuscation over the dorsal area of all femora. The wing is very dark brown but is black in the pterostigmal area. The hyaline spots of the wing, all with indistinct borders, are very small and round except for a few larger spots in the middle of cells rt and r2+3 and in cells dm and cuaj (fig. 324). The golden brown tomentose scutum is without stripes and bears yellow setulae. P. dupla and variabilis differ in having a striped scutum. The generally larger wing spots and the presence of black scutal setulae separate coloradensis from snowi. The faint brown-gray tomentose scutum with white setulae distinguish the very similar footeorum from snowi. distribution.

hosts.

Map 57.

Senecio triangularis is a host, according to Novak (1974b) and Novak and

Foote (1975). Our unpublished records show rearings from Arnica cor difolia. discussion.

This high-mountain species is not common in tephritid collections.

Paroxyna steyskali Novak (Fig. 332; Map 58) Paroxyna steyskali Novak 1974b: 36 (male holotype, NMNH; Oreg.). - Arnaud 1979: 331 (type data). recognition.

P. steyskali is a rather small fly with yellow legs. The paired tergal

spots of the abdomen are large and brown-tomentose but are obscure. More than three-fourths of the wing is subhyaline, being dark only near its apex (fig. 332); in this respect it differs from jamesi, the wing of which is darkened in its terminal third. P. jamesi is larger than steyskali, with its body usually longer than 3.5 mm; steyskali is usually shorter than 3.25 mm. The oviscape of steyskali is as long as the preceding five abdominal tergites; that of occidentalis is only as long as the three preceding ones.

299

Map 58. Distribution of Paroxyna steyskali and P. variabilis.

300

Systematic Treatment of the Genera

Map 58. This fly is commonly found at high altitudes, whereas jamesi is more prevalent in the lowlands. distribution.

host.

Not

known.

Paroxyna tenebrosa (Coquillett) (Fig. 337; Map 51) Tepbritis tenebrosa Coquillett 1899c: 264 (male holotype, NMNH; Custer County, Colo.). - Aldrich 1905: 612 (in catalog). - Novak 1974b: 78 (type data). Spatbulina tenebrosa: Cockerell 1917: 17 (Colo.). Paroxyna tenebrosa: Quisenberry 1951: 59 (taxonomy). — Foote 1965a: 666 (in catalog). — Novak 1974b: 38 (revision). recognition.

The identification of tenebrosa is simplified by the wing pattern

(fig. 337). Approximately one-third of the wing, bounded by vein Sc, the costa, and vein R4+5, is solid brown (fig. 337, c) except for two hyaline spots extending posteri¬ orly from the costa through cell r1 (fig. 337, b), the more basal one continuing across cell r2+3 and into cell r4+5. The dark brown area extends posteriorly across the apex of cell dm and covers vein dm-cu. Cell c is hyaline except for a small dark spot against the costa (fig. 337, a). Except for their apices, the femora are dark brown. Map 51. According to Novak (1974b), tenebrosa occurs mainly at high elevations throughout its range. distribution.

host.

Not

known.

Paroxyna variabilis (Doane) (Fig. 322; Map 58) Tepbritis variabilis Doane 1899: 188 (female lectotype, WSU; Corvallis, Oreg.). Aldrich 1905: 612 (in catalog). - Grossbeck 1912: 378 (type data). - Cole and Lovett 1921: 326 (Oreg.). - Foxlee 1957: 37 (B.C.). - Byers et al. 1962: 181 (type data). - Foote 1962: 178 (lectotype designation). - Foote 1966b: 125 (lectotype designation). - Novak 1974b: 39 (type data). - Zack 1984: 33 (type data). Euribia variabilis: Sweet 1930: 123, 131 (plant associations). Paroxyna variabilis: Quisenberry 1951: 59 (taxonomy). - Foote 1962: 178 (type designation). - Foote and Blanc 1963: 50 (review, Calif.). - Foote 1965a: 666 (in catalog). - Wasbauer 1972: 123 (hosts). - Novak 1974b: 39 (revision). recognition.

P. variabilis is a relatively large, dark-winged Paroxyna and is

distinguished by a combination of characters including: cell c dark brown except for a small hyaline spot near each end (fig. 322, a); legs brown, the femora further darkened by grayish black tomentum; scutum light blue-pollinose with a narrow central and two lateral prominent copper-colored longitudinal stripes, the latter pair

301

Peronyma usually extending posteriorly over the gray scutellum; the hyaline wing spots dis¬ tinctly bordered; and the apices of cells r2+3 and r4+5 each with at least one hyaline spot (fig. 322, b). P. dupla is similar to this species but has a brown scutum with faint brown stripes; the hyaline wing spots are indistinctly bordered; and the apex of cell r4+5 may have a small, indistinct spot which is lacking in cell r2+3. distribution.

Map 58. P. variabilis occurs at high elevations.

Novak (1974b) reported variabilis from Agoseris grandiflora and Microseris laciniata. hosts.

Genus Peronyma Loew Peronyma Loew 1873: 250 (type species, Trypeta sarcinata Loew 1862a: 218, by monotypy) (= quadrifasciata (Macquart)). - Williston 1896: 121 (in key). Coquillett 1899c: 267 (in key). — Aldrich 1905: 605 (in catalog). - Williston 1908: 286 (in key). — Coquillett 1910: 587 (type data). - Cresson 1914a: 276 (taxon¬ omy). — Hendel 1914b: 87 (in key). - Bezzi 1920: 6 (in key). - V. T. Phillips 1923: 121 (in key); 130 (revision). — Curran 1932b: 2 (in key). - F. H. Benjamin 1934: 24 (review, Fla.). — Curran 1934: 289, 293 (in key). — Aczel 1953: 125 (distribu¬ tion). — Foote 1965a: 660 (in catalog). — Wasbauer 1972: 123 (hosts). — Freidberg 1984: 129 (galls). Trypeta (Peronyma): Osten Sacken 1878: Tomoplagina Curran 1932b: 14 (type 1932b: 14, by original designation) (= jamin 1934: 24 (synonymy). — Arnaud

190 (in catalog). species, Tomoplagina maculata Curran quadrifasciata (Macquart)). — F. H. Ben¬ and Owen 1981: 151 (type data).

Peronyma is a genus of moderate-sized, robust flies with banded wings. It is unique in many respects among the North American Tephritidae and will not easily be confused with other genera. The scutellum is inflated and bilobate (fig. 37), with a shallow median depression visible from both posterior and dorsal as¬ pects; the lower outer corners of the face lack well-developed white setae; a dark spot is present on the parafacial between the eye and the face; the frons is about two times as wide as its length; the dorsocentral bristles are almost exactly in line with the supra-alar bristles; all scutal and frontal bristles arise from small dark swellings surrounding their bases; and a distinct apical elongation of cell cup is present. The wing pattern (fig. 341) also is unique in that the prominent brown bands are directed recognition.

Figure 341. Right wing, Peronyma quadrifasciata (Macq.).

Systematic Treatment of the Genera

diagonally from the wing base to the margins, and each band has a narrow, darker margin. In this respect, the pattern is reminiscent of Tomoplagia, which differs markedly in nearly every other way. discussion. This genus, although very distinctive, may be related to Eurosta. Many older pinned specimens of Peronyma have an oily appearance similar to that in the former genus, and the wide frons, swollen scutellum, and parafacial spot are found in several other genera that do not occur in the United States.

Peronyma quadrifasciata (Macquart) (Figs. 37, 341; Map 60) Tephritis quadrifasciata Macquart 1843a: 383 (male syntypes, UMO; Ga.). Macquart 1843b: 218 (taxonomy). - Osten Sacken 1878: 188 (in catalog). Aldrich 1905: 605 (in catalog). — F. H. Benjamin 1934: 24 (synonymy). Trypeta quadrifasciata: Osten Sacken 1858: 80 (in catalog). - Loew 1862c: 60 (taxonomy). - Loew 1873: 337 (synonymy). Trypeta sarcinata Loew 1862a: 218 (female syntype, NMW; S. Car.). - Loew 1864: 88 (taxonomy). - Loew 1873: 247 (review). - F. H. Benjamin 1934: 24 (syn¬ onymy). Peronyma sarcinata: Loew 1873: 250, 329 (taxonomy). - Aldrich 1905: 605 (in catalog). - Coquillett 1910: 587 (type data). - Hendel 1914b: 87 (type data). Sturtevant 1918: 35 (Ala., description). - V. T. Phillips 1923: 130 (review). — Brimley 1938: 383 (N. Car.). - Aczel 1949: 190 (in catalog). - Aczel 1953: 125 (taxonomy). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 123 (host). Trypeta (Peronyma) sarcinata: Loew 1873: 337 (synonymy). - Osten Sacken 1878: 190 (in catalog). Tomoplagina maculata Curran 1932b: 14 (female holotype, AMNH; Gotha, Fla.). Arnaud and Owen 1981: 151 (type data). - F. H. Benjamin 1934: 24 (synonymy). - Aczel 1953: 125 (synonymy). Peronyma maculata: F. H. Benjamin 1934: 24 (review, Fla.). - Curran 1934: 294 (figure of head). — Hennig 1952: 220 (in catalog immature stages). Tepbrella maculata: Bates 1935: 108 (review). recognition. As for the genus. distribution. Map 60. P. quadrifasciata is not known to occur south of the United States. host. P. quadrifasciata was reported from Heterotbeca tricbopbylla by Benjamin (1934). A. L. Norrbom examined a single male specimen from Georgia labeled “Tephritis 4-fasciata” by Macquart in the Bigot collection (UMO) at Oxford. discussion.

There is no doubt this represents the species which has always gone under the name

303

Procecidochares

Peronyma sarcinata Loew in the American literature. Macquart’s specimen is to be designated as the lectotype.

Genus Procecidochares Hendel Oedaspis Loew [misidentification]: Loew (part) 1873: 260, 328 (taxonomy, in key). — Williston 1896: 122 (in key). — Coquillett 1899c: 267 (in key). — Aldrich (part) 1905: 606 (in catalog). — Williston 1908: 284 (in catalog). - Cresson (part) 1914a: 277 (taxonomy). - Phillips 1923: 136 (synonymy). Trypeta (Oedaspis) [misidentification]: Osten Sacken 1878: 191 (in catalog). Oedaspis-solidago Patton 1897: 247 (type species, Trypeta atra Loew, by monotypy). Application for the suppression of this name is to be made to the Interna¬ tional Commission on Zoological Nomenclature. Procecidochares Hendel 1914b: 91 (type species, Trypeta atra Loew 1862a, by original designation). - Hendel 1914c: 42 (review). - Bezzi and Tavares 1916: 158, 161 (taxonomy, in key). — Bezzi 1920: 7 (in key). — V. T. Phillips (part) 1923: 122, 136 (in key; review, e. U.S.). - Hendel 1927: 87 (taxonomy). - Aldrich (part) 1929: 1 (revision). — Curran 1932b: 4 (in key). - F. H. Benjamin 1934: 22 (review, Fla.). - Curran 1934: 291 (in key). - Aczel 1953: 125 (key to species). - Foote 1960c: 671 (taxonomy, hosts). - Foote and Blanc 1963: 50 (review, Calif.). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 123 (hosts). - Furniss and Barr 1975: 20 (galls). — Wangberg 1978b: 148 (biology). — Silverman and Goeden 1980: 283 (biology, hosts, natural enemies). - Wangberg 1980: 401 (biology). - Freidberg 1984: 129 (gall biology). - Dodson 1986: 99 (behavior). - Maddison and Bartlett 1989: 32 (zoogeography). Prececidochares [error]: Aldrich 1929: 11 (in catalog). recognition.

Members of Procecidochares are small to medium-sized flies with

a yellowish head, black thorax, dark brown to black abdomen, and black or yellow femora. The wing is hyaline with four prominent, brown transverse bands (fig. 343), the basal one usually very short. The head bears two to five frontal bristles, some of them not strictly paired, and usually only one pair of orbitals; the posterior pair, when present, are extremely short and delicate. The scutellum is shining black, prominently swollen, and rounded dorsally. The scutum bears one or two pairs of dorsocentral bristles and displays two paired, shining black areas laterally before and behind the transverse suture. Along with much of the pleural region and the abdomi¬ nal dorsum, it bears numerous contrasting flattened white or yellowish setae in various patterns. Procecidochares is easily distinguishable from the somewhat similar genera Callachna, Peronyma, Cecidocharella, Stenopa, and Procecidocharoides by using the characters given in the key to genera. As far as known, the larvae of all species form galls on their host plants. As our studies have progressed, we have gained the impression that many, if discussion.

Systematic Treatment of the Genera not all, the species are restricted in their host preferences to a much greater extent than previously realized. Some of the taxonomic characters employed in the past to identify adults, such as the distribution patterns of flattened white setae on the scutum, are easily disrupted when specimens are preserved; we have tried to use alternative characters, especially in the key to species, that we hope circumvent some of these difficulties. Several investigators have shown that sibling or cryptic species exist in this genus, and we also have seen a number of undescribed species from the United States and Mexico. Some of these unresolved cases are treated in the ensuing discussions. Two Mexican species, not known to occur in the United States or Canada, have been introduced into the Australasian Region for the control of noxious weeds. P. utilis Stone has been introduced into Hawaii where it is effective in controlling the plant known there as Maui pamakani (Ageratina adenophorum), and it is now effectively controlling the same plant in parts of Australia, where it is called Crofton weed. P. alani Steyskal was more recently introduced from Mexico into Hawaii, where it is also established and successful in the control of hamakua pamakani [Ageratina rip aria) (Julien 1982).

Key to U.S. and Canadian Species of Procecidochares 1. Pterostigma lighter brown in basal fourth to half than in apical part; 2 pairs orbital bristles, the posterior pair quite short and delicate. .flavipes Aldrich Pterostigma evenly brown; 1 pair orbital bristles .2 2. Coxae and femora entirely yellow.3 Coxae and femora darkened to widely black, at least on basal halves of femora .4 3. Anepisternum (fig. 5) shining; basal band of wing darker brown than other bands (fig. 343, a) .polita (Loew) Anepisternum tomentose; basal band of wing concolorous with or lighter than the other bands (fig. 344, a).montana (Snow) 4. At least lower third of anepisternum tomentose.pleuralis Aldrich Anepisternum entirely shining .5 5. Basal and discal bands of wing at least partially joined (fig. 346, a) . .stonei Blanc and Foote Basal and discal bands entirely separated, although sometimes narrowly, by a hyaline space .6 6. Proximal hyaline triangle nearly equilateral (fig. 347, a), distinctly wider than distal triangle at posterior margin .atra (Loew) Proximal triangle much narrower, the two triangles about equally wide along posterior margin (fig. 348) .7 7. Only 1 pair of dorsocentral bristles, situated posterior to transverse suture ..8

Two pairs of dorsocentral bristles, one pair anterior to transverse suture .9 8. Parafacial less than half as wide as 1st flagellomere; wing as in fig. 348 . . .minuta (Snow)

Procecidocbares a

basal

discal

subapical

proximal triangle

apical

distal triangle

Figures 342-347. Right wings, Procecidocbares spp.; 342, flavipes Aldr.; 343, polita (Lw.); 344, montana (Snow); 345, pleuralis Aldr.; 346, stonei Blanc and Foote; 347, atra (Lw.).

Parafacial about half as wide as 1st flagellomere; wing as in fig. 349 .... .blantoni Hering 9. Polished area between posterior dorsocentral bristle and wing base divided by a single or double row of flattened whitish setae; wing as in fig. 350 .grindeliae Aldrich This polished area entirely without flattened whitish setae .10 10. Parafacial nearly half as wide as 1st flagellomere; wing as in fig. 351 .... .australis Aldrich Parafacial less than l/s as wide as 1st flagellomere; wing as in fig. 352 ... .anthracina (Doane)

Procecidochares anthracina (Doane) (Fig. 352; Map 59) Oedaspis antbracina Doane 1899: 180 (female lectotype, WSU; Ida.). — C. F. Baker 1904: 30 (Nev.). — Aldrich 1905: 606 (in catalog). — Foote 1966b: 121 (type designation). — Zack 1984: 32 (type data). Cecidocbares antbracina: Bezzi 1910: 25 (review). Procecidocbares antbracina: Bezzi and Tavares 1916: 162 (in key). - V. T. Phillips (part) 1923: 138 (taxonomy, synonymy). — Aldrich 1929: 5 (revision). — Knowlton

a

Figures 348-352. Right wings, Procecidochares spp. 348, minuta (Snow); 349, blantoni Her.; 350, grindeliae Aldr.; 351, australis Aldr.; 352, anthracina (Doane).

Map 59. Distribution of Procecidochares anthracina, P. minuta, and P. montana.

306

Procecidocbares

and Harmston 1937: 145 (Utah). — Foote 1960c: 672 (description). — Foote and Blanc 1963: 51 (review, Calif.). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 123 (host). A medium-sized species, anthracina possesses the following characters: frons dark yellow tomentose, 1.1 to 1.2 times as wide as long; parafacial distinctly less than half as wide as width of first flagellomere; gena about 0.15 times as high as eye; two pairs of dorsocentral bristles, one anterior and one posterior to the transverse suture; anepisternum entirely shining; oviscape about as long as last three abdominal tergites; all femora largely black except for yellow apices; wing pattern as in fig. 352, the basal band distinctly separated from discal band; apical and subapical bands not as widely separated, sometimes fused in a narrow area in vicinity of vein R2+3. This species closely resembles australis and grindeliae in having a shining anepisternum, two pairs of dorsocentral bristles, black coxae and femora, and similar wing patterns (figs. 350—352). The comparatively narrow parafacial separates it from australis, and the absence of white setae on the shining area be¬ tween the posterior dorsocentral bristle and the wing base distinguishes it from grindeliae. recognition.

distribution.

Map 59.

P. anthracina has been reared from a bud gall on Solidago californica Nuttall (Goeden in litt.) and from an unidentified species of Erigeron (Wasbauer 1972). It has also been swept from Chrysothamnus sp. (Foote and Blanc 1963). hosts.

During preliminary studies of this genus, we have seen a large num¬ ber of specimens we presume to be anthracina from widely scattered localities in the western United States. Whether they are conspecific with anthracina has yet to be discussion.

determined.

Procecidochares atra (Loew) (Figs. 39, 347; Map 60) Trypeta atra Loew 1862a: 219 (male and female syntypes, NMW; N.Y.). - Loew 1864: 89 (taxonomy). - Loew 1873: 256 (review). - Aldrich 1929: 8 (type data). Oedaspis atra: Loew 1873: 260, 329 (taxonomy). - W. A. Snow 1894: 164 (taxon¬ omy). - Doane 1899: 181 (taxonomy). - Wulp 1899: 408 (in catalog). - Johnson 1900b: 687 (N.J.). - F. FI. Snow 1903: 219 (Kans.). - Aldrich 1905: 606 (in catalog). — Washburn 1906: 82 (Minn.). — Johnson 1910: 801 (N.J.). — Felt 1918: 193, 198 (galls). - Sturtevant 1918: 36 (Ala.). - Britton 1920: 204 (Conn.). - Cole and Lovett 1921: 325 (Oreg.). - Huber 1927: 97 (parasite). - Felt 1940: 318,323 (galls). Trypeta (Oedaspis) atra: Osten Sacken 1878: 191 (in catalog). Oedaspis-solidago atra: Patton 1897: 247 (description adult, gall). Oedaspis setigera Coquillett 1899c: 262 (male and female syntypes, NMNH; Va.). -

308

Systematic Treatment of the Genera

Map 60. Distribution of Peronyma quadrifasciata, Procecidocbares atra, and P. stonei.

Aldrich 1905: 606 (in catalog). - Washburn 1906: 80 (Minn.). - Johnson 1913b: 83 (Fla.). - Bezzi and Tavares 1916: 161 (synonymy). - Aldrich 1929: 7 (syn¬ onymy, type data). Oediaspis [error] atra: Thompson 1907: 71 (biology). Cecidochares atra: Bezzi 1910: 22 (review). - Bezzi and Tavares 1916: 161 (in key). Cecidochares setigera: Bezzi 1910: 23 (review). Procecidocbares atra: Hendel 1914b: 91 (type data). - Hendel 1914c: 42 (review). Bezzi and Tavares 1916: 159 (taxonomy, in key). -V. T. Phillips (part) 1923: 138 (review). - Johnson 1925b: 262 (N. Engl.). - Leonard 1928: 853 (N.Y.). - Aldrich 1929: 7 (revision). - Curran 1930: 77 (N.Y.). - Brimley 1938: 383 (N. Car.). V. T. Phillips 1946: 58, 119 (description, biology larva, hosts). - Aczel 1949: 188 (in catalog). — Hennig 1952: 210 (in catalog immature stages). — Aczel 1953: 125

Procecidochares (in key). — Steyskal 1957: 94 (eye color). — Foote 1960c: 672 (description). — Foote 1965a: 660 (in catalog). - Wasbauer 1972: 123 (hosts). Procecidochares setigera: V. T. Phillips 1923: 138 (review). - Johnson 1925b: 262 (N. Engl.). recognition. One of the largest species discussed in this handbook, atra is

characterized by the following: frons densely tomentose, usually golden in color but sometimes darker, about 1.5 times as wide as high; parafacial at least 0.5 times as wide as width of first flagellomere; gena about 0.25 times as high as eye; usually two pairs of dorsocentral bristles, one in front of and one behind the transverse suture; anepisternum entirely shining; oviscape about as long as the last two abdominal tergites; all femora largely black except for yellow apices; basal and discal bands of wing (fig. 347) distinctly separated; subapical and apical bands usually fused or touching in cell r1 distad of pterostigma. P. atra is distinctive among all the black¬ legged species with a shining anepisternum in that the proximal hyaline triangle is broad and equilateral in shape (fig. 347, a), and the distal triangle is distinctly narrower at the posterior wing margin. The wing pattern of pleuralis is similar, but the two hyaline triangles are more nearly equal in size and shape, and the anepister¬ num is tomentose in the latter species. This wingpattern distinguishes atra regardless of the number of its dorsocentral bristles. distribution. Map 60. hosts. Wasbauer (1972) reported atra from several species of Solidago. His rec¬

ord from a species of Aster is questionable. discussion. P. atra is widespread and is the most commonly encountered Pro¬

cecidochares species occurring in the eastern half of the United States. We have seen specimens from the western part of its range having only one pair of dorsocentral bristles; additional morphological (and accompanying biological) differences may be found in such individuals, but whether these individuals constitute a different species has not been determined.

Procecidochares australis Aldrich (Fig. 351; Map 61) Procecidochares atra var. australis Aldrich 1929: 9 (female holotype, NMNH; Waco, Tex.). Procecidochares australis: F. H. Benjamin 1934: 22 (review, Fla.). — V. T. Phillips 1946: 59, 119 (description, biology larva, hosts). — Hennig 1952: 210 (in catalog immature stages). — Foote 1965a 660 (in catalog). — Stegmaier 1968b: 48 (biology, hosts). - Huettel and Bush 1972: 465 (host selection). - Wasbauer 1972: 123 (hosts). - Huettel and Bush 1973: 145 (host selection). Procecidochares sp. A of Huettel and Bush 1972: 467 (genetics, biology, host). — Huettel and Bush 1973: 145 (enzymes, biology, host).

310

Systematic Treatment of the Genera

Map 61. Distribution of Procecidochares australis, P. blantoni, and P. flavipes.

recognition.

F. australis is medium-sized with the following characters: frons

yellow-tomentose, at least 1.5 times as wide as high; parafacial about 0.5 times as wide as width of first flagellomere; gena about 0.25 times as high as eye; two pairs of dorsocentral bristles, one anterior and one posterior to the transverse suture; anepisternum entirely shining; oviscape almost as long as the last three abdominal tergites; femora either yellowish (but distinctly tinged with suffused gray areas, especially dorsally, as in Florida specimens), or distinctly black along at least twothirds of the length as in specimens from Texas; wing pattern as in fig. 351, the basal and discal bands narrowly separated; subapical and apical bands narrowly separated or touching in region of vein R2+3. Most Florida specimens of australis may be identified by the partial darkening of the femora, whereas the dark-legged Texas populations more closely resemble antbracina and grindeliae. P. australis differs from grindeliae in lacking the flat, whitened setae on the shining area between the wing base and the posterior dorsocentral bristle, and from antbracina principally by the much wider parafacial. DISTRIBUTION.

Map 61.

Benjamin (1934) studied Florida specimens reared from Heterotbeca subaxillaris and Conyza canadensis (as Erigeron pusillus), much less commonly on the hosts.

latter than on the former. Heterotbeca subaxillaris and H. latifolia are the only plants recorded as hosts of australis in Texas (Huettel and Bush 1972, 1973) (see also the ensuing discussion for additional host information).

Procecidochares Huettel and Bush (1972, 1973) have shown that part of the popula¬ tions of australis they studied in Texas can be differentiated genetically and by host preference, thus indicating the presence of a sibling species there. It prefers the camphor daisy, Mach aeranther a phyllocephala, which is closely related to Hetero¬ theca and is widespread in the southern United States and Mexico but apparently is not infested outside of Texas. No studies have been conducted for distinguishing this sibling morphologically from australis as defined here. discussion.

All the Florida specimens of australis described by Benjamin are much lighter in color and give the impression of being more fragile than the Texas specimens found by Huettel and Bush. The much lighter yellow legs of the former tempt one to group them in the key with polita and montana, which are entirely yellow-legged; the more robust and darker Texas individuals suggest they might belong to a different species. However, in this handbook, these two groups of australis are treated under one name because their host preferences and many of their morphological characters are so similar.

Procecidochares blantoni Hering (Fig. 349; Map 61) Procecidochares blantoni Hering 1940c: 12 (female holotype, BMNH; Salem, Oreg.). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 123 (hosts.). A small to medium-sized species, blantoni can be recognized in part by the following characters: frons yellowish tomentose, less than 1.5 times as wide as high; parafacial about 0.5 times as wide as width of first flagellomere; gena 0.1 to 0.15 times as high as eye; a single pair of dorsocentral bristles, situated posterior to the transverse suture; anepisternum entirely shining; oviscape about as long as the last three abdominal tergites; femora black except for yellow apices; wing pattern as in Fig. 349, the basal and discal bands narrowly separated from each other; subapical and apical bands widely separated to touching on vein R2+3. P. blantoni most closely resembles minuta in having black femora, a shining anepister¬ num, only one pair of dorsocentral bristles, and a similar wing pattern, but it differs from that species in its much wider parafacial and longer antenna and oviscape. recognition.

DISTRIBUTION.

Map 61.

Hering (1940c) described blantoni as having been reared (“gezuchtet”) from Heterotheca (as Coreopsis) oregana, but other host relationships of this species are uncertain. Heterotheca villosa and Heterotheca villosa var. sessiflora, originally listed as hosts for minuta, were stated by Wasbauer (1972) to be probable hosts of blantoni. Further study of the taxonomic and host relationships of blantoni and hosts.

minuta are required. Little is known about the morphological characters and biology of blantoni. If individuals of this species with two pairs of dorsocentral bristles are ever discussion.

312

Systematic Treatment of the Genera

identified, this species may be extremely difficult to distinguish from anthracina and its relatives.

Procecidochares flavipes Aldrich (Figs. 42, 342; Map 61) Procecidochares flavipes Aldrich 1929: 5 (female holotype, CAS; Revillagigedo Is., Mexico). - Foote and Blanc 1963: 51 (review, Calif.). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 124 (host). - Arnaud 1979: 331 (type data). One of the smaller species in the genus, flavipes possesses the following characters: frons yellow tomentose, about 1.2 times as wide as high; recognition.

parafacial distinctly narrower than half the width of first flagellomere; gena very narrow, less than 0.1 times as wide as eye height; two pairs of orbital bristles present, the posterior pair very short and slender; only one pair of dorsocentral bristles, situated behind the transverse suture; anepisternum densely tomentose; oviscape about as long as the last two abdominal tergites; all femora blackened, the apical one-fifth or one-sixth of each yellow; wing pattern as in fig. 342, the basal and discal bands narrowly separated; subapical and apical bands usually separated but some¬ times touching along vein R2+3* This is the only species of Procecidochares treated in this handbook having either two pairs of orbital bristles or the pterostigma “bi¬ colored,” with a light brown basal area contrasting with a darker apical one (% 342). DISTRIBUTION.

Map 61.

Wasbauer cited a Brickellia sp. as a host, and we have seen specimens reared from flower heads of at least two species in this plant genus by R. D. Goeden hosts.

(see ensuing discussion). G. L. Bush (unpublished data) indicates that a Brickellia species is the most likely host of the Mexican holotype of this species. discussion.

The only previous key to the species of this genus (Aldrich 1929)

gives the initial impression that three species of yellow-legged Procecidochares are present in the United States because the holotype of flavipes, a Mexican species with yellow femora, was included. However, in contrast to the holotype, all the specimens of flavipes from California we have seen possess primarily black femora and appear to be generally more robust. In the absence of detailed studies of these and possibly other differences, and because the distinguishing characters discussed above are com¬ mon to the holotype as well as to California material, we assume that only one species is involved.

Procecidochares grindeliae Aldrich (Fig. 350; Map 62) Procecidochares grindeliae Aldrich 1929: 6 (male holotype, NMNH; Alameda County, Calif.). - V. T. Phillips 1946: 119 (hosts). - Foote 1960c: 672 (descrip-

313

Procecidochares

Map 62. Distribution of Procecidochares grindeliae, P. pleuralis, and P. polita.

tion). — Foote and Blanc 1963: 52 (review, Calif.). — Foote 1965a: 660 (in catalog). — Wasbauer 1972: 124 (hosts). — Arnaud 1979: 331 (type data). Euribia grindeliae: Bates 1934a: 295 (taxonomy). A medium-sized species with the following characters: frons dark yellow to golden tomentose, 1.1 to 1.2 times as wide as high; parafacial less than 0.5 times the width of first flagellomere; gena 0.1 to 0.15 times as high as eye; two pairs of dorsocentral bristles, one behind and one in front of the transverse suture; anepisternum entirely shining; oviscape nearly as long as the four terminal abdomi¬ nal tergites; all femora mostly black except for yellow apices; wing pattern as in fig. 350, the basal and discal bands separated but sometimes approaching each other in cell br; subapical and apical bands usually widely separated. P. grindeliae, like antbracina and australis, has a shining anepisternum, black coxae and femora, two pairs of dorsocentral bristles; the wing patterns of these three species are similar (figs. 350-352). P. grindeliae is distinguished from those two species by the presence of a single or double row of flattened whitish setae dividing the shining area between the recognition.

posterior dorsocentral bristle and the wing base. distribution.

hosts.

Map 62.

P. grindeliae appears to be restricted to the plant genus Grindelia, the only

host record listed in the literature we have seen. Except for the distinguishing characters described above, we find it difficult to separate grindeliae from antbracina with confidence. Future biological discussion.

Systematic Treatment of the Genera investigations may show significant host and behavioral characteristics that will be additional means of distinguishing them.

Procecidochares minuta (Snow) (Fig. 348; Map 59) Oedaspis minuta Snow 1894: 164 (male holotype, UKL; Mont.). - Aldrich 1905: 606 (in catalog). — Byers et al. 1962: 180 (type data). Cecidocbares minuta: Bezzi 1910: 25 (review). — Ballou 1926: 29 (gall). Rhagoletis minuta: Aldrich 1913: 220 (biology). Procecidochares minuta: Bezzi and Tavares 1916: 162 (in key). — Aldrich 1929: 10 (revision). - Janes and Thomas 1932: 103 (Utah). - Knowlton and Harmston 1937: 145 (Utah). - Hering 1940c: 12 (taxonomy). — V. T. Phillips 1946: 119 (host). - Foote 1962: 176 (type data). - Foote and Blanc 1963: 52 (review, Calif.). - Foote 1965a: 660 (in catalog). - Novak et al. 1967: 147 (host, gall). - Harris and Piper 1970: 134 (hosts). - Wasbauer 1972: 124 (hosts). - Wangberg 1980: 401 (systematics, biology). Oedaspis atra Loew [misidentification]: Felt 1918: 198 (description gall). Procecidochares sp. A of Wangberg 1980: 402 (biology, hosts). Procecidochares sp. B of Wangberg 1980: 408 (biology, host). Procecidochares sp. C of Wangberg 1980: 413 (biology, hosts). recognition.

Individuals of minuta are small and may be recognized in part by

the following: frons yellow tomentose, 1.1 to 1.2 times as wide as high; parafacial distinctly less than half as wide as first flagellomere; gena about 0.2 times as high as eye; only one pair of dorsocentral bristles, situated close behind the transverse su¬ ture; anepisternum subshining but not at all tomentose; oviscape about as long as the two terminal abdominal tergites; all femora black except for yellow apices; wing pattern as in fig. 348, the basal and discal bands narrowly separated, at least in cell br (fig. 348, a); subapical and apical bands narrowly separated or joined anteriorly. Except for stonei and some specimens of atra, minuta and blantoni are the only two U.S. species of Procecidochares having black legs, a shining anepisternum, and only one pair of dorsocentral bristles. The two last-named species may be separated from the former two by characters given in the key and in the respective discussions. Individuals of minuta are considerably smaller than those of blantoni, the parafacial is much narrower, the antenna is shorter, and the orbital bristle is inserted in a narrow, more or less darkly shining band attaining the vertex of the head. distribution.

Map 59.

The genera Ambrosia, Aster, Brickellia, Chrysothamnus, Corethrogyne, Solidago, and Viguiera were cited by Wasbauer (1972), among which Chrysotham¬ hosts.

nus appears to be dominant. In fact, minuta has been identified from a much wider variety of hosts, and preliminary host observations by a number of investigators indicate that an undetermined number of sibling species are very probably involved. Dodson (1986, 1987a) published host records of a species “near minuta

315

Procecidochares Wangberg (1980) has shown that minuta as diagnosed in this hand¬ book comprises at least three sibling species that are apparently closely restricted in their host preferences, and further evidence from other sources, especially G. L. Bush (in litt.) and R. D. Goeden (in litt.), indicates that others may exist as well. G. C. Steyskal (unpublished data) indicates that at least some of these forms may be distin¬ guishable by external morphological characters, but his studies were preliminary and did not lead to satisfactory conclusions. Biological research on discrete populations of minuta, such as those conducted by Wangberg and now under way by Goeden in California, will very probably be required before morphological characters can be used to identify them accurately. discussion.

Procecidochares montana (Snow) (Fig. 344; Map 59) Oedaspis montana Snow 1894: 163 (male holotype, UKL; Mont.). — Byers et al. 1962: 180 (type data). Cecidochares montana: Bezzi 1910: 25 (review). Procecidochares montana: Bezzi and Tavares 1916: 162 (in key). - Aldrich 1929: 4 (revision). - Foote 1962: 176 (type data). — Foote 1965a: 660 (in catalog). A fairly small species, montana possesses the following charac¬ ters: frons brown to yellow tomentose, 1.1 to 1.2 times as wide as high; parafacial at least half as wide as width of first flagellomere; gena relatively wide, about 0.25 to 0.33 times as high as eye; only one pair of dorsocentral bristles, situated behind the transverse suture; anepisternum entirely tomentose, oviscape about as long as the two terminal abdominal tergites; legs entirely yellow; wing pattern as in fig. 344, the basal and discal bands tending to approach each other in cell br (fig. 344, a) and the subapical and apical bands touching or separate. Only montana, polita, and some specimens of flavipes have completely yellow coxae and femora; montana is distin¬ guishable from polita in having an entirely tomentose anepisternum, the four wing recognition.

bands concolorous, and the oviscape appreciably shorter. distribution.

host.

Not

Map 59.

known.

DISCUSSION.

No

information is available to us concerning the biology of

mon¬

tana.

Procecidochares pleuralis Aldrich (Fig. 345; Map 62) Procecidochares pleuralis Aldrich 1929: 9 (female holotype, NMNH; Fort Huachuca, Ariz.). - V. T. Phillips 1946: 119 (host). - Foote 1965a: 660 (in catalog). - Wasbauer 1972: 124 (host). - Arnaud 1979: 331 (type data).

Systematic Treatment of the Genera

P. pleuralis is the largest Procecidochares in the United States and Canada. It is characterized by the following: frons yellow to golden tomentose, 1.6 recognition.

to 1.7 times as wide as high, the lunule wide and very prominent; parafacial almost as wide as width of first flagellomere; the eye relatively narrow in relation to its width; gena about 0.25 times as high as eye; only one pair of dorsocentral bristles, situated behind the transverse suture; anepisternum tomentose; oviscape almost as long as the entire abdomen; all femora largely black except for yellow apices; wing pattern as in fig. 345, the basal and discal bands widely separated; subapical and apical bands sometimes touching along vein R2+3. In addition to the characters given above, pleuralis may be distinguished from all other Procecidochares discussed here¬ in by the wing pattern, in which both hyaline triangles are extensively developed and about the same width along the posterior wing margin (fig. 345, a). distribution.

Map 62. Our unpublished data indicates pleuralis may occur in

Mexico. The type specimens studied and described by Aldrich are labeled “from gall on sunflower stem.” This is the only host record known to us. host.

discussion.

No additional information concerning the biology of pleuralis is

available.

Procecidochares polita (Loew) (Fig. 343; Map 62) Trypeta polita Loew, 1862c: 77 (female syntypes, MCZ; D.C.). - Osten Sacken 1869: 299 (gall). - Loew 1873: 256 (review). - Beutenmuller 1892: 274 (gall). Jarvis 1907: 58 (gall). - Girault 1913: 340 (gall). - Aldrich 1929: 1 (type data). Oedaspis polita: Loew 1873: 260, 329 (taxonomy). - Osten Sacken 1880: 53 (host). - Keen 1885: 55 (Pa.). - Johnson 1900b: 687 (N.J.). - Aldrich 1905: 606 (in catalog). - Washburn 1905: 118 (Minn.). - Johnson 1910: 801 (N.J.). - Stebbins (part) 1910: 52 (gall). - Johnson 1913b: 83 (Fla.). - Felt 1918: 193 (gall). Britton 1920: 204 (Conn.). -Brimley 1938: 383 (N. Car.).-Felt 1940: 319 (gall). Trypeta (Oedaspis) polita: Osten Sacken 1878: 191 (in catalog). Oedaspis-solidago polita: Patton 1897: 247 (description adult, gall). Oediaspis [error] polita: Thompson 1907: 71 (biology). Cecidochares polita: Bezzi 1910: 24 (review). Procecidochares polita: Bezzi and Tavares 1916: 162 (in key). - V. T. Phillips (part) 1923: 137 (review). - Johnson 1925b: 262 (N. Engl.). - Leonard 1928: 852 (N.Y.). - Aldrich 1929: 3 (revision). - Johnson 1930: 151 (Nantucket). - V. T. Phillips 1946: 119 (host). - Foote 1965a: 661 (in catalog). - Wasbauer 1972: 124 (hosts). P. polita is a medium-sized to large species with the following characteristics: frons yellow and highly tomentose, 1.4 to 1.5 times as wide as high; recognition.

317

Frocecidocbares width of parafacial about half the width of first flagellomere; gena wide, about onefourth the eye height; only one pair of dorsocentral bristles, situated behind the transverse suture; anepisternum shining; oviscape about as long as the four terminal abdominal tergites; legs entirely yellow; wing pattern as in fig. 343, the basal and discal bands widely separated, the basal band darker in color than the other three (fig. 343, a); subapical and apical bands also widely separated. This species and montana are the only Frocecidocbares treated in this handbook having entirely yellow coxae and femora, but a number of species occurring in Mexico also share this character. The anepisternum of polita is distinctly shining, the basal wing band is darker than the other three bands, and the aculeus is long; in montana, the anepister¬ num is tomentose, all four wing bands are the same shade of brown, and the aculeus is shorter. distribution.

hosts.

Map 62.

Several species of Solidago were the only hosts listed by Wasbauer (1972).

Aldrich indicated that many early records of polita refer to atra. Other than the foregoing host data, no information is available to us concerning the biology of this species. discussion.

Procecidochares stonei Blanc and Foote (Fig. 346; Map 60) Frocecidocbares stonei Blanc and Foote 1961: 77 (female holotype, NMNH; San Ysidro, Calif.). — Foote and Blanc 1963: 53 (review, Calif.). — Foote 1965a: 661 (in catalog). — Tauber and Tauber 1968: 553 (biology). — Wasbauer 1972: 124 (hosts). — Silverman and Goeden 1980: 283 (hosts). P. stonei is a fairly large species having the following characters: head in lateral view relatively elongated, in some specimens almost two times as high as wide; frons dark yellow, tomentose, about 1.4 to 1.5 times as wide as high; parafacial almost as wide as width of first flagellomere; gena about 1.5 times as high as eye; one pair of dorsocentral bristles, situated behind the transverse suture; anepisternum subshining but distinctly not tomentose; oviscape almost as long as the four terminal abdominal tergites; femora black except for yellow apices; wing pat¬ tern as in fig. 346, the basal and discal bands usually fused posteriorly and the subapical and apical bands narrowly separated, sometimes touching. This species is distinguished from all other known Procecidochares by the fused basal and discal bands of the wing (fig. 346, a) and by the elongated nature of the head when viewed recognition.

laterally. distribution.

Map 60.

Part of the type material of stonei was reared from Viguiera laciniata. Wasbauer (1972) reported it from at least one species of Cbrysotbamnus and from hosts.

318

Systematic Treatment of the Genera

Corethrogyne filaginifolia and Heterotheca villosa, but according to R. D. Goeden (in litt.), these are questionable records.

discussion.

Our preliminary (unpublished data) studies show that at least one

undescribed species of Procecidochares somewhat similar to stonei exists in southern California. It very probably uses a different host (Goeden and Ricker 1976a, Silverman and Goeden 1980), otherwise it might well be confused with stonei. We believe that the Mexican alani, not yet found in the United States, also is closely related.

Genus Procecidocharoides Foote Trypeta (Oedaspis) (part): Osten Sacken 1878: 191 (in catalog). Oedaspis (part): Aldrich 1905: 606 (in catalog). Procecidochares (part): V. T. Phillips 1923: 122 (in key); 136 (review). - Aldrich 1929: 1 (revision). Procecidocharoides Foote 1960c: 671 (type species, Trypeta (Oedaspis) penelope Osten Sacken 1877: 346, by original designation). - Foote and Blanc 1963: 54 (review, Calif.). - Foote 1965a: 661 (in catalog).

Moderate-sized flies with three pairs of frontal bristles; two pairs of orbital bristles, the posterior pair reclinate or convergent; two pairs of scutellar bristles; two pairs of dorsocentral bristles, one anterior and one posterior to the recognition.

transverse suture; scutellum slightly swollen, shining or subshining (bare or lightly tomentose); and vein r-m situated well distad of the center of cell dm. The wing exhibits at least four prominent dark marks on a hyaline disk as follows: one subbasal band, one discal band traversing the wing at a slight angle, one apical band extending along the costa from the vicinity of the pterostigma to the apex of vein M, and one subapical band extending diagonally from the vicinity of the pterostigma to the apex of vein CuAj. In addition, a mark, which varies from a variable clouding (fig. 354, b) to a distinct dark band connected to the discal band in cell dm (fig. 353, b), is always present in cell cua! between the discal and subapical bands. This genus is distinguished from Procecidochares, which it strongly resembles, by the presence of well-developed posterior orbital bristles and a variable dark mark between the discal and subapical bands (figs. 353, b; 354, b). It also somewhat resembles Stenopa and Cecidocharella, but the pair of presutural dorsocentral bristles and the charac¬ ters presented in the key to genera distinguish it sufficiently. Except for penelope, Procecidocharoides is rarely encountered, and none of the species is of any known economic importance. Two of the species treated discussion.

here occur in Mexico, and we have seen other specimens, possibly representing undescribed species, from as far south as Panama.

319

Procecidocharoides

Figures 353-356. Right wings, Procecidocharoides spp. 353, caliginosus Foote; 354, flavissimus Foote; 355, penelope (O.S.); 356, pullatus Foote. Figures 357, 358. Thoracic pleuron, Pro¬ cecidocharoides spp. 357, flavissimus Foote; 358, penelope (O.S.).

Key to Known Species of Procecidocharoides 1. Apical wing band broadly connected to discal band, hyaline area between apical and subapical bands not extending anteriorly beyond vein R2+3 (fig. 353, a); band occupying center of cell cuaA fully connected to discal band in cell m (fig. 353, b); postocular setae mixed dark and light ... . .caliginosus Foote Apical band more narrowly connected to the discal band, hyaline area between apical and subapical bands extending anteriorly into cell r^ (fig. 355, a); band occupying center of cell cuaj never fully connected to discal band in cell dm (fig. 354, b), or indistinct; all postocular setae lightcolored .^ 2. Dark band occupying center of cell cua3 present (fig. 354, b), scutum with posterior and lateral margins broadly yellow (fig. 357, a); scutellum yel¬ low with black basal corners .. ..flavissimus Foote Dark band occupying center of cell cuaj indistinct or absent (figs. 355, b;

320

Systematic Treatment of the Genera

356, b); scutum and scutellum dark brown or black except for postpronotal lobes and usually notopleural area (fig. 358, a) .3 3. Apical and subapical bands never fully connected to discal band (fig. 355, a); posterior orbital bristles convergent (e. North America) . .pen el ope (Osten Sacken) Apical and subapical bands fully, although sometimes narrowly, connected to discal band (fig. 356, a); posterior orbital bristles reclinate (sw. U.S.) .pullatus Foote

Procecidocharoides caliginosus Foote (Fig. 353; Map 63) Procecidocharoides caliginosa Foote 1960c: 674 (female holotype, CAS; Magdalena Mts., Socorro County, N. Mex.). Procecidocharoides caliginosus: Foote 1965a: 661 (in catalog). - Arnaud 1979: 331 (type data). The wing pattern of caliginosus is distinctive in that the apical band is broadly connected to the discal band, as is the dark mark occupying the recognition.

center of cell cua^ in the other species, the latter mark is distinct only in flavissimus and never so broadly connected in cell dm (fig. 354, b). The postocular setae are mixed dark and light in caliginosus, whereas they are all light-colored in the other three species. The white setae on the scutum of caliginosus are arranged in four

Map 63. Distribution of Procecidocharoides caliginosus, P. flavissimus, P. penelope, and P. pullatus.

Procecidocharoides distinct longitudinal rows which are separated medially by the dorsocentral bristles. The basal two-thirds of the middle and hind femora are dark brown or black. distribution.

Map 63. We have also seen one specimen from the state of Mex¬

ico, Mexico. host.

Not

known.

Procecidocharoides flavissimus Foote (Figs. 354, 357; Map 63) Procecidocharoides flavissima Foote 1960c: 672 (female holotype, NMNH; Yreka, Siskiyou County, Calif.). — Foote and Blanc 1963: 54 (review, Calif.). Procecidocharoides flavissimus: Foote 1965a: 661 (in catalog). - Arnaud 1979: 332 (type data). P. flavissimus differs from the other three species of Pro¬ cecidocharoides occurring north of Mexico in the color of the scutum. In penelope, pullatus, and caliginosus, the scutum, except sometimes the notopleural area, and the scutellum are dark brown or black. In flavissimus, the lateral and posterior margins of the scutum are broadly yellow (fig. 357, a), and the scutellum is yellow except for the black lateral anterior corners. The wing bands of flavissimus are distinctly darker and better defined than in penelope (fig. 354), the hyaline spot between the discal and subapical bands is usually more pronounced (fig. 354, b), and the distance between veins r-m and dm-cu is greater than the length of r-m. recognition.

distribution.

host.

Not

Map 63.

known.

Procecidocharoides penelope (Osten Sacken) (Figs. 41, 355, 358; Map 63) Trypeta (Oedaspis) penelope Osten Sacken 1877: 346 (male and female syntypes, MCZ; Manlius, N.Y.). - Osten Sacken 1878: 191 (in catalog). Oedaspis penelope: Aldrich 1905: 606 (in catalog). Cecidochares penelope: Bezzi 1910: 25 (review). Procecidochares penelope: Bezzi and Tavares 1916: 162 (in key). — V. T. Phillips 1923: 137 (review). — Johnson 1925b: 262 (N. Engl.). — Leonard 1928: 852 (N.Y.). - Aldrich 1929: 3 (description). Procecidocharoides penelope: Foote 1960c: 672 (revision). — Foote 1965a: 661 (in catalog). - Stolzfus 1974: 12 (biology, description of larva). In contrast to flavissimus, the scutum and scutellum of penelope are broadly shining black; this color envelopes the postnotum or mediotergite, recognition.

322

Systematic Treatment of the Genera

spreads broadly around the posterior end of the thorax, forms a large dark spot immediately above the hind coxa, and covers most of the katepisternum (fig. 358, b). In penelope, the wing bands are wider, contain dark yellow areas, especially in the vicinity of the pterostigma, and only a suggestion of the dark spot between the discal and subapical bands is evident (fig. 355, b). In flavissimus, the scutum and thoracic pleura are much more broadly yellow (fig. 357), the scutellum is yellow centrally with black areas at its anterior basal corners, and the wing pattern differs as dis¬ cussed under that species. Of the four species treated here, penelope is unique in having distinctly convergent posterior orbital bristles. distribution.

HOST.

Map 63.

Eupatorium rugosum is the only known host. Stoltzfus (1984) reported

that the larva feeds in the flower without forming a gall and that it leaves to pupariate in the soil. In the older literature, penelope was included in the genus Procecidocbares. It is by far the most widespread and commonly encountered of the species of Procecidocbaroides. discussion.

Procecidocharoides pullatus Foote (Figs. 43, 356; Map 63) Procecidocbaroides pullata Foote 1960c: 673 (female holotype, NMNH; Chiricahua, Ariz.). Procecidocbaroides pullatus: Foote 1965a: 661 (in catalog). - Arnaud 1979: 332 (type data). P. pullatus and caliginosus are quite similar in possessing an en¬ tirely shining black abdomen and oviscape (variable in Mexican males), which imme¬ recognition.

diately distinguish them from penelope and flavissimus. In contrast to caliginosus, pullatus has entirely yellow legs, and the anterior half of the thoracic pleural area is yellowish white, two characters that readily distinguish this species. Map 63. P. pullatus also has been reported from the state of Puebla, Mexico (Foote 1980). distribution.

host.

Not

known.

Genus Rhagoletis Loew Zonosema Loew 1862b: 43 (type species, Tepbritis alternata Fallen 1814: 162, by subsequent designation of Rondani 1870a: 6). - Osten Sacken 1878: 190 (in catalog).

322

Systematic Treatment of the Genera

spreads broadly around the posterior end of the thorax, forms a large dark spot immediately above the hind coxa, and covers most of the katepisternum (fig. 358, b). In penelope, the wing bands are wider, contain dark yellow areas, especially in the vicinity of the pterostigma, and only a suggestion of the dark spot between the discal and subapical bands is evident (fig. 355, b). In flavissimus, the scutum and thoracic pleura are much more broadly yellow (fig. 357), the scutellum is yellow centrally with black areas at its anterior basal corners, and the wing pattern differs as dis¬ cussed under that species. Of the four species treated here, penelope is unique in having distinctly convergent posterior orbital bristles. distribution.

HOST.

Map 63.

Eupatorium rugosum is the only known host. Stoltzfus (1984) reported

that the larva feeds in the flower without forming a gall and that it leaves to pupariate in the soil. In the older literature, penelope was included in the genus Procecidocbares. It is by far the most widespread and commonly encountered of the species of Procecidocbaroides. discussion.

Procecidocharoides pullatus Foote (Figs. 43, 356; Map 63) Procecidocbaroides pullata Foote 1960c: 673 (female holotype, NMNH; Chiricahua, Ariz.). Procecidocbaroides pullatus: Foote 1965a: 661 (in catalog). - Arnaud 1979: 332 (type data). P. pullatus and caliginosus are quite similar in possessing an en¬ tirely shining black abdomen and oviscape (variable in Mexican males), which imme¬ recognition.

diately distinguish them from penelope and flavissimus. In contrast to caliginosus, pullatus has entirely yellow legs, and the anterior half of the thoracic pleural area is yellowish white, two characters that readily distinguish this species. Map 63. P. pullatus also has been reported from the state of Puebla, Mexico (Foote 1980). distribution.

host.

Not

known.

Genus Rhagoletis Loew Zonosema Loew 1862b: 43 (type species, Tepbritis alternata Fallen 1814: 162, by subsequent designation of Rondani 1870a: 6). - Osten Sacken 1878: 190 (in catalog).

323

Rhagoletis

Rhagoletis Loew 1862b: 44 (type species, Musca cerasi Linnaeus 1758: 600, by monotypy). — Loew 1873: 263, 328 (taxonomy, in key). — Williston 1896: 122 (in key). — Doane 1898: 69 (key to species). - Coquillett 1899c: 267 (in key). Aldrich (part) 1905: 606 (in catalog). - Williston 1908: 286 (in key). — Aldrich (part) 1909: 69 (revision). — Bezzi 1910: 14 (key to species). — Coquillett 1910: 599 (type data). — Cresson 1914a: 276 (taxonomy). - Hendel 1914b: 91 (in key). — V. T. Phillips 1923: 122, 134 (in key; review, e. U.S.). - Aldrich 1929: 2 (taxonomy). — Cresson 1929: 401 (revision). — Curran 1932a: 3 (key to species, taxonomy). - Curran 1932b: 3 (in key). - F. H. Benjamin 1934: 12 (review, Fla.). — Curran 1934: 289 (in key). — V. T. Phillips 1946: 59 (immature stages). — Rohdendorf 1961: 89 (review, Palearctic species). — Shewed 1961: 1045 (taxonomy). - Foote and Blanc 1963: 54 (review, Calif.). - Foote 1965a: 674 (in catalog). — Bush 1966a: 431 (revision, Nearctic species). — Foote 1967b: 40 (in catalog, Neotropical Region). - Bush 1969a: 237 (speciation). - Bush 1969b: 669 (ecology). — Wasbauer 1972: 125 (hosts). — Bush 1974: 3 (speciation). — Boiler and Prokopy 1976: 223 (biology). - Bush and Kitto 1977: 94 (speciation). - Hardy 1977: 114 (in catalog, Oriental Region). — Kandybina 1977: 124 (immature stages). — Prokopy 1977b: 305 (trophic relations). — Berlocher 1980a: 131 (elec¬ trophoretic key to species). — Prokopy 1980: 37 (mating behavior). — Berlocher 1981: 1 (phylogeny). — Foote 1981: 1 (revision, Neotropical species). — Berlocher and Bush 1982: 136 (phylogeny). — Diehl and Bush 1983: 276 (host trace ele¬ ments). — Rossiter et al. 1983: 77 (symbiotic bacteria). — Foote 1984: 120 (in catalog, Palearctic Region). — E. J. Harris 1989: 75 (pest status). - D. J. Howard 1989: 121 (symbionts). Acidia Robineau-Desvoidy [misidentification]: Loew 1873: 235, 327 (taxonomy, in

key). — Aldrich (part) 1905: 603 (in catalog). Spilographa Loew [misidentification]: Loew (part) 1873: 245, 327 (in key, taxon¬ omy). - Aldrich (part) 1905: 604 (in catalog). - Williston 1908: 286 (in key). Trypeta (Acidia) [misidentification]: Osten Sacken (part) 1878: 189 (in catalog). Trypeta (Rhagoletis): Osten Sacken 1878: 191 (in catalog). Trypeta (Spilographa) [misidentification]: Osten Sacken (part) 1878: 191 (in catalog). Zonosema: Coquillett (part) 1899c: 267 (in key). Trypeta [misidentification]: Aldrich (part) 1905: 604 (in catalog). Tephritis [misidentification]: Aldrich (part) 1905: 611 (in catalog). recognition. Flies of the genus Rhagoletis are medium to small in size, with an either black or yellowish brown thorax and abdomen, usually yellow legs, and prominently banded wings. One of the most distinctive features is the apically pointed first flagellomere (as in fig. 34, a), a character it shares in North America only with the genera Acidogona, Xenochaeta, and Zonosemata. Rhagoletis may be distinguished from the former two genera by its unicolorous rather than mixed black and white bristles and setulae and from the latter by the position of the dorsocentral bristles, which always arise anterior to the midpoint between the acrostichal and supra-alar bristles (fig. 23), and by the additional characters given in the key to

324

Systematic Treatment of the Genera

genera. Vein r-m in Rhagoletis is situated approximately at the midpoint of cell dm (fig. 57, a); two pairs of scutellar, three pairs of frontal, and two pairs of orbital bristles are present; the posterior orbitals are usually reclinate (convergent in acuticornis); the scutellum is not swollen and in the black-bodied species always has a prominent creamy white or yellow spot. Extensive morphological studies have been carried out on the adult stage (see Bush 1966a and other publications referenced in the synonymies of the individual species), and some morphological work has been done on immature stages of the Nearctic species. Therefore, except for Steyskal’s (1973) key to the larvae of Juglans-infesting species and possibly the electrophoretic key to species by Berlocher (1980a), few means are available for the precise identification of species on the basis of immature stages. If proper host information is available, one might assume the identity of the larvae of some oligotrophic species such as basiola (restricted to Rosa spp.), juniperina (restricted to Juniperus spp.), and so on. On the other hand, such assump¬ tions are always dangerous whenever precise identifications are sought, as demon¬ strated by Berlocher’s (1984a) discovery of an electromorph on Cornus quite similar to, if not inseparable morphologically from, tabellaria, previously believed to be a distinctive Corwws-inhabiting species. The adults of many Rhagoletis species are comparatively distinctive and relatively easy to identify by external characters. discussion.

Rhagoletis is by far the largest genus of the subtribe Carpomyina. It

may be paraphyletic as currently composed; some species groups may be more closely related to Carpomya Costa, Zonosemata, or other genera. Berlocher and Bush (1982) analyzed the relationships of many of the species groups on the basis of isozyme data. In the New World, Rhagoletis occurs not only in North America but in Central America, the West Indies, and much of the South American continent, where a total of 15 species has been recorded. Twenty-four species or subspecies of the genus are known from Europe, Asia, and North Africa, and a few have been recorded in the Oriental Region. This group of flies is comparatively well studied except in remote areas, so that not many undescribed species, at least on the basis of morphological characters, remain to be discovered within the United States and Canada. The economic impact of Rhagoletis is paramount in the United States and Canada, primarily because of the widespread infestation of apples and cherries by several of the species discussed herein. Because of its wide distribution, it is probably more economically important in our region than Anastrepha, a genus of acknowledged importance in the subtropical regions of the United States where citrus is a significant part of the area’s agricultural economy. The importance of R. pomonella is attested to by the immense body of research partially listed in our synonymy for that species; more phases of taxonomy, biology, and control are known for this species than for almost any other fruit fly in the world except perhaps for the Mediterranean fruit fly, Ceratitis capitata. The user of this handbook is referred to the publication of Bush (1966a), which is one of the most detailed taxonomic revisions ever to appear for any North American

Rhagoletis fruit fly genus. Abundant data on origins, hosts, biology, cytology, and many other aspects of North American species of Rhagoletis not within the scope of this hand¬ book are readily available in that publication and will be most helpful to those who require more detailed information. A clear understanding of Rhagoletis wing band terminology will be most helpful to users of the keys and subsequent descriptions of the species discussed herein. This terminology is detailed in fig. 359 and is defined as follows: (a) The subbasal band originates at the humeral vein and usually terminates in cell

cup. (b) The discal band originates on the pterostigma, crosses vein r-m, and runs to or in the direction of the apex of vein A] + CuA2 (c) The subapical band, when present, originates at a point on the costa well distad of the pterostigma, crosses vein dm-cu, and runs to the posterior wing margin just basad of the apex of vein CuAg an important exception is found in the pomonella group, in which this band is absent (figs. 374-377). (d) The anterior apical band, usually joined to the subapical band anteriorly, borders the costa or runs close to it and terminates in most species at or closely beyond the apex of vein M. (e) In some species, a posterior apical band also is present (fig. 367); it originates either on the subapical band or, if the subapical band is not present, in the vicinity of vein r-m on the discal band and terminates either on or between the apices of vein CuAj and M. (f) The intercalary band (fig. 361), which arises at the costa between the discal and subapical bands, is short and present in only two North American species of Rhagoletis. The following key, based primarily on wing pattern, facilitates the preliminary sorting of unidentified specimens into one of the major species groups. The data in this key also summarize the species groups of North American Rhagoletis and their hosts, as developed and presented by Bush (1966a). Note that the groupings ob¬ tained by wing pattern alone do not exactly correspond to those based on a combina¬ tion of characters and that two of the North American species (acuticornis and fausta) do not belong to any of the formally designated species groups as defined by Bush (1966a).

Key to Groups of Species of Rhagoletis Based on Wing Pattern 1. An intercalary band present. R. alternata group (figs. 361, 362) R. basiola Rosaceae R. meigenii Berberidaceae No such band present .

326

Systematic Treatment of the Genera 2. Subapical band completely lacking, the two apical bands joining the discal band and forming an F-shaped pattern in the apical half of the wing R. pomonella group (figs. 374—377) R. pomonella Rosaceae R. zephyria Caprifoliaceae R. mendax Ericaceae R. cornivora Cornaceae Subbasal, discal, and subapical bands all present.3 3. Apical band not joined to subapical band in cell r1? represented by only 2 discrete dark spots at apices of R2+3 and R4+5. R. acuticornis (fig. 360) Solanaceae Apical band joined to subapical band in cell r1? usually unbroken.4 4. Apical band divided into 2 arms at its junction with vein R2^3 (outermost arm sometimes broken). R. cingulata group (figs. 370—373) R. cingulata Rosaceae R. indifferens Rosaceae R. osmanthi Oleaceae R. chionanthi Oleaceae Apical band entire, undivided .5 5. Anterior and posterior apical bands present, both joined to the subapical band... R. fausta (fig. 367) Rosaceae R. striatella group (fig. 369) R. striatella Solanaceae Only an anterior apical band present, sometimes represented by 2 discrete dark spots .6 6. Apical band touching costa along its entire length . R. suavis group (figs. 363-366, 368) R. suavis Juglandaceae R. completa Juglandaceae R. juglandis Juglandaceae R. boycei Juglandaceae R. zoqui Juglandaceae R. ribicola group (part) (fig. 380) R. berberis Berberidaceae Apical band separated from costa by a hyaline area in at least cells r2 + 3 and r4 + 5

.

R. ribicola group (part) (fig. 382) R. ribicola Saxifragaceae R. tabellaria group (figs. 359, 378, 379, 381, 383) R. tabellaria Cornaceae, Ericaceae R. electromorpha Cornaceae R. juniperina Cupressaceae R. persimilis Not known R. ebbetsi Not known Because wing pattern alone is not sufficient for precise species identification of North American Rhagoletis, the following key must be used for that purpose.

subbasal

discal

subapical anterior apical

b

apical

Figures 359-368. Right wings, Rhagoletis spp. 359, tabellaria (Fitch), showing named crossbands; 360, acuticornis (Steysk.); 361, basiola (O.S.); 362, meigenii (Lw.); 363, juglandis Cress.; 364, completa Cress.; 365, boycei Cress.; 366, zoqui Bush; 367, fausta (O.S.); 368, suavis (Lw.).

327

Systematic Treatment of the Genera

Key to U.S. and Canadian Species of Rhagoletis (Adapted from Bush 1966a) 1. Posterior orbital bristles convergent (as in fig. 27, a); apical wing band incomplete, represented by only 2 discrete dark spots (fig. 360, a) .. .. .acuticornis (Steyskal) Posterior orbital bristles reclinate; apical band entire (although sometimes divided), connected to subapical band (fig. 359) .2 2. Scutellum concolorous cream to yellowish white, without distinct spot 3 Scutellum basically dark with distinct cream to yellow, circular, trapezoidal, or V-shaped spot (figs. 384, a; 385, a).10 3. Wing pattern with a small intercalary band crossing cells r1 and r7+3 be¬ tween discal and subapical crossbands (fig. 361); body brown to yellow (R. alternata species group) .4 Wing pattern without such a band (fig. 363, b); body yellow or black (R. suavis species group) .5 4. Subapical wing band complete from costa to vein CuAj (fig. 361, a); apical band narrow, occupying no more than apical Ve of cell r4+5 (fig. 361, b); infesting hips of Rosa spp.basiola (Osten Sacken) Subapical band extending no farther forward than vein R2 + 3 (fig. 362, a); apical band wide, occupying V2 to 2/s of cell r4+5 (fig. 362, b); infesting fruits of Berberis vulgaris ...meigenii (Loew) 5. Wing pattern with indistinct subbasal crossband (fig. 363, a); body entirely yellow; infesting Juglans spp.juglandis Cresson Wing pattern with dark, distinct subbasal crossband (fig. 364, a); body yellow to black .6 6. Discal band not normally joined to subapical band (fig. 364, b); if joined (fig. 367, a), then body yellow to yellowish tan and subscutellum entirely brown or with dark brown stripes (fig. 386).7 Discal band broadly joined to subapical band (figs. 367, a; 368, a); sub¬ scutellum entirely black or yellow, never brown or with vertical stripes .9 7. Veins R2+3 and R4+5 with horizontal fuscous markings between subapical and apical crossbands (figs. 365, b; 366, a) .8 Veins R2+3 and R4+5 without fuscous markings in this area; wing pattern as in fig. 364; infesting Juglans spp.completa Cresson 8. Thorax and abdomen black; wing pattern as in fig. 365; infesting Juglans SPP.boycei Cresson Thorax and abdomen predominantly tan to yellow; black markings limited to anepisternal and katepisternal areas (fig. 389, b); wing pattern as in fig. 366; infesting Juglans spp.; Mexico only.zoqui Bush 9. Body black; abdomen without white bands; wing pattern as in fig. 367; infesting Prunus spp.fausta (Osten Sacken) Body yellow to yellowish tan; wing pattern as in fig. 368; infesting Juglans SPP.suavis (Loew) 10. Scutellar spot V-shaped (fig. 384, a); subapical and posterior apical band present; wing pattern as in fig. 369; infesting husk tomato (Physalis spp.) .striatella Wulp

Figures 369-378. Right wings, Rhagoletis spp. 369, striatella Wulp; 370, indifferens Curr.; 371,

cingulata (Lw.).; 372, chionanthi Bush; 373, osmanthi Bush; 374, zephyria Snow; 375, pomonella (Walsh) (U.S.); 376, mendax Curr.; 377, cornivora Bush; 378, persimilis Bush.

329

330

Systematic Treatment of the Genera

Scutellar spot oval or trapezoid in shape (fig. 385, a); either the subapical or posterior apical band absent (although anterior apical band is forked in cingulata group) .11 11. Apical band of wing forked (fig. 370, a), or upper arm of fork separated from apical band by a hyaline area (fig. 371, a) {R. cingulata species

group).12 Apical band of wing entire (fig. 374, c) .15 12. Black shading present on posterior surface of fore coxa; epandrium (fig. 388) black; anterior arm of apical band nearly always entire (fig. 370, a), rarely forming an isolated spot at apex of vein R3 + 4 as in fig. 371, b; occurring west of 110°W.indifferens Curran Fore coxa almost entirely yellow; epandrium yellow; occurring east of 110°W . 11 13. Anterior arm of apical band nearly always forming an isolated spot at apex of vein R4+5 (fig. 371, b), rarely connected to subapical band; infesting fruits of black cherry; eastern U.S., Canada to Florida. .cingulata (Loew) Anterior arm of apical band nearly always connected to subapical band (figs. 372, a; 373, a); infesting fruits of Oleaceae.14 14. Aculeus 0.9-1.0 mm long; infesting Chionanthus spp.; adults active midJuly to early August; southeastern U.S.; wing as in fig. 372 . . chionanthi Bush Aculeus longer than 1.0 mm; infesting fruits of Osmantbus spp.; adults active October to January: Florida only; wing as in fig. 373 . .osmanthi Bush 15. Subbasal crossband joined to discal crossband posteriorly (fig. 374, a) 16 Subbasal and discal crossbands separate (fig. 380, b) .22 16. Subapical band absent; anterior and posterior apical bands present and joined to discal band to form an F-shaped pattern (figs. 374-377) {R. pomonella species group) .17 Only 1 apical band present, joined to subapical band (fig. 378). 20 17. Posterior apical band usually at least half as wide as discal band (fig. 374, b); aculeus no longer than 0.90 mm; infesting only Symphorocarpos spp. .zephyria Snow Posterior apical band usually less than half as wide as discal band (fig. 376, a); aculeus in northern specimens longer than 0.90 mm; hosts otherwise; occurring east of 110°W except pomonella also in Washington, Oregon, California, and Utah .18 18. Heavy dark brown to black shading often present on posterior surface of fore femur (fig. 387, a); infesting Rosaceae.pomonella (Walsh) Posterior surface of fore femur entirely yellow; infesting plants other than Rosaceae.. 19. Aculeus 0.75—0.85 mm long; infesting fruits of Ericaceae; wing as in fig. 376 .mendax Curran Aculeus more than 0.9 mm long; infesting fruits of Cornus spp.; wing as in fig. 377 .cornivora Bush 20. Aculeus relatively long, about 0.8 mm; host not known; wing as in fig. 378 .persimilis Bush

Figures 379-383. Right wings, Rbagoletis spp. 379, electromorpha Berl.; 380, berberis Curr.; 381, ebbettsi Bush; 382, ribicola Doane; 383, juniperina Marcovitch. Figures 384, 385. Dorsal view, scutellum, Rbagoletis spp. 384, striatella Wulp; 385, pomonella (Walsh). Figure 386. Posterior view of subscutellum and mediotergite, Rbagoletis completa Cress. Figure 387. Posterior view of fore femur, Rbagoletis pomonella (Walsh). Figure 388. Side view of epandrium, Rbagoletis cingulata (Lw.). Figure 389. Lateral view of thorax, Rbagoletis sp. Figures 390, 391. Basal posterior quarter of wing disk, Rbagoletis spp. 390, ribicola Doane; 391, juniperina Marc.

331

Systematic Treatment of the Genera

Figures 392, 393. Hind view, head, Rhagoletis spp. 392, ribicola Doane; 393, juniperina Marc. Figures 394, 395. Posterior external view, male genitalia, Rhagoletis spp. (redrawn from Westcott 1982). 394, pomonella (Walsh); 395, zephyria Snow.

Aculeus relatively short, about 0.7 mm; infesting dogwood berries (Cornus spp.) and blueberries (Vaccinium spp.) .21 21. Extreme apex of cell r4+5 along costa almost entirely hyaline (fig. 359, a);

subapical band bordering vein dm-cu dark or faint. .tabellaria (Fitch) More than 0.3 of extreme apex of cell r4+5 darkened by apex of apical band (fig. 379, a); subapical band always faint, narrowly bordering vein dm-cu .electromorpha Berlocher 22. Apical band touching costa along its entire length (fig. 380, c); infesting Oregon grape (Mahonia spp.).berberis Curran Apical band separated from costa by a hyaline area (fig. 381, b).23 23. Discal band joined to subapical band by an infuscated region in cell r4^s (fig. 381, a) .ebbettsi Bush Discal band not joined at all to subapical band (fig. 382, a).24

333

Rbagoletis 24. Usually with single strong anepisternal bristle (fig. 389, a); cell cup not sharply pointed (fig. 390, a); posterior surface of head with black horseshoe-shaped pattern (fig. 392); wing pattern as in fig. 382; infesting currants and gooseberries (Ribes spp.) .ribicola Doane Usually 2 or 3 strong anepisternal bristles; cell cup sharply pointed (fig. 391, a); black only across upper V3 of posterior surface of head, the lower 2/3 yellow (fig. 393); wing pattern as in fig. 383; infesting juniper berries (Juniperus spp.) .juniperina Marcovitch

Rhagoletis acuticornis (Steyskal) (Fig. 360; Map 64) Uropbora acuticornis Steyskal 1979: 43 (male holotype, NMNH; El Paso, Tex.). Rbagoletis acuticornis: Norrbom 1989: 59 (description, taxonomy, synonymy). R. acuticornis differs from all other species of Rbagoletis in having convergent posterior orbital bristles (as in fig. 27, a). It also has a distinctive wing pattern in which the apical band is reduced to two discrete spots touching the costa (fig. 360, a), completely separated from each other and from the subapical band. Vein CuA2 is nearly straight at its apex, thus producing, at most, only the slightest suggestion of an apical extension of cell cup. The scutum is shining black with little if any tomentum anteriorly and posteriorly; a white anepisternal stripe is present, but the postpronotal lobes are black; the scutellum is cream-colored centrally; the ab¬ dominal tergites are entirely black; and the legs are tawny with the basal third of the recognition.

middle femur and basal two-thirds of the hind femur black. distribution.

host.

Map 64.

The holotype was reared from a berry of Lycium berlanderi (Solanaceae).

discussion.

Norrbom (1989) transferred acuticornis from Uropbora. He dis¬

cussed in detail the relationships of this species and concluded that it does not logically belong to any of the species groups of Rbagoletis established and discussed by Bush (1966a) and Foote (1981). Other than striatella, it is the only member of the genus found in America north of Mexico that has a solanaceous host.

Rhagoletis basiola (Osten Sacken) Rose hip maggot (Fig. 361; Map 65) Trypeta flavonotata [misidentification]: Osten Sacken 1858: 79 (in catalog). - Loew 1862b: 58 (synonymy). - Loew 1873: 244, 245 (review). - Cockerell 1898: 149 (N. Mex.). Zonosema flavonotata [misidentification]: Loew 1873: 245 (taxonomy). Coquillett 1899c: 261 (synonymy, taxonomy). - Johnson 1900b: 687 (N.J.). V. T. Phillips 1923: 119 (review). - Johnson 1925b: 261 (N. Engl.). - Johnson

Systematic Treatment of the Genera

Rhagoletis acuticornis o Rhagoletis fausta • Rhagoletis ribicola A

Map 64. Distribution of Rhagoletis acuticornis, R. fausta, and R. ribicola.

1927: 217 (Mt. Desert). - Leonard 1928: 851 (N.Y.). - Johnson 1930: 151 (Nantucket). Spilographa flavonotata [misidentification]: Loew 1873: 327 (in key). - Doane 1900: 47 (taxonomy). - F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 604 (in catalog). - Washburn 1905: 118 (Minn.). - Tucker 1907: 104 (Colo.). - Johnson 1910: 801 (N.J.). - Muesebeck 1950: 254 (parasites). Trypeta (Zonosema) basiolum Osten Sacken 1877: 348 (female lectotype, MCZ; Brookline, Mass.). - Osten Sacken 1878: 190 (in catalog). - Bush 1966a: 510 (lectotype designation). Trypeta (Spilographa) flavonotata [misidentification]: Osten Sacken 1878: 190 (in catalog).

335

Rhagoletis

Map 65. Distribution of Rhagoletis basiola.

Zonosema basiolum: Coquillett 1899c: 261 (synonymy, taxonomy). - V. T. Phillips 1923: 128 (synonymy). — Johnson 1925b: 261 (N. Engl.). Spilographa setosa Doane 1899: 178 (female lectotype, NMNH; Vollmer, Ida.). — Coquillett 1899c: 261 (synonymy, taxonomy). - Doane 1900: 47 (taxonomy). Aldrich 1905: 604 (in catalog). - Grossbeck 1912: 378 (type data). - Robertson 1924: 15 (biology). - Stone 1951: 47 (synonymy, lectotype designation). - Foote 1966b: 124 (type data). - Zack 1984: 32 (type data). Zonosena [error] flavonotata: Johnson 1903b: 106 (N. Mex.). Spilographa basiolum: Aldrich 1905: 604 (in catalog).

Systematic Treatment of the Genera

Zonosema setosa: V. T. Phillips 1923: 128 (review). - Foxlee 1957: 37 (B.C.). “Trypetid”: Keifer 1930: 579 (description larva). Zonosema flavinotata [error]: Strickland 1938: 204 (Alta.). Zonosemata setosa: V. T. Phillips 1946: 98, 129 (description, biology larva; host). Hennig 1952: 220 (catalog immature stages). Rhagoletis basiola: Stone 1951: 45 (review). - Burks 1955: 81 (parasite). - Balduf 1957: 163 (puparia). — Balduf 1958: 7 (puparium). — Balduf 1959: 11 (review). Foote 1960d: 148 (taxonomy). - Foote and Blanc 1963: 55 (review, Calif.). Foote 1965a: 674 (in catalog). - Bush 1966a: 510 (revision). - Wasbauer 1972: 125 (hosts). - Wharton and Marsh 1978: 149 (parasites). - Prokopy and Berlocher 1980: 1319 (host). - Averill and Prokopy 1981: 222 (pheromone). Rhagoletis setosa: Stone 1951: 47, 48 (synonymy). Rhagoletis alternata [misidentification]: Muesebeck 1950: 254 (parasites). - Burks 1955: 81 (parasites). - Fischer 1964: 305, 359 (parasites). recognition. R. basiola and meigenii are distinctive among North American Rhagoletis in possessing an intercalary wing band (fig. 361) situated between the discal and subapical bands. In addition, the head, body, and legs are almost entirely yellowish, another feature by which both species can be distinguished from most others in America north of Mexico. R. basiola is distinguished from meigenii by two major features of the wing pattern: the subapical band is complete from the costa to vein CuAj (fig. 361, a), and the apical band is relatively narrow, occupying at most the apical one-sixth of cell r4+5 (fig. 361, b). See the recognition section of meigenii for parallel characters. distribution. Map 65. The most widely distributed North American Rhagoletis, basiola is commonly found in Alaska, much of Canada, and most of the United States. Although so far never collected in Mexico, basiola may occur in some of the northern parts of that country, where roses grow in a mild winter climate (Bush 1966a). hosts. R. basiola was reported by Balduf (1959) from 19 species of the genus Rosa; it appears to be restricted entirely to this plant genus. discussion. The publications of Balduf (1957, 1958, 1959) set forth most of what is known about the biology of basiola. Stone (1951) discussed its nomenclature and relationship to the European alternata (Fallen). Berlocher and Bush (1982) considered it to be a member of the alternata species group, which is otherwise Palearctic in distribution.

Rhagoletis berberis Curran (Fig. 380; Map 66) Rhagoletis berberis Curran 1932a: 8 (female holotype, AMNH; Hood R., Oreg.). Pickett 1937: 53 (taxonomy, male genitalia). - Jones 1942: 44 (hosts). - V. T. Phillips 1946: 61, 119 (description, biology larva; host). - Forsell 1947: 167

Rbagoletis

Map 66. Distribution of Rbagoletis berberis, R. meigenii, and R. striatella.

(Puget Sound). - Hennig 1952: 217 (catalog immature stages). - Foxlee 1957: 37 (B.C.). - Rohdendorf 1961: 93 (taxonomy). - Foote and Blanc 1963: 56 (review, Calif.). - Foote 1965a: 674 (in catalog). - Bush 1966a: 509 (revision). - Madsen 1970: 13 (biology). - Wasbauer 1972: 126 (hosts). - Wharton and Marsh 1978: 149 (parasite). - Arnaud and Owen 1981: 151 (type data). Rbagoletis berberis Jerm [error]: Fischer 1964: 358 (parasite). In addition to its host relationships, berberis is distinctive among all North American Rbagoletis in possessing the following combination of charac¬ recognition.

ters: the subbasal, discal, and subapical bands are all present and distinctly separated by hyaline areas (fig. 380); the single (anterior) apical band is undivided, fully joined to the subapical band, and follows the costa without any hyaline separation from it (fig. 380, c); the body is mainly black with a prominent yellow to whitish scutellar spot; and only the upper half of the posterior region of the head is darkened (as in fig. 393). DISTRIBUTION.

Map 66. R. berberis has not been recorded from anywhere in the

New World south of the United States. According to Bush (1966a), berberis has been recorded from two species of Mabonia (Oregon grape). Other species of that plant genus also may serve as hosts. Curran’s (1932a) host record of Berberis nervosa probably referred to hosts.

Mabonia.

338

Systematic Treatment of the Genera The ribicola group, including that species and berberis, was defined by Bush (1966a) only on characters of the male genitalia; the wing pattern alone is of no value in defining the group as presented in the abbreviated key. The biology of this species has not been studied under field conditions. discussion.

Rhagoletis boycei Cresson (Fig. 365; Map 67) Rhagoletis boycei Cresson 1929: 413

(male holotype, ANSP; Carr Canyon,

Huachuca Mts., Cochise County, Ariz.). - Barrett 1932: 296 (host). - Curran 1932a: 6 (in key). — Boyce 1934: 377 (review). — V. T. Phillips 1946: 119 (host). Wasbauer 1964: 5 (review). - Foote 1965a: 674 (in catalog). - Bush 1966a: 492 (revision). - Wasbauer 1972: 126 (hosts). - Wharton and Marsh 1978: 149 (para¬ site). — Foote 1981: 57 (in key, taxonomy). R. boycei is the only black-bodied species of Rhagoletis among those infesting walnuts (suavis group). It may be distinguished from all other black¬ recognition.

bodied Rhagoletis by the unmarked, wholly yellowish white scutellum. The wing pattern resembles those of juglandis and zoqui (Mexico) in that veins R4+5 or M, or both, are partially covered by narrow, elongated, dark infuscations (figs. 365, b; 366, a) in addition to the four major transverse bands. R. boycei is easily distinguished from juglandis by its dark body color and by the presence of a well-developed subbasal band (faint in juglandis) (see fig. 363, a).

Map 67. Distribution of Rhagoletis boycei, R. juglandis, and R. suavis.

339

Rhagoletis

Map 67. R. boycei was reported by Bush (1966a) from northern Sonora, Mexico, but has not been found south of that Mexican state. distribution.

R. boycei is restricted to walnuts, three species of Juglans having been recorded by Wasbauer (1972). hosts.

Apparently this is a rather rare species which is difficult to find in heavy infestations (Bush 1966a). discussion.

Rhagoletis chionanthi Bush (Fig. 372; Map 68) Rhagoletis cingulata Loew [misidentification]: F. H. Benjamin (part) 1934: 13 (host). — V. T. Phillips (part) 1946: 120 (host). Rhagoletis chionanthi Bush 1966a: 492 (female holotype, NMNH; Apopka, Fla.). Wasbauer 1972: 126 (hosts). R. chionanthi possesses a wing pattern (fig. 372) typical of the cingulata group as described in the abbreviated key and the recognition section under cingulata. In Florida, this species may be distinguished from cingulata by its considerably larger size, longer aculeus (about 0.95 mm versus 0.85 mm), and host preference. From osmantbi, chionanthi may be distinguished by its difference in recognition.

• Rhagoletis chionanthi o

Rhagoletis cingulata

▲ Rhagoletis indifferens A

Rhagoletis osmanthi

Map 68. Distribution of Rhagoletis chionanthi, R. cingulata, R. indifferens, and R. osmanthi.

340

Systematic Treatment of the Genera

seasonal distribution (see key), the slightly shorter aculeus (about 0.95 mm versus about 1.05 mm), and the extent of yellow coloration (see recognition characters for osmantbi). distribution. Map 68. According to Bush (1966a), the distribution of cbionantbi may be much wider than shown here, but it probably does not extend ouside the United States. host. R. cbionantbi has been reared only from Cbionantbus virginicus, which has a distribution much wider than the known range of this fly (Bush 1966a). discussion. Very little biological information is available for this apparently rare species.

Rhagoletis cingulata (Loew) Cherry fruit fly, cherry maggot, eastern cherry fruit fly, whitebanded cherry fruit fly, trypete des cerises (Figs. 371, 388; Map 68) Trypeta cingulata Loew 1862c: 76 (female syntype, MCZ; “Middle States”). - Loew 1873: 263 (taxonomy). — Lowe 1899: 693 (description, host). Rbagoletis cingulata: Loew 1873: 263, 329 (taxonomy). - Doane 1898: 70 (in key). - Slingerland 1899: 23 (biology). - Johnson 1900b: 687 (N.J.). - Slingerland 1902: 28 (nomenclature, biology). - Chittenden 1904: 70 (biology). - Aldrich 1905: 606 (in catalog). - Aldrich 1909: 69 (in key). - Gibson 1909: 73 (syn¬ onymy).-Johnson 1909: 113 (distribution). - Bezzi 1910: 15 (review).-Johnson 1910: 801 (N.J.). - Banks 1912: 33 (description larva). - Illingworth 1912a: 135 (host). - Illingworth 1912b: 191 (taxonomy). - Herrick 1913: 79 (biology). Caesar 1915: 107 (biology). - Britton 1920: 204 (Connecticut). - V. T. Phillips 1923: 135 (review). - Glasgow and Gambrell 1925: 1 (review). - Johnson 1925b: 261 (N. Engl.). - Pettit 1925: 1 (review). - Leonard 1928: 851 (N.Y.). - Cresson 1929: 408 (review). — C. T. Greene 1929: 494 (description larva, pupa). — Pettit and Tolies 1930: 1 (review). - Gambrell 1931: 226 (biology). - Glasgow 1931: 431 (hosts). - Curran 1932a: 7 (in key, taxonomy). - Petch and Maltaise 1932: 56 (Que.). — Farleman 1933: 825 (biology). — F. H. Benjamin (part) 1934: 13 (review, Fla.). - Pickett 1937: 53 (taxonomy, male genitalia). - Quayle (part) 1938: 431 (review). - Cameron 1941: 14 (parasite). - Middlekauff 1941: 621 (biology, para¬ sites). - V. T. Phillips 1946: 65, 120 (description, biology larva; host). - Cox 1952: 1 (hosts, biology). — Hennig 1952: 217 (catalog immature stages). — Peterson 1953. 333, 335, 337 (figures of larva). — Simkover 1953: 896 (crossing). — Frick et al. 1954: 1 (biology). — Blanc and Keifer 1955: 77 (crossing, taxonomy). — Frings and Frings 1955: 431 (chemoreceptors). - Muesebeck 1956: 99 (parasite). - Raine and Andison 1958: 593 (biology). - Christenson and Foote 1960: 171 (biology). Still 1960: 967 (trap). - Fischer (part) 1964: 286 (parasite). - Foote 1965a: 674 (in catalog, as subspecies). - Bush 1966a: 473 (revision). - Wasbauer 1972: 126

341

Rhagoletis

(hosts). — Weems 1972a: 1 (review). — AliNiazee and Brown 1974: 93 (bibliogra¬ phy). — Jubb and Cox 1974: 613 (biology). — Prokopy 1975d: 420 (trap). — Prokopy 1977a: 593 (lures). — Wharton and Marsh 1978: 149 (parasites). — Berlocher 1980a: 131 (electrophoretic key). - Muesebeck 1980: 32 (parasite). Trypeta (Rhagoletis) cingulata: Osten Sacken 1878: 191 (in catalog). Trypeta pomonella [misidentification]: A. J. Cook 1889: 152 (hosts). - Cordley (part) 1889: 192 (taxonomy, hosts). As described in the abbreviated key to species groups, the subapical band of the wing pattern in cingulata extends completely across the disk and is entirely separated from the discal band (figs. 370—373). The apical band is broadly joined to the subapical band anteriorly and is divided apically into two arms, the posterior one always complete and terminating near the apex of vein M, the anterior one usually broken to produce a fuscous spot at the apex of vein R4+5 (fig. 371, b). About 65—80% of cingulata individuals show this spot, whereas in indifferens the spot is present in at most 5% of individuals; the anterior arm is never completely broken in cbionantbi and osmantbi (figs. 372, 373). See the recognition section for indifferens for additional characters separating these two species. In Florida, cingulata may be distinguished from cbionantbi and osmantbi by the break in the anterior arm of the apical band in most specimens, an oviscape length/width ratio of about 6, the aculeus about 0.85 mm long, the generally smaller size, and the host recognition.

preferences. distribution.

Map 68. R. cingulata has been reported from Texcoco in central

Mexico (Foote 1981). hosts.

The black cherry, Prunus serotina, is the principal native host of cingulata.

At least six additional species of Prunus also are recorded. discussion.

R. cingulata was one of the first Rhagoletis species to be recognized

in the United States as economically important, and today it remains of prime significance in commercial cherry production in the eastern half of the country.

Rhagoletis completa Cresson Walnut husk fly, mouche des brous du noyer (Figs. 364, 386; Map 69) Rhagoletis juglandis [misidentification]: Boyce 1929a: 269 (biology). - Boyce 1929b: 861 (biology). Rhagoletis suavis completa Cresson 1929: 412 (male holotype, ANSP; Chino, Calif.). - Boyce 1930: 249 (biology). - Barrett 1932: 296 (hosts). - Arnaud 1979: 332 (type data). Rhagoletis suavis var. completa: Keifer 1930: 579 (description, larva). Rhagoletis completa: Boyce 1931: 1018 (diapause). - Boyce 1932: 1053 (biology). Curran 1932a: 6 (in key). - Boyce 1933: 813 (biology). - Marlatt 1933: 3 (para-

342

Systematic Treatment of the Genera

Map 69. Distribution of Rhagoletis completa.

sites). — Boyce 1934: 363 (biology). — Pickett 1937: 53 (taxonomy, male genitalia). — Quayle 1938: 388 (review). — Boyce and Bartlett 1941: 318 (lures). — V. T. Phillips 1946: 64, 120 (biology, description larva; host). - Benjamin and Hodson 1949: 707 (lures, distribution). - Hennig 1952: 217 (catalog immature stages). Peterson 1953: 333, 337 (figures of larvae). - R. L. Metcalf et al. 1955: 222 (physiology). - Clausen 1956a: 178 (parasites). - Clausen 1956b: 25 (parasites). Michelbacher and Sisson 1956: 13 (review, Calif.). - Barnes and Ortega 1958: 532 (lure). - Barnes and Osborn 1958: 686 (lures). - Christenson 1958: 11 (lure). Essig 1958: 604 (review). — Moffitt 1958: 551 (sex determination). — Barnes 1959: 90 (flight range). - Barnes and Ortega 1959: 279 (lures). - Ebeling 1959: 348 (review). - Christenson and Foote 1960: 171 (biology). - Foote and Blanc 1963: 58 (review, Calif.). - Wasbauer 1964: 5 (review). - Foote 1965a: 674 (in catalog). Harper 1965: 82 (dispersal). - Bush 1966a: 488 (revision). - Bateman 1972: 495 (biology). - Cirio 1972: 99 (biology). - Wasbauer 1972: 126 (hosts). - Steyskal 1973: 522 (in key, larvae in walnut husks). - Tsiropolous 1976: 83 (bacterial associates). - Wharton and Marsh 1978: 149 (parasites). - Berlocher 1980a: 131 (electrophoretic key). - Muesebeck 1980: 36, 40 (parasites). - Riedl et al. 1981: 21 (attractants). - Berlocher 1984c: 392 (genetics). - Berlocher 1984d: 906 (genet¬ ics). - Elzinga and Broce 1986: 208 (labellar morphology). - Legner and Goeden 1987: 739 (parasites).

recognition. The four transverse wing bands are present and are usually dis¬ tinctly separated by hyaline areas (fig. 364), except in occasional specimens in

343

Rhagoletis

which the discal and subapical bands are connected posteriorly (fig. 364, b). In the former case, the pattern closely resembles that of berberis (fig. 380), but the host relationships of these two species are quite different. The thorax and abdomen of completa are golden yellow with darker markings (completely black in berberis), and the scutellum is concolorous yellow (black with a distinct yellow spot in ber¬ beris).

distribution.

Map 69. R. completa also is known to occur in Monterrey, Nueva

Leon, Mexico. No fewer than seven species of Juglans were reported as hosts by Wasbauer (1972). Bush (1966a) indicated that the several occurrences of completa reported in peach (Prunus persica) is not surprising because, under certain condi¬ tions, that fruit is highly susceptible to attack by many species of fruit flies. hosts.

Rhagoletis cornivora Bush (Fig. 377; Map 70) Rhagoletis pomonella [misidentification]: F. H. Benjamin (part) 1934: 14 (note, adult). — Hall 1943: 202 (cross-breeding). — V. T. Phillips (part) 1946: 120 (host). — Christenson and Foote 1960: 184 (biology). Rhagoletis sp.: Hall 1938: 56 (biology, Ont.). Rhagoletis cornivora Bush 1966a: 470 (male holotype, MCZ; Lincoln, Mass.). Wasbauer 1972: 127 (hosts). - Wharton and Marsh 1978: 149 (parasite). Berlocher 1980a: 131 (electrophoretic key). — Berlocher 1984a: 240 (taxonomy, hosts). The wing pattern of cornivora (fig. 377) resembles those of pomonella, zephyria, and mendax in that the subapical band is missing, and the anterior and posterior apical bands are broadly connected to the discal band to form an F-shaped pattern in the distal half of the wing. R. striatella possesses a similar pattern in the apex of the wing (fig. 369) but has separate discal and subapical bands and may be separated easily from cornivora by this and other characters given in the key. R. cornivora may be distinguished from mendax by the length of the aculeus (see key to species) and by host data. Northern specimens of cornivora are distinct morphologically from those of mendax by the shape of the surstylus (see Bush 1966a), a character not used in this handbook. The recognition of Florida specimens recognition.

of cornivora depends almost entirely on host data.

distribution.

hosts.

Map 70.

Four species of Cornus were recorded as hosts by Bush (1966a) and

Wasbauer (1972).

Systematic Treatment of the Genera

▲ Rbagoletis cornivora o Rbagoletis mendax • Rbagoletis zepbyria

Map 70. Distribution of Rbagoletis cornivora, R. mendax, and R. zepbyria.

Rhagoletis ebbettsi Bush (Fig. 381; Map 71) Rhagoletis ebbettsi Bush 1966a: 504 (male holotype, NMNH; 4 mi. e. Ebbetts Pass, Alpine County, Calif.). In wing pattern, R. ebbettsi resembles other species of the tabellaria group (see abbreviated key) in that the subbasal, discal, and subapical bands RECOGNITION.

are present, and the single (anterior) apical band is apically undivided and separated from the costa by a hyaline area. It is distinguishable from those species by an extensive connection between the discal and subapical bands anterior to vein M (fig. 381, a). A connection between these bands exists in some other North American

345

Rhagoletis

Rhagoletis species (suavis, [fig. 368, a]; some specimens of striatella; fausta [fig. 367, a]; and completa [fig. 364, b]), but in all those species, the connection is more posterior, and the apical band joins the costa without an intervening hyaline area.

DISTRIBUTION.

host.

Not

Map 71.

known.

discussion.

R. ebbettsi is extremely rare. To our knowledge, only one specimen

(the rather badly damaged holotype) has ever been seen.

Systematic Treatment of the Genera

Rhagoletis electromorpha Berlocher (Fig. 379; Map 71) Rhagoletis electromorpha Berlocher 1984a: 237 (male holotype, INHS; Morton Arboretum, Lisle, Ill.).

The wing patterns of tabellaria, persimilis, and electromorpha (figs. 359, 378, and 379, respectively) are quite similar in exhibiting the following recognition.

combinations of characters: subbasal, discal, and subapical bands present, the subbasal and discal bands distinctly connected posteriorly and the single (anterior) apical band undivided apically and separated from the costa by a hyaline area. The subbasal and discal bands are united, and the apical band is separated from the costa in the pomonella group, but the subapical band is absent. Bush (1966a) states that the only differences between tabellaria and persimilis lie in the characters of the internal male genitalia and the spermathecae of the female. Among North American Rhagoletis, only tabellaria and electromorpha have a membranous sac at the base of the distiphallus in the male (Berlocher 1984a). According to Berlocher (1984a), the following overlapping, comparative characters should aid in distinguishing electro¬ morpha from tabellaria:

R. electromorpha: (a) Subapical band a lighter color narrowly bordering vein dm-cu. (b) Half or more of apex of apical band terminating anterior to vein M (fie 379, a). (c) Dark marking more prominent on posterior region of head. (d) Scutellar spot extending farther toward apex of scutum. R. tabellaria: (a) Subapical band not or only slightly lightened bordering vein dm-cu. (b) Nearly entire apex of apical band terminating posterior to vein M (fig. 359, a). (c) Dark marking less prominent on posterior region of head. (d) Scutellar spot more distantly removed from apex of scutum.

DISTRIBUTION. Map 71.

hosts.

Berlocher (1984a) listed Cornus drummondi and C. racemosa as hosts.

Berlocher (1984a) distinguished electromorpha primarily on the basis of electrophoretic characters. It is considered to be an electromorph because discussion.

only such characters completely distinguish it from tabellaria. Morphological differ¬ ences between the two species overlap and were discovered subsequent to the elec¬ trophoretic investigations.

347

Rbagoletis

Rhagoletis fausta (Osten Sacken) Black cherry fruit fly, black-bodied cherry fruit fly, trypete noire des cerises (Fig. 367; Map 64) Trypeta (Acidia) fausta Osten Sacken 1877: 346 (female lectotype, MCZ; Mt. Wash¬ ington, N.H.). - Osten Sacken 1878: 189 (in catalog). - Bush 1966a: 518 (lec¬ totype designation). Trypeta fausta: Slosson 1895: 7 (Mt. Washington). Rhagoletis fausta: Coquillett 1899c: 260 (taxonomy). - Chagnon 1901: 13 (Can.). Aldrich 1909: 70 (in key). - Bezzi 1910: 19 (review). - Illingworth 1912b: 193 (biology). - Herrick 1913: 79 (biology). - Caesar 1915: 107 (biology). - Severin 1918b: 325 (biology, distribution). -V. T. Phillips 1923: 136 (review). - Glasgow and Gambrell 1925: 1 (review). — Johnson 1925b: 262 (N. Engl.). - Pettit 1925: 1 (review). — Bird 1927: 128 (Man.). - Johnson 1927: 217 (Mt. Desert). — Leonard 1928: 851 (N.Y.). — Cresson 1929: 404 (review). - Gambrell 1931: 226 (biology). — Glasgow 1931: 431 (hosts). - Curran 1932a: 6 (in key). — Farleman 1932: 1048 (oviposition). — Petch and Maltaise 1932: 56 (Que.). - Eichmann 1936: 7 (biology, Mont.). — Pickett 1937: 53 (taxonomy, male genitalia). — Essig 1938: 604 (review). — Quayle 1938: 431 (review). — Crandell 1939: 632 (parasite). — Mackie 1940: 157 (review, Calif.). — Cameron 1941: 14 (parasite). — Middlekauff 1941: 623 (biology). — Burks 1943: 583 (parasite). — Breakey 1945: 119 (biology). — V. T. Phillips 1946: 66, 120 (description, biology larva; host). - Forsell 1947: 167 (Puget Sound, host). - Cox 1952: 1 (biology, hosts). - Hennig 1952: 217 (catalog immature stages). - Peterson 1953: 333, 335, 337 (figures of larva). — Proverbs 1953: 18 (B.C.). - Frick et al. 1954: 42 (biology). - Clausen 1956a: 178 (para¬ sites). — Clausen 1956b: 25 (parasites). — Essig 1958: 604 (review). - Raine and Andison 1958: 593 (biology). - Still 1960: 967 (trap). - Foote and Blanc 1963: 60 (review, Calif.). - Fischer 1964: 279, 293 (parasites). - Foote 1965a: 674 (in catalog). - Bush 1966a: 518 (revision). - Newcomer 1966: 39 (review). - Madsen 1970: 13 (biology). - Banham 1972: 25 (review, B.C.). - Wasbauer 1972: 127 (hosts). - AliNiazee and Brown 1974: 93 (bibliography). - Jubb and Cox 1974: 613 (biology). - Prokopy 1975c: 298 (pheromones). - Prokopy 1976a: 899 (biolo¬ gy). - Prokopy 1977a: 593 (lures). - Wharton and Marsh 1978: 149 (parasites). Berlocher 1980a: 131 (electrophoretic key). - Muesebeck 1980: 32, 36 (parasites). Acidia fausta: Aldrich 1905: 603 (in catalog). Rhagoletis intrudens Aldrich 1909: 70 (female holotype, NMNH; Victoria, B.C.). Gibson 1909: 73 (biology, synonymy). - Aldrich 1910: 99 (nomenclature). Illingworth 1912b: 193 (synonymy). - Cresson 1929: 404 (hosts, synonymy). Bush 1966a: 518 (synonymy). Rhagolitis [error] fausta: Van Duzee 1911: 244 (host, Ont.). recognition. In the preceding abbreviated key to species groups, we placed fausta together with striatella because the posterior apical band in both species arises

348

Systematic Treatment of the Genera

from the subapical band in the vicinity of vein r-m, making an F-shaped pattern in the apical half of the wing similar to that in the pomonella group (however, in the latter, note that the subapical band is missing and the apical bands are connected to the discal band). In wing pattern alone, fausta is unique among North American

Rhagoletis in combining a very broad connection between the discal and subapical bands in cell dm (fig. 367, a) with the presence of both an anterior and posterior apical band, the latter arising in much the same location as in striatella. In striatella, the discal and subapical bands are separate or connected only along the posterior wing margin (Foote 1981). In many respects, the wing pattern of fausta resembles that of suavis (fig. 368), but fausta has both anterior and posterior apical bands and an isolated hyaline spot in the distal half of cell cuaj (fig. 367, b). R. suavis has a yellowish body; that of fausta is black and without yellowish bands at the posterior margins of the abdominal tergites. distribution. Map 64.

R. fausta infests the fruit of Pvunus ewiavginata over most of its western range, whereas in the east, the pin or fire cherry (P. pennsylvanica) is the preferred hosts.

host. Five other species of Prunus have been identified as hosts (Wasbauer 1972). discussion. We follow Bush (1966a) in considering the eastern and western

populations of fausta as a single species because no morphological differences be¬ tween the two can be found. Berlocher and Bush (1982) suggested that fausta is most closely related to the suavis species group; we have already pointed out how these two species are similar in wing pattern.

Rhagoletis indifferens Curran Western cherry fruit fly (Figs. 57, 370; Map 68)

Rhagoletis cingulata: Wilson and Lovett 1913: 160 (biology). — Cole and Lovett 1921: 325 (Oreg.). - Lovett 1923: 1 (biology). - Bird 1927: 128 (Man.). - F. H. Benjamin 1934: 13 (taxonomy). - Essig 1938: 604 (review). - Hatch 1938: 197 (bibliography, Wash.). - Jones 1938: 75 (biology). - Keifer 1938: 662 (Calif.). Quayle 1938: 431 (review). - Mackie 1940: 157 (review, Calif.). - Breakey 1945: 119 (biology). - Jones 1945a: 55 (genitalia, crossing). - Jones 1945b: 116 (biolo¬ gy). - Eide 1946: 55 (description, biology). - Eide 1947: 119, 123 (biology, parasite). - Forsell 1947: 167 (Puget Sound, hosts).-Eide et al. 1949: 1 (biology). — Frick 1949: 99 (biology). — Newcomer 1950: 31 (review, biology). — Frick 1952: 262 (biology, lure). - Frick 1953: 893 (larval survey). - Frick et al. 1954: 1 (biology). - Kamal 1954: 959 (biology). - Jones and Wallace 1955: 616 (biology). — Clausen 1956a: 178 (parasites). — Clausen 1956b: 25 (parasites). — Essig 1958: 604 (review). - Fleschner 1963: 114 (parasites). - Fischer (part) 1964: 286 (para¬ site). - Muesebeck 1980: 40 (parasite). Cherry fruit fly: Frank 1924: 42 (biology).

349

Rhagoletis

Rhagoletis indifferens Curran 1932a: 8 (male holotype, AMNH; Hood R., Oreg.). — Curran 1934: 286 (figure of wing). — Pickett 1937: 53 (taxonomy, male genitalia). -Jones 1942: 44 (hosts). — V. T. Phillips 1946: 66, 120 (description, biology larva; hosts). — Hennig 1952: 217 (catalog immature stages). - Simkover 1953: 896 (crossing). - Bush 1966a: 477 (revision). - Newcomer 1966: 39 (review). - Arrand and Peters 1968: 40 (B.C.). - Madsen 1970: 13 (biology). - Banham 1971: 29 (hosts). — Banham 1972: 25 (review, B.C.). — Wasbauer 1972: 28 (hosts). — AliNiazee 1973a: 103 (nomenclature, biology). — AliNiazee 1973b: 75 (biology). — Banham 1973: 13 (traps). — AliNiazee 1974a: 909 (biology). - AliNiazee 1974b: 1201 (behavior). — AliNiazee and Brown 1974: 93 (bibliography). — AliNiazee 1975: 1011 (biology). - AliNiazee 1976: 397 (biology). - AliNiazee and Brown 1977: 1227 (laboratory rearing). — Brown and AliNiazee 1977: 678 (laboratory rearing). - AliNiazee 1978: 1133 (traps). - Wharton and Marsh 1978: 149 (para¬ sites). — AliNiazee 1979: 1101 (phenology). - Berlocher 1980a: 131 (electrophore¬ tic key).

Rhagoletis cingulata indifferens: Blanc and Keifer 1955: 1977 (crossing, taxonomy). - Muesebeck 1956: 99 (parasites). - Frick 1957: 584 (biology). - Christenson and Foote 1960: 171 (biology). - Ellertson 1961: 116 (hosts). - Foote and Blanc 1963: 57 (review, Calif.). - Foote 1965a: 674 (in catalog).

R. indifferens is similar to cingulata in wing pattern (figs. 370, 371) (see abbreviated key and recognition characters for cingulata), but the anterior arm of indifferens is broken to produce an apical spot in only about 5% of individu¬ recognition.

als in contrast to cingulata, in which this spot is much more commonly encountered (fig. 371, b). In addition, other characters that distinguish indifferens from cingulata are as follows:

R. indifferens: (a) Apical yellow shading on posterior margin of tergite 5 of male lacking. (b) Black shading always present on posterior surface of fore coxa. (c) Epandrium dark-colored.

R. cingulata: (a) Apical yellow shading on posterior margin of tergite 5 of male present. (b) Fore coxae concolorous yellow. (c) Epandrium light-colored (fig. 388, a). Most individuals of indifferens may be distinguished from those of chionanthi and

osmanthi by the differences in geographical distribution and hosts and by the gener¬ ally smaller size and lesser development of the wing bands (compare figs. 370, 372, and 373). distribution. Map 68. HOSTS. The wild pin cherry,

Prunus emarginata, is the native host (Frick 1957).

Five additional species of Prunus were recorded by Wasbauer (1972).

Systematic Treatment of the Genera

discussion. Although

indifferens was originally described by Curran as a valid

species, a considerable amount of morphological and biological information is pre¬ sented in the literature under this name as a subspecies of cingulata.

Rhagoletis juglandis Cresson (Fig. 363; Map 67)

Rhagoletis juglandis Cresson 1920: 65 (male holotype, ANSP; Carr Canyon, Huachuca Mts., Cochise County, Ariz.). — Cresson 1929: 413 (review). — Barrett 1932: 296 (hosts). - Curran 1932a: 6 (in key). - Boyce 1934: 376 (review). - Essig 1938: 604 (review). - V. T. Phillips 1946: 67, 120 (description, biology larva; host). - Hennig 1952: 218 (catalog immature stages). - Essig 1958: 604 (review). -Muesebeck 1961: 57 (parasite). - Foote 1965a: 674 (in catalog). - Bush 1966a: 490 (revision).-Wasbauer 1972: 128 (hosts).-Steyskal 1975b: 231 (in key, larvae in walnut husks). - Wharton and Marsh 1978: 149 (parasite). - Muesebeck 1980: 36 (parasite). - Nation 1981: 121 (sex pheromone glands).

recognition.

R. juglandis is the only North American Rhagoletis with the fol¬

lowing combination of characters: thorax and abdomen entirely yellow and un¬ marked, subbasal wing band faint or in some specimens nonexistent, veins R4+5 and sometimes M with an elongated, narrow fuscous covering. The only other com¬ pletely yellow-bodied Rhagoletis in North America, basiola and meigenii, are re¬ stricted to Rosaceae and Berberidaceae, respectively, and have quite different wing patterns with an intercalary band (figs. 361, 362). distribution. Map 67.

R. juglandis extends south into central Mexico and

possibly farther. Juglans spp. occur well into Central America.

hosts.

R. juglandis has been reared from only two species of Juglans in the

United States and Mexico.

discussion.

R. juglandis is sympatric with boycei over part of its range.

Rhagoletis juniperina Marcovitch (Figs. 383, 391, 393; Map 72)

Rhagoletis juniperinus Marcovitch 1915: 171 (male lectotype, MCZ; Six Mile, Ithaca, N.Y.). - V. T. Phillips 1923: 135 (review). - Leonard 1928: 852 (N.Y.). Cresson 1929: 406 (synonymy). - V. T. Phillips 1946: 69, 120 (description, biolo¬ gy larva; host). - Hennig 1952: 218 (catalog immature stages). - Foote 1965a: 675 (in catalog). — Bush 1966a: 500 (lectotype designation).

Rhagoletis juniperina: Hering 1958: 4 (taxonomy). - Rohdendorf 1961: 93 (emen¬ dation). - Bush 1966a: 500 (revision). - Wasbauer 1972: 128 (hosts).

Rhagoletis

o

Rhagoletis juniperina

•

Rhagoletis tabellaria

Map 72. Distribution of Rhagoletis juniperina and R. tabellaria.

recognition.

R. juniperina and ribicola are the only two North American Rha¬

goletis with the following combination of wing characters (figs. 382, 383): subbasal, discal, and subapical bands all present, distinct, and sometimes well separated from each other; one apical (anterior) band, broadly joined to the subapical, not divided at its apex as in members of the cingulata group (figs. 370-373) and distinctly sepa¬ rated from the costa by a narrow hyaline area in the apices of cells r2-1-3 and ^+5. These two species are readily distinguished from tabellaria, persimilis, and electro-

morpha by the absence of a posterior connection between the subbasal and discal bands, from ebbettsi (fig. 381) by the complete separation of the discal and subapical bands, and from completa by the hyaline separation of the apical band from the costa. In addition to differences in host preference, juniperina and ribicola are dis-

352

Systematic Treatment of the Genera

tinguishable from each other by the absence in the former of a complete, dark horseshoe-shaped mark on the posterior surface of the head (compare figs. 392 and 393) and by the narrow apical prolongation of cell cup (compare figs. 390 a and 391, a).

R. juniperina is not known to occur south of the United States despite the presence of Juniperus spp. in Mexico. distribution. Map 72.

Juniperus have been recorded as hosts, but this fly also occurs outside the recorded range of those species, suggesting that other Juniperus hosts. Several species of

species also may be hosts (Bush 1966a).

discussion.

R. juniperina is widespread north of Mexico but is not often en¬

countered.

Rhagoletis meigenii (Loew) (Fig. 362; Map 66)

Trypeta meigenii Loew 1844: 316 (syntypes, sex and depository not stated; Eu¬ rope?).

Zonosema meigenii: Loew 1862b: 44 (taxonomy). Rhagoletis meigenii: Hendel 1927: 74 (taxonomy). - Foote 1984: 122 (in catalog, Palearctic Region). — White 1988: 39 (in key, British species).

recognition.

R. meigenii and basiola are the only two Rhagoletis in America

north of Mexico having an intercalary band in the wing (fig. 361, a) and almost entirely brown to yellowish head, body, and legs. R. meigenii can be distinguished from basiola by wing characters: the subapical band in the former extends from vein CuA2 only to (at most) vein R2+3 (fig. 362, a), and the apical band is comparatively wide, occupying approximately the apical one-third of cell r4+5 (fig. 362, b). distribution. Map 66.

R. meigenii is widespread in northern and central Eu¬

rope and the western portions of the former USSR and is to be regarded as an immigrant species in the United States, having been found so far only on the limited area shown on the map.

host. In the Old World,

meigenii breeds in fruits of Berberis vulgaris, the alter¬

nate host of the stem rust of wheat. This plant has been introduced and is the probable host of meigenii in the United States.

meigenii has been present in the United States since at least 1986. Like basiola, it is a member of the alternata species discussion. Collection records indicate that

group.

353

Rbagoletis

Rhagoletis mendax Curran Blueberry maggot, blueberry fruit fly, mouche de Pairelle (Fig. 376; Map 70)

Rbagoletis pomonella [misidentification]: Britton 1906: 260 (host).-Johnson (part) 1910: 802 (N.J.). - Illingworth (part) 1912a: 129 (review, biology). - O’Kane (part) 1914: 1 (biology). - Woods 1914: 398 (host). - Woods 1915: 293 (parasite). - Woods 1916: 252 (review). - Gahan 1919: 167 (possible parasite). - Herrick (part) 1920: 89 (biology). - Patch and Woods 1922: 77 (biology). - V. T. Phillips (part) 1923: 136 (review). - Brues (part) 1924: 129 (races). - Guyton and Knull (part) 1925: 3 (review, Pa.). - Porter (part) 1928: 1 (review). - Cresson (part) 1929: 409 (hosts). - Gambrell (part) 1931: 226 (biology). - Lathrop and McAlis¬ ter 1931: 1 (biology). - Lathrop and Nickels 1931: 260 (biology). - McLaine 1931: 68 (Canada). - Phipps 1931: 132 (review, Maine). - Lathrop and Nickels 1932: 1 (biology). - McAlister 1932: 415 (biology). - Lathrop and Newton 1933: 143 (biology of parasite). - McAlister and Anderson (part) 1935: 675 (biology, crossing). - Pickett (part) 1937: 53 (male genitalia, crossing). - Pickett and Neary (part) 1940: 551 (crossing, hosts). - E. Cameron 1941: 15 (parasite). - Chandler 1943: 115 (biology). - Beckwith 1945: 43 (review, N.J.). - Tomlinson 1951: 266 (biology). - Commonw. Inst. Entomol. (part) 1954: Map No. 48 (distribution).

Rbagoletis mendax Curran 1932a: 7 (male holotype, AMNH; Maine). - F. H. Benjamin 1934: 15 (taxonomy). - Darlington 1943: 217 (biology). - V. T. Phillips 1946: 70, 120 (description, biology larva; hosts). - Hennig 1952: 218 (in catalog immature stages). - Lathrop 1952: 1 (review). - Peterson 1953: 331 (figure of larva). - Christenson and Foote 1960: 183 (biological races). - Eaton and Hall 1961: 26 (note, biology). - Maxwell and Wood 1961: 26 (review, Maritime Prov.). - Foote 1965a: 674 (in catalog). - Bush 1966a: 466 (revision). - Simon 1969: 169 (serology). - Roth and Richardson 1970: 496 (hosts). - Wasbauer 1972: 128 (hosts). - Prokopy and Coli 1978: 45 (trapping). - Wharton and Marsh 1978: 149 (parasites). - Berlocher 1980a: 131 (electrophoretic key). - Arnaud and Owen 1981: 151 (type data). - Smith and Prokopy 1981: 462 (biology). - Smith and Prokopy 1982: 388 (mating behavior). recognition. The wing pattern of mendax (fig. 376) resembles closely those of

pomonella, zepbyvia, and cotnivova in that the subapical band is absent, and the anterior and posterior apical bands are connected instead to the discal band, forming an F-shaped pattern in the apical half of the wing. R. striatella has a somewhat similar wing pattern but may be distinguished from mendax by the characters given in the key. R. mendax may be distinguished from pomonella by the lack of dark markings on the posterior surface of the fore femur, the shorter aculeus, and the smaller wing band ratio (see recognition characters for pomonella). R. mendax,

pomonella, and cornivora can be distinguished reliably from each other in Florida only by the use of host data, as individuals of pomonella are smaller and the females have a shorter aculeus than in the northern parts of its range.

Systematic Treatment of the Genera

DISTRIBUTION. Map 70.

R. mendax has long been known to infest commercial plantings of blue¬ berries, and no fewer than 11 additional species of Vaccinium have been recorded as HOSTS.

hosts (Wasbauer 1972). Three species of Gaylussacia also are regarded as preferred hosts (Bush 1966a).

discussion. We remind users of this handbook that many of the taxonomic,

host, distributional, and biological records of mendax appear under the name

pomonella. Geddes et al. (1987) presented a complete general review of this species.

Rhagoletis osmanthi Bush (Fig. 373; Map 68)

Rhagoletis cingulata Loew: F. H. Benjamin (part) 1934: 13 (host). - V T Phillips (part) 1946: 120 (host).

Rhagoletis osmanthi Bush 1966a: 478 (male holotype, NMNH; Osceola County Fla.). - Wasbauer 1972: 130 (host).

R. osmanthi has a wing pattern (fig. 373) typical of the cingulata group as described in the abbreviated key and recognition section for cingulata. It has a longer aculeus (about 1.05 mm) than either cingulata (about 0.85 mm) or RECOGNITION.

chionanthi (about 0.95 mm). In osmanthi, the posterior cranial regions of the head are never darkened, there are never any dark leg maculations or shading, and the pleural regions of the thorax and the dorsum of the abdomen are more extensively yellow than in either cingulata or chionanthi. distribution. Map 68.

R. osmanthi is known from only one host plant, Osmanthus americanus, the wild tea-olive or devil wood. According to Bush (1966a), other species of Os¬ manthus also may serve as hosts. HOSTS.

Rhagoletis persimilis Bush (Fig. 378; Map 71)

Rhagoletis persimilis Bush 1966a: 503 (male holotype, CNC; Robson, B.C.). RECOGNITION. The wing patterns of

tabellaria, persimilis, and electromorpha

(figs. 359, 378, and 379, respectively) are similar in exhibiting the following combi¬ nation of characters: subbasal, discal, and subapical bands present, the first-named distinctly connected to the discal band posteriorly and the single (anterior) apical band undivided apically and separated from the costa by a hyaline area. The subbasal and discal bands are united, and the apical band is separated from the costa in the pomonella group (figs. 374-377), but the subapical band is absent. The only

355

Rhagoletis character separating this species from tabellaria is the relative length of the aculeus, that of the latter averaging about 0.7 mm, the former about 0.8 mm. Other distin¬ guishing characters are found only in the male genitalia. DISTRIBUTION. Map 71.

host. Not known.

discussion. R. persimilis is extremely rare; only a few specimens have ever been

discovered.

Rhagoletis pomonella (Walsh) Apple maggot, apple maggot fly, apple fruit fly, apple worm, railroad worm, mouche de la pomme (Figs. 61, 375, 385, 387, 394; Map 73) Note: The literature of pomonella is so extensive that the compilation of a com¬ plete bibliography has not been possible. Two bibliographies already in existence, those by Rivard (1968) and Dean and Chapman (1973), will provide the reader with most of the references that have appeared up to or slightly beyond 1970. The follow¬ ing synonymy presents those useful to anyone who wishes to investigate this species at a biosystematics level. Trypeta albiscutellata Say in T. W. Harris 1835b: 600 (nomen nudum). - Osten Sacken 1858: 78 (in catalog). - Loew 1862c: 57 (taxonomy). - Johnson 1925a: 97 (synonymy). Trypeta pomonella Walsh 1867: 343 (female lectotype, MCZ; Ill.). - B. D. Walsh 1868: 33 (description, biology). - Walsh and Riley 1869: 59 (N.Y.). - Loew 1873: 265 (review). — Comstock 1882: 195 (biology, hosts). — A. J. Cook 1889: 152 (biology). - G. C. Davis 1889: 291 (biology, hosts). - Harvey 1890a: 1 (biology). Harvey 1890b: 190 (biology). - Riley and Howard 1891b: 253 (biology). Harvey 1893: 99 (host). - Weed 1896: 31 (biology). - Bush 1966a: 457 (lectotype designation). Rhagoletis pomonella: Loew 1873: 267, 329 (taxonomy). - Osten Sacken 1880: 53 (host). - F. H. Snow 1894: 164 (taxonomy). - Doane 1898: 69, 71 (in key; taxonomy). — Doane 1899: 181 (distribution). — Johnson 1900b: 687 (N.J.). — Patch and Munson 1904: 169 (review). - Aldrich 1905: 606 (in catalog). Washburn 1905: 118 (Minn.). - Tucker 1907: 104 (Colo.). - Aldrich 1909: 69 (in key). — Bezzi 1910: 18 (review).-Johnson (part) 1910: 802 (N.J.). - O’Kane 1910: 169 (biology). - Banks 1912: 32 (description of larva). - Illingworth (part) 1912a: 125 (review, biology). - Ross 1913: 67 (biology). - O’Kane (part) 1914: 1 (biolo¬

gy), _ Good 1915: 54 (biology). - Winn and Beaulieu 1915: 153 (Que.). - Woods (part) 1915: 293 (parasite). - Good 1916: 168 (parasite). - Brittain and Good 1917: 1 (review, N.S.). - Caesar 1917: 8 (description, biology). - Brittain 1918: 37 (biology). - Brittain 1919: 15 (biology). - Caesar and Ross 1919: 1 (review). -

356

Systematic Treatment of the Genera

Rhagoletis pomonella Rbagoletis pomonella (state record only)

Map 73. Distribution of Rhagoletis pomonella.

Britton 1920: 204 (Conn.). - Herrick (part) 1920: 89 (biology). - Wellhouse 1920: 388 (host). — Porter and Alden 1921: 62 (parasite). — Knight 1922: 488 (review, damage). - Porter 1922: 165 (parasite). - Wellhouse 1922: 1085 (biology, host). - Champlain and Knull 1923: 215 (biology). - V. T. Phillips (part) 1923: 136 (review). - Snodgrass 1923: 260 (morphology of larva). - Brues (part) 1924: 129 (races). - Curran 1924a: 63 (description of adult). - Curran 1924b: 56, 57 (description of adult). - Snodgrass 1924: 1 (morphology of larva). - Guyton and Knull (part) 1925: 3 (review, Pa.). - Johnson 1925b: 261 (N. Engl.). - Johannsen 1926: 159 (N.Y.). - Pettit 1926a: 1 (review). - Pettit 1926b: 1 (review). - Ries 1927: 144 (biology). - Leonard 1928: 852 (N.Y.). - Porter (part) 1928: 1 (review). - Caesar 1929: 93 (biology). - Cresson (part) 1929: 409 (review). - C. T. Greene 1929: 494 (description of larva, pupa). - Pettit 1929: 1 (review). - Allen 1931:

357

Rh ago let is 338 (bacterial associates). — Chapman 1931: 686 (biology). — Fluke and Allen 1931: 77 (biology). — Gambrell (part) 1931: 226 (biology). — Garman 1931: 59 (host). — Marshall 1931: 82 (lures). — Pettit and Hutson 1931: 44 (review, Mich.). - Allen and Ricker 1932: 557 (bacterial associates). — Curran 1932a: 6, 7 (review). - Dean 1932: 210 (morphology of alimentary canal). - Petch and Maltaise 1932: 56 (Que.). - Phipps and Dirks 1932: 576 (biology). - Allen and Fluke 1933: 1108 (biology). — Chapman 1933: 3 (biology). — Dean 1933: 1 (morphology of alimen¬ tary canal). — Farleman 1933: 825 (biology). — Marshall 1933: 47 (attractants). — Phipps and Dirks 1933a: 344 (dispersal). — Phipps and Dirks 1933b: 349 (biolo¬ gy). — Allen et al. 1934: 228 (bacterial associates). — Bates 1934b: 1 (morphology of larva). — F. H. Benjamin (part) 1934: 14 (review, Fla.). — Bourne et al. 1934: 352 (dispersal). — Chapman and Hammer 1934: 1 (review, hosts). — Dean 1935a: 1 (female reproductive system). — Dean 1935b: 504 (biology). — Dirks 1935: 198 (biology). — McAlister and Anderson (part) 1935: 675 (biology; crossing). — Butt 1937: 487 (morphology of larva). — Hall 1937: 46 (biology). — Pickett (part) 1937: 53 (male genitalia, crossing). — Brimley 1938: 383 (N. Car.). — Dean 1938: 241 (adult nutrition). - Hall (part) 1938: 53 (biology). - Pickett and Neary (part) 1940: 551 (crossing). - E. Cameron 1941: 14, 15 (parasites). - Chapman and Hess 1941: 1 (biology). — Dean 1941: 123 (lures). — Garman 1941: 17 (biology).— Middlekauff 1941: 621 (biology; parasites). - Dean 1942a: 397 (morphology of female reproductive system). — Dean 1942b: 291 (review). — Hall (part) 1943: 202 (crossing). - Hodson 1943: 545 (lures). - Lathrop and Dirks 1944: 199 (biology). - Lathrop and Dirks 1945: 330 (emergence). - V. T. Phillips 1946: 71, 120 (description, biology larva; hosts). — Hodson 1948: 61 (lures). — R. G. Palmer 1948: 993 (emergence). - Garman and Townsend 1952: 3 (review; parasites). Hennig 1952: 218 (catalog of immature stages). - Baulne 1953: 195 (para¬ site). - Peterson 1953: 333, 335, 337 (figures of larva). - Strickland 1953: 277 (morphology). - Commonw. Inst. Entomol. (part) 1954: Map No. 48 (distribution). - Lathrop 1955: 7 (review). - Muesebeck 1956: 99, 102 (parasites). - Christenson and Foote 1960: 171 (biology). - Glass 1960: 1 (description; emer¬ gence). - Neilson 1960: 464 (lures). - Still 1960: 967 (trap). - Oatman 1961: 70 (review, Wis.). - Neilson 1962: 924 (taxonomy). - Stevenson 1963: 1154 (biolo¬ gy).

_ Wasbauer 1963: 1 (taxonomy). - Fischer 1964: 291 (parasites). - Leroux

and Mukerji 1964: 60 (biology; Que.). - Neilson 1964: 810 (biology). - Neilson and McAllen 1964a: 333 (artificial rearing). - Neilson and McAllen 1964b: 904 (artificial rearing). — Oatman 1964a: 349 (biology). — Oatman 1964b: 529 (attrac¬ tants). - Oatman 1964c: 676 (biology). - Oatman et al. 1964: 978 (Wis.). Wasbauer 1964: 5 (review). - Foote 1965a: 674 (in catalog). - Howitt and Connor 1965: 134 (lures). - Maxwell 1965: 110 (traps). - McAllen and Neilson 1965: 168 (biology).-Neilson 1965: 1056 (rearing). - Neilson and McAllen 1965a: 542 (rearing). - Neilson and McAllen 1965b: 276 (biology). - Bush 1966a: 457 (revi¬ sion). - Neilson and Wood 1966: 997 (biology). - Prokopy 1966: 231 (biology). Wilde and Goble 1966: 145 (trapping). - Baerwald and Boush 1967: 682 (biolo¬ gy). - Boush and Baerwald 1967: 865 (parasite). - Boush and Matsumura 1967: 918 (bacterial associates). - Neilson 1967: 217 (biology). - Prokopy 1967a: 4

358

Systematic Treatment of the Genera

(lures). - Prokopy 1967b: 950 (lures). - Prokopy 1967c: 1161 (diet). - Rivard 1967: 895 (parasites). - Maxwell 1968: 103 (biology). - Maxwell and Parsons 1968: 1157 (biology). - Miyazaki et al. 1968: 513 (bacterial associates). — Prokopy 1968a: 403 (biology). - Prokopy 1968b: 1082 (lures). - Prokopy 1968c: 318 (diapause). - Rivard 1968: 5 (biology; bibliography). - Boulanger et al. 1969: 748 (rearing). - Boush et al. 1969: 19 (diet). - Jaques et al. 1969: 850 (disease). Maxwell 1969: 945 (traps). - Maxwell and Parsons 1969: 1310 (biology). Moore 1969: 1076 (lures). - Neilson 1969: 1028 (rearing). - Simon 1969: 169 (serology). - Kring 1970: 466 (lures). - Lienk 1970: 1684 (host). - Roth and Richardson 1970: 496 (hosts). - Shervis et al. 1970: 294 (host). - Monteith 1971a: 52 (predators). - Monteith 1971b: 507 (parasites). - Monteith 1971c: 38 (review). - Neilson 1971: 648 (biology). - Prokopy et al. 1971: 1405 (behavior).Stanton 1971: 61 (biology). - Bateman 1972: 494 (biology). - Biggs 1972: 349 (behavior). - Kamasaki et al. 1972: 1292 (rearing). - Monteith 1972: 257 (preda¬ tors). - Prokopy 1972a: 326 (biology). - Prokopy 1972b: 720 (lures). - Prokopy and Bush 1972a: 275 (behavior). - Prokopy and Bush 1972b: 597 (host). Prokopy et al. 1972: 97 (behavior). - Rivard 1972: 62 (biology). - Stanton 1972: 254 (rearing). — Wasbauer 1972: 131 (hosts). — Weems 1972b: 1 (review). — Carter 1973: 23 (associates). - Dean and Chapman 1973: 1 (biology). - Knell and Stoffolano 1973: 1221 (histology, larva). - Neilson 1973: 555 (rearing). - Prokopy 1973: 953 (lures). - Prokopy and Bush 1973a: 873 (biology). - Prokopy and Bush 1973b: 927 (biology). - Prokopy and Bush 1973c: 1197 (biology). - Prokopy et al. 1973: 743 (lures). - Rivard 1973: 126 (Que.). - Buriff and Davis 1974: 572 (trapping). - Cameron and Morrison 1974a: 1025 (sampling). - Cameron and Morrison 1974b: 13 (parasite). - Pree 1974: 1019 (development). - Reissig 1974a: 484 (traps, lures). - Reissig 1974b: 733 (biology). - Rock and Yeargan 1974: 508 (biology). - Moericke et al. 1975: 497 (biology). - Neilson 1975: 909 (biology). - Prokopy 1975a: 197 (lures). - Prokopy 1975b: 905 (behavior). Prokopy 1975d: 420 (trap). - Reissig 1975a: 445 (trapping). - Reissig 1975b: 534 (trapping). - Trottier et al. 1975: 211 (traps). - Monteith 1976: 28 (predators). H. S. Myers et al. 1976: 39 (biology). - Neilson 1976: 885 (review, N.S.). Neilson et al. 1976: 527 (traps). - Prokopy 1976b: 6 (traps). - Prokopy et al. 1976: 170 (pheromones). - Reissig 1976: 634 (traps, lures). - Agee et al. 1977: 359 (eye structure). - Cameron and Morrison 1977: 769 (biology). - Monteith 1977. 23 (predators). — Prokopy 1977a: 593 (lures). — Prokopy and Spatcher 1977: 960 (receptors). - Reissig 1977: 631 (lures). - Monteith 1978: 3 (parasites). - Poinar et al. 1978: 19 (microorganisms). - Prokopy and Owens 1978: 409 (vision). - Prokopy and Webster 1978: 481 (pheromones; parasites). - Reissig and Smith 1978: 155 (biology; host). - Sharp 1978: 199 (behavior; flight). - Webster and Stoffolano 1978: 844 (reproductive system). - Wharton and Marsh 1978: 149 (parasites). - Berg 1979: 19 (in key to larvae). - Crnjar et al. 1979: 283 (phe¬ romone). - Prokopy and Hauschild 1979: 696 (lures). - Sharp 1979: 70 (wing dimensions). - Webster et al. 1979: 41 (reproduction). - Averill and Prokopy 1980: 34 (pheromone). - Berlocher 1980a: 131 (electrophoretic key). - Berlocher 1980b: 63 (genetics). - Muesebeck 1980: 32, 44 (parasites). - Prokopy and

359

Rhagoletis Berlocher 1980: 1319 (host). - Smith and Prokopy 1980: 585 (mating behavior). AliNiazee and Penrose 1981: 245 (review, Oreg.). - Nation 1981: 121 (sex phe¬ romone glands). - Prokopy 1981a: 477 (pheromone system). - Prokopy 1981c: 181 (epideictic pheromones). - Roitberg and Prokopy 1981: 540 (pheromones). Crnjar and Prokopy 1982: 393 (pheromone receptors). - Prokopy 1982: 30 (be¬ havior). - Prokopy et al. 1982a: 1 (oviposition). - Prokopy et al. 1982b: 76 (oviposition). - Ratner and Stoffolano 1982: 555 (oesophageal bulb). - Roitberg et al. 1982: 307 (foraging behavior). - Roitberg and Prokopy 1982: 437 (foraging behavior). — Westcott 1982: 25 (taxonomy). — Berlocher and Smith 1983: 337 (genetics). — Joos et al. 1984: 9 (review, Calif.). — Ratner and Stoffolano 1984: 191 (morphology). — Stoffolano and Yin 1987: 41 (morphology). Trypeta (Rhagoletis) pomonella: Loew 1873: 337 (taxonomy). — Osten Sacken 1878: 191 (in catalog). — Quaintance 1908: 1 (review). recognition. R. pomonella, together with zepbyria, mendax, and cornivora,

are among the most readily recognized species of Rhagoletis by virtue of their wing pattern (figs. 374—377), which consists of a slightly oblique discal band to which the anterior and posterior apical bands are connected, forming a characteristic F-shaped pattern in the apical half of the wing. The absence of the subapical band distin¬ guishes the species of the pomonella group from all other species of Rhagoletis. R. striatella, which also has an F-shaped apical wing pattern, is distinguished from pomonella by the color pattern of the scutellum and the additional characters given in the key to species. R. pomonella is separable from the other three species of the pomonella group by the presence in most specimens of heavy black shading on the posterior surface of the fore femur (fig. 387, a), and, in specimens from the northern part of its range, by a generally larger body size and by the longer aculeus (0.901.49 mm). For means of distinguishing pomonella from zepbyria, see the recognition section for zepbyria. In the southern part of its range, specimens of pomonella generally are smaller than in the north. For that reason and because of a conse¬ quently shorter aculeus, females are not separable from those of mendax and cor¬ nivora by the use of morphological characters. Mexican specimens of pomonella resemble those that occur in the United States and Canada but generally are larger and possess a light spot near the base of the apical wing band. distribution. Map 73. Also known from northern and central Mexico, larvae

and adults from Crataegus originating in localities at fairly high altitudes are com¬ monly intercepted by the USDA Animal and Plant Health Inspection Service. One doubtful record indicates the possible occurrence of pomonella in northern Colom¬ bia. hosts. The host relationships of pomonella have been widely investigated and

prove to be quite complex. Bush (1966a) summarized this situation well; we are not aware of a later comprehensive analysis. Wasbauer (1972) listed pomonella as having been found in at least 27 different plant species belonging to at least nine different genera, all of which belong in the Pomoideae and Prunoideae of the family Rosaceae.

360

Systematic Treatment of the Genera These genera are Amelanchier, Aronia, Cotoneaster, Crateagus, Malus, Prunus, Pyracantha, Pyrus, and Sorbus. Symphoricarpos, a genus not mentioned by Bush (1966a), was reported as a forced laboratory host by Pickett and Neary (1940).

R. pomonella is undoubtedly the native tephritid of greatest eco¬ nomic importance in the United States and extreme southern Canada, and an im¬ discussion.

mense body of literature applies to many aspects of its life history and economic importance. This species was found in Oregon in 1979, a situation reviewed by Joos et al. (1984). Subsequently, it has spread to southern Washington, northern Califor¬ nia, and Utah.

Rhagoletis ribicola Doane American currant fruit fly, dark currant fly (Figs. 382, 390, 392; Map 64) Rhagoletis ribicola Doane 1898: 69 (female lectotype, WSU; Wash.). - Piper and Doane 1898: 1 (description, biology). - Doane 1899: 181 (distribution). - Aldrich 1905: 607 (in catalog). - Aldrich 1909: 69 (in key). - Bezzi 1910: 15 (review). Grossbeck 1912: 378 (note, cotype). - Marcovitch 1915: 173 (taxonomy). Severin 1917b: 258 (distribution, Calif.). - Cresson 1929: 406, 407 (synonymy, host). — Curran 1932a: 6 (in key). — Pickett 1937: 53 (taxonomy, male genitalia). — Essig 1938: 605 (review). - Hatch 1938: 197 (bibliography, Wash.). - V. T. Phillips 1946: 72, 120 (description, biology larva; hosts). - Forsell 1947: 167 (host, Puget Sound). - Hennig 1952: 218 (catalog immature stages). - Essig 1958: 605 (re¬ view). - Foote and Blanc 1963: 61 (review, Calif.). - Foote 1965a: 674 (in catalog). - Bush 1966a: 505 (revision, lectotype designation). - Foote 1966b: 124 (type data). - Madsen 1970: 13 (biology). - Wasbauer 1972: 132 (hosts). - Zack 1984: 32 (type data).

The single apical band is entire, fully joined to the subapical band, and is separated from the costa by a hyaline area (fig. 382, b). In these respects the recognition.

wing resembles those of ebbettsi, electromorpba, juniperina, persimilis, and tabellaria. In addition to host preference, the following differences exist: from tabellaria, persimilis, and electromorpba, ribicola may be distinguished by the absence of a dark posterior connection between the subbasal and discal bands (fig. 382); from ebbettsi by the lack of a connection between the discal and subapical bands; and from juniperina by the yellow (or only partially shaded) fore femur. R. ribicola also resembles berberis in wing pattern, but in the latter species, the apical band touches the costa throughout its entire length (fig. 380, c). All leg segments except the coxae are yellow in ribicola males, whereas in females the femora are brownish black with yellowish apices. The posterior region of the head exhibits a darkened horseshoe¬ shaped area that extends nearly to the oral margin (fig. 392).

Rhagoletis distribution.

Map 64. R. ribicola is confined to the western United States and

Canada. The host relationships of ribicola are poorly known and remain uninvesti¬ gated. The species is restricted to Ribes, of which three species have been recorded as hosts. hosts.

In North America, the ribicola group includes only ribicola and berberis. The group was defined by Bush (1966a) only on the basis of genitalic characters but is divided in the abbreviated key, which is based solely on wing pattern. discussion.

Rhagoletis striatella Wulp (Figs. 369, 384; Map 66) Rhagoletis striatella Wulp 1899—1900: 408 (female holotype, BMNH; Guerrero, Mexico). — V. T. Phillips 1923: 136 (review). - Curran 1928b: 100 (Ont.). — Cresson 1929: 405 (review). — Curran 1932a: 6 (in key). - Aczel 1954a: 78 (in key). — Foote 1965a: 674 (in catalog). — Foote 1965b: 239 (type data, taxonomy). — Bush 1966a: 514 (revision). — Wasbauer 1972: 132 (host). — Berg 1979: 19 (in key, larva). — Foote 1981: 34 (in key, taxonomy, hosts). - Norrbom 1983: 11 (host). R. striatella is one of the most distinctive species of Rhagoletis in North America. As indicated in the abbreviated key, the wing pattern (fig. 369) resembles those of species of the pomonella group in that the anterior and posterior apical bands form a distinctive F-shaped pattern in the apical half of the wing. However, striatella differs from all of the species of the pomonella group by the presence of a subapical wing band, the lack of a posterior connection between the subbasal and discal bands, the extremely long aculeus and oviscape, the shape of the yellow to whitish scutellar spot (fig. 384, a), the very wide gena in comparison recognition.

with the eye height, and its preference for solanaceous hosts. DISTRIBUTION.

Map 66. R. striatella also occurs at relatively high altitudes in

Mexico, but it has not been recorded south of that country. R. striatella is apparently of considerable economic importance in Mex¬ ico, where it infests commercial plantings of husk tomatoes (Physalis sp.) used for the preparation of sauces (Foote 1981). A Physalis species also is a host in eastern North hosts.

America (Norrbom 1982). Bush (1966a) indicates that striatella may have originated in Mex¬ ico. The species group to which it belongs is otherwise entirely Neotropical (Foote discussion.

1981).

362

Systematic Treatment of the Genera Rhagoletis suavis (Loew) Northern walnut husk fly, walnut husk maggot (Fig. 368; Map 67) Trypeta suavis Loew 1862c: 75 (male holotype, MCZ; “Middle States”). - Loew 1873: 236 (review). Acidia suavis: Loew 1873: 235 (taxonomy). — Coquillett 1899c: 260 (taxonomy). — Aldrich 1905: 603 (in catalog). - Washburn 1905: 118 (Minn.). - Britton 1920: 203 (Conn.). Trypeta (Acidia) suavis: Osten Sacken 1878: 189 (in catalog). Rhagoletis suavis: Coquillett 1899c: 260 (taxonomy). - Babb 1902: 242 (descrip¬ tion larva, pupa). - Washburn 1906: 80 (Minn.). - Aldrich 1909: 69 (in key). Aldrich 1910: 99 (biology). - Banks 1912: 32 (description larva). - Cresson 1920: 66 (taxonomy). - Brooks 1921: 1 (biology). - Bridwell 1923: 262 (biology). V. T. Phillips 1923: 136 (review). - Johnson 1925b: 262 (N. Engl.). - Leonard 1928: 852 (N.Y.). - Cresson 1929: 411 (review). - Gambrell 1931: 226 (biology). - Barrett 1932: 296 (hosts). - Beck 1932: 497 (biology). - Curran 1932a: 6 (in key). - Bates 1934b: 1 (morphology larva). - Boyce 1934: 375 (review). - Ries 1935: 717 (biology). - Stirrett 1936: 69 (Ont.). - Pickett 1937: 53 (taxonomy, male genitalia). - Brimley 1938: 383 (N. Car.). - M. E. Phillips 1939: 325 (larval morphology). - V. T. Phillips 1946: 74, 120 (description, biology larva; hosts). Benjamin and Hodson 1949: 707 (lure, distribution). - Hennig 1952: 218 (catalog immature stages). - Peterson 1953: 331, 333, 335, 337 (figures of larvae). Christenson and Foote 1960: 171 (biology). - Wasbauer 1964: 5 (review). - Foote 1965a: 675 (in catalog). - Bush 1966a: 483 (revision). - Dean 1969: 940 (host). Wasbauer 1972: 132 (hosts). - Steyskal 1973: 522 (in key, larvae in walnut husks). - Wharton and Marsh 1978: 149 (parasites). - Berlocher 1980a: 131 (elec¬ trophoretic key). - Muesebeck 1980: 32, 44 (parasites). All four transverse wing bands are present in suavis, but the wing pattern (fig. 368) is distinctive in having a broad connection between the discal and recognition.

subapical bands at the approximate center of the wing, together with an uninter¬ rupted hyaline incursion from the posterior margin basad of the subapical band (fig. 368, b). In this respect, the wing pattern is similar to that of fausta (fig. 367), but the hyaline incursion in the latter species is interrupted by a black area to form a rounded hyaline spot near the apex of cell dm (fig. 367, b). In addition, only the anterior apical band is present in suavis; in fausta, both anterior and posterior apical bands are present. The thorax and abdomen of suavis are yellow with brownish to dark brown markings, and the scutellum is entirely yellow. Map 67. Although occurring close to the Canadian border, suavis appears to be restricted to the United States. distribution.

Primarily restricted to Juglans, suavis has been reared from four species of that plant genus. Wasbauer (1972) listed peach as a recorded host. hosts.

363

Rhagoletis

This is the most commonly encountered and widespread species of Rhagoletis occurring in walnut and butternut in the eastern half of the United States. discussion.

Rhagoletis tabellaria (Fitch) (Fig. 359; Map 72) Tephritis tabellaria Fitch 1855a: 770 (types probably lost [Bush 1966a: 495]; N.Y.) — Fitch 1856: 66 (repeats Fitch 1855a). — Aldrich 1905: 612 (in catalog). Trypeta tabellaria: Loew 1862c: 60 (taxonomy). - Loew 1873: 263 (review). — Osten Sacken 1878: 80 (in catalog). Rhagoletis tabellaria: Loew 1873: 265, 329 (taxonomy). — Doane 1898: 69 (in key). -Johnson 1900b: 687 (N.J.). - Aldrich 1905: 607 (in catalog). - Aldrich 1909: 69 (in key). — Bezzi 1910: 18 (review, world). — Johnson 1910: 801 (N.J.). — V. T. Phillips 1923: 135 (review). — Plank 1923: 99 (host). — Johnson 1925b: 261 (N. Engl.). — Leonard 1928: 852 (N.Y.). - Cresson 1929: 401 (N. Engl.). - Glasgow 1931: 437 (host). - Pickett 1937: 53 (taxonomy, male genitalia). - Strickland 1938: 204 (Alta.). — Ouellet 1941: 130 (Que.). — V. T. Phillips 1946: 120 (host).— Forsell 1947: 167 (host, Puget Sound). — Frick et al. 1954: 59 (trapping). - Foxlee 1957: 37 (B.C.). - Steyskal 1957: 94 (description eye color). - Foote and Blanc 1963: 61 (review, Calif.). — Fischer 1964: 305, 358 (parasites). — Foote 1965a: 675 (in catalog). — Bush 1966a: 495 (revision). - Madsen 1970: 13 (biology). Wasbauer 1972: 133 (hosts). - Wharton and Marsh 1978: 149 (parasites). Berlocher 1984a: 237 (taxonomy, hosts). Trypeta (Rhagoletis) tabellaria: Loew 1873: 338 (taxonomy). - Osten Sacken 1878: 191 (in catalog). Rhagoletis tabernella [error]: Cresson 1929: 402 (in catalog). recognition.

The wing patterns of tabellaria, persimilis, and electromorpha

(figs. 359, 378, and 379, respectively) are similar in having the following combina¬ tion of characters: subbasal, discal, and subapical bands present, the subbasal and discal bands distinctly connected posteriorly, and the single (anterior) apical band undivided apically and separated from the costa by a hyaline area. The subbasal and discal bands are united, and the apical band is separated from the costa in the pomonella group, but the subapical band is absent in those species (figs. 374-377). R. tabellaria is distinguished from persimilis mainly on the basis of male and female genitalic characters. The aculeus of persimilis is slightly longer than that of tabellaria (about 0.8 mm versus about 0.7 mm, respectively). For the characters that distinguish tabellaria from electromorpha, see the recognition section for the latter species. distribution.

Map 72.

Bush (1966a) discussed the host relationships of the eastern and western populations of tabellaria in some detail. The eastern populations attack only Cornus HOSTS.

spp., whereas those in the west are restricted to Vaccinium.

364

Systematic Treatment of the Genera

According to Bush (1966a), two distinct allopatric and alltrophic populations exist. The differences between these two populations have not been investigated. DISCUSSION.

Rhagoletis zephyria Snow Snowberry maggot (Figs. 374, 395; Map 70) Rhagoletis zephyria Snow 1894: 164 (male lectotype, UKL; So. Calif.). - Doane 1898: 69 (in key, synonymy). - Aldrich 1909: 69 (synonymy). - Bezzi 1910: 18 (synonymy). - Curran 1924a: 63 (description). - Curran 1924b: 56 (description). - Porter 1928: 4 (taxonomy, hosts). - Cresson 1929: 409 (synonymy, taxonomy). - Curran 1932a: 6 (in key). - F. H. Benjamin 1934: 16 (review, Fla.). - Pickett 1937: 53 (taxonomy, male genitalia). - V. T. Phillips 1946: 120 (hosts). - Hennig 1952: 218 (catalog immature stages). - Christenson and Foote 1960: 171 (biolo¬ gy). - Byers et al. 1962: 180 (type data). - Foote 1962: 178 (type designation, synonymy). - Foote and Blanc 1963: 62 (review, Calif.). - Wasbauer 1963: 1 (taxonomy). - Wasbauer 1964: 5 (taxonomy). - Foote 1965a: 675 (in catalog). Bush 1966a: 464 (revision). - Madsen 1970: 13 (biology). - Wasbauer 1972: 133 (hosts). - Wharton and Marsh 1978: 149 (parasite). - Arnaud and Owen 1981: 151 (type data). - Averill and Prokopy 1982: 315 (pheromone). - Wescott 1982: 25 (taxonomy). Rhagoletis zephrina [error]: Woodworth 1913: 137 (Calif.). Rhagoletis pomonella [misidentification]: Treherne 1917: 329 (review, B.C.). Downes 1919: 2 (taxonomy, hosts). - Gahan 1919: 164 (parasite). - Cole and Lovett 1921: 325 (Oreg.). - Essig 1938: 604 (review). - Strickland 1946: 166 (Alta.). - Forsell 1947: 167 (Puget Sound). - Essig 1958: 604 (review). Rhagoletis symphoricarpi Curran 1924a: 63 (male holotype, CNC; Victoria, B.C.). - Curran 1924b: 56 (description). - Porter 1928: 4, 7 (taxonomy, hosts). Cresson (part) 1929: 409 (synonymy, taxonomy). - Gahan 1930: 2 (parasite). Curran 1932a: 3, 6 (in key). - V. T. Phillips 1946: 76, 120 (description, biology larva; hosts). - Flennig 1952: 218 (catalog immature stages). — Frick et al. 1954: 59 (trapping). — Foxlee 1957: 37 (B.C.). — Byers et al. 1962: 181 (type data). — Foote 1962: 178 (type data). - Foote 1965a: 675 (in catalog). RECOGNITION. The wing pattern of zephyria closely resembles those of pomo¬ nella, mendax, and cornivora (figs. 375-377); the discal band slants diagonally across the wing and is broadly connected to the anterior and posterior apical bands, which are relatively long and widely separated at their apices, forming an F-shaped pattern in the distal half of the wing. R. striatella has a somewhat similar wing pattern but may be readily distinguished from zephyria by the presence of a subapical band and by the additional characters presented in the key. With the advent of pomonella into the northwestern United States, a means was required to distinguish that species from zephyria with the greatest possible degree of accuracy. The differ-

365

Rbagoletotrypeta ences between these two species were summarized by Wasbauer (1963), Bush (1966a), and Westcott (1982) as follows: R. pomonella: (a) Aculeus length 0.90 to 1.49 mm (very little or no overlap with zephyria). (b) Wing of female very slightly less broadly rounded distally and with a subtly lighter color pattern. (c) Wing band width ratio (posterior apical/discal) usually less than 0.50. (d) Surstyli of male (fig. 394) parallel in posterior view. (e) Larvae found in Rosaceae but not in Caprifoliaceae. R. zephyria: (a) Aculeus length 0.66—0.88 mm (very little or no overlap with pomonella). (b) Wing of female very slightly more broadly rounded apically and with subtly darker color pattern. (c) Wing band width ratio (posterior apical/discal) usually greater than 0.50. (d) Surstyli of male divergent (fig. 395) in posterior view. (e) Larvae found only in Symphoricarpos spp. (Caprifoliaceae). distribution.

Map 70.

R. zephyria is restricted to the plant genus Symphoricarpos. Although only two species have definitely been recorded as hosts, others also may be involved hosts.

(Bush 1966a).

Genus Rhagoletotrypeta Aczel Rbagoletotrypeta Aczel 1954b: 137 (type species, Rhagoletotrypeta xanthogastra Aczel 1951: 315, by original designation). - Foote 1965a: 675 (in catalog). - Foote 1966a: 803 (review). - Sabrosky 1971: 85, 87 (correction, catalog). - Wasbauer 1972: 133 (hosts). - Steyskal 1981: 707 (partial revision). The two U.S. species discussed herein resemble many species of Rhagoletis in having a predominantly black thorax and abdomen with whitish markings and a hyaline wing with dark to black transverse bands. The following features distinguish the genus from Rhagoletis: first flagellomere rounded apically; a prominent facial carina that widens perceptably at the anterior oral margin; scutum with a prominent median white stripe widening posteriorly; and the oviscape almost recognition.

bulbous basally but narrowing abruptly apically. discussion.

Rhagoletotrypeta is a little-known genus with three species groups,

two of which are restricted to continental South America. The third group is North American and contains a Mexican species, annulata Aczel, in addition to the two

366

Systematic Treatment of the Genera

discussed herein. The aculeus is distinctive in these three species, with the tip con¬ stituting more than half its length. All of the species whose biology is known breed in the fruit of Celtis species, although one South American species also has been re¬ ported to attack plum. Rhagoletotrypeta belongs to the subtribe Carpomyina, a mainly Holarctic and Neotropical group of fruit-breeding species. Rkagoletis and Zonosemata are the only other two genera of this subtribe in our region.

Key to Species of Rhagoletotrypeta North of Mexico Dark bands of wing relatively broad; apical band filling all of cells r1 and r2+3 distal to subapical band (fig. 396, a); scutum with the 4 dark longi¬ tudinal stripes separating the gray tomentose areas narrow (fig. 398, a)

.rohweri Foote Dark bands of wing relatively narrow; apical band filling no more than half of cell r2+3 distal to subapical band (fig. 397, a); scutum with the 4 dark stripes separating the gray tomentose areas broad (fig. 399, a).

.uniformis Steyskal Rhagoletotrypeta rohweri Foote (Figs. 396, 398; Map 74) Rhagoletotrypeta sp.: Foote 1965a: 675 (in catalog). Rhagoletotrypeta rohweri Foote 1966a: 804 (female holotype, NMNH; Camden, N.J.). - Wasbauer 1972: 133 (host). - Steyskal 1981: 708 (taxonomy, in key)!

Figures 396, 397. Right wings, Rhagoletotrypeta spp. 396, rohweri Foote; 397, uniformis Steyskal. Steyskal

3"’ SCUtal pattem’ RhaSoletotrypeta spp. 398, rohweri Foote; 399, uniformis

367

Rhagoletotrypeta The wing of rohweri is characterized by a broad, dark, apical band that completely fills cells rl and r2+3 apicad of its junction with the subapical band (fig. 396, a) and a slightly tapering discal band (fig. 396, b). These bands are considerably narrower in uniformis (compare figs. 396 and 397). The scutal mark¬ ings consist of four narrow, dark, parallel lines (as shown in fig. 398, a) rather than the two broader ones as shown for uniformis in fig. 399, a. The aculeus has a strong subbasal constriction and its tip has a slight preapical dorsal bend. recognition.

distribution.

host.

Map 74.

R. rohweri has never actually been reared from hackberry (Celtis sp.),

therefore the report by Wasbauer (1972) of this plant as a host is circumstantial although probably correct. Small larvae resembling those of Rhagoletis spp. have been found in hackberries at the type locality of this species where adults were trapped, but the larva of rohweri has never been positively identified. discussion.

R. rohweri is a rare species that has not been encountered since

1962.

Rhagoletotrypeta uniformis Steyskal (Figs. 397, 399; Map 74) Rhagoletotrypeta annulata Aczel [misidentification]: Foote 1966a: 806 (taxonomy). — Wasbauer 1972: 133 (host). Rhagoletotrypeta uniformis Steyskal 1981: 709 (male holotype, NMNH; San An¬ tonio, Tex.). RECOGNITION.

R. uniformis is distinguished from rohweri by the narrower wing

bands; the apical one at most fills the apical half of the part of cell r2+3 posterior to the anterior apical band (fig. 397, a), and the discal band distinctly narrows as it crosses vein CuA1 (fig. 397, b). It is further distinguished by the presence of two very broad, dark scutal stripes on each side of the central light stripe, each separated by a narrow stripe of light gray tomentum (fig. 399). The aculeus, like that of rohweri, has a strong subbasal constriction, but its tip is straight or slightly ventrally curved. The aculeus of annulata is much different in lacking a subbasal constriction and having the tip flattened in the sagittal plane and being trilobed apically. distribution.

HOST.

Map 74.

Celtis laevigata. R. uniformis emerged in the spring from hackberries col¬

lected in San Antonio, Texas, the previous fall. discussion.

Steyskal (1981) describes the genitalia and the larva of this rare

species. Foote (1966a) misidentified this species as annulata Aczel, a closely related species now known only from Mexico.

368

Systematic Treatment of the Genera

• Rkagoletotrypeta rohweri o Rhagoletotrypeta uniformis ▲ Rhynencina longirostns A Stenopa affinis

Map 74. Distribution of Rhagoletotrypeta rohweri, R. uniformis, Rhynencina longirostris, and

Stenopa affinis.

Genus Rhynencina Johnson Rhynencina Johnson 1922: 24 (type species, Rhynencina longirostris Johnson 1922: 24, by original designation). — Shewed 1961: 1045 (taxonomy). — Foote 1965a: 659 (in catalog). - Steyskal 1979: 5 (in key). Aleomyia Phillips 1923: 123 (type species, Aleomyia alpha Phillips 1923: 123 by original designation) (= longirostris Johnson). - Curran 1932a: 2 (in key). Curran 1932b: 3 (in key); 12 (review). Euribia (part): Curran 1934: 287 (in key). - Hering 1941a: 128 (in key). Rhynencina is a monotypic genus with unique head characters that cannot be confused with those of any other U.S. or Canadian genus. The face recognition.

from lateral view is slanted forward at an angle to the vertical axis of the head, and the anterior oral margin is produced anteriorly well beyond the anterior margin of the parafacial (fig. 19, a). The proboscis is geniculate, and the labellum is longer than the head (fig. 19, c). The palpi (fig. 19, b) are comparatively large and broad. As in Urophora, cell cup is closed by a transverse or outwardly convex vein CuA^ without any sign of a distal elongation at its apex, and only one pair of orbital bristles is present. Also, the wing bands (fig. 400) are narrow and broken as in many species of Urophora, but the shape of the head with its elongated face and the extremely long labellum distinguish Rhynencina from species of Urophora occurring north of Mexico.

369

Stenopa

Figure 400. Right wing, Rhynencina longirostris Johns.

Steyskal (1979) presented a key to the known world genera of Myopitini, the tribe containing all those genera in which cell cup is closed transversely or convexly at its apex and only one pair of orbital bristles is present (White and Clement 1987). Of those genera, only Rhynencina and Urophora occur in the United States and Canada. See the discussion of Urophora for additional comments on the discussion.

status of these two distinctive genera.

Rhynencina longirostris Johnson (Figs. 19, 400; Map 74) Rhynencina longirostris Johnson 1922: 24 (male and female syntypes, MCZ, PDAH; Mt. Alto, Pa.). - Foote 1965a: 659 (in catalog). Aleomyia alpha Phillips 1923: 124 (syntypes male and female, CUI, NMNH; Plummer’s I., Md.). - Curran 1932a: 2 (in key). - Curran 1932b: 12 (in key). Flering 1941a: 128 (in key). - Foote 1965a: 659 (synonymy). Euribia longirostris: Bates 1934a: 295 (taxonomy). - Hering 1941a: 128 (in key). recognition.

distribution.

host.

Not

As

for

the genus.

Map 74.

known.

discussion.

Although no other species have ever been associated with longi¬

rostris in the genus Rhynencina, several Neotropical species described by Steyskal (1979) belong here. A Mesoamerican species appears to be the most closely related to longirostris (Freidberg and Norrbom, in preparation).

Genus Stenopa Loew Stenopa Loew 1873: 234 (type species, Trypeta vulnerata Loew 1873: 232, by original designation); 327 (in key). - Williston 1896: 121 (in key). - Coquillett 1899c: 267 (in key). - Aldrich 1905: 603 (in catalog). - Williston 1908: 286 (in key). - Coquillett 1910: 608 (type data). - Cresson 1914a: 276 (taxonomy). Hendel 1914b: 88 (in key). - V. T. Phillips 1923: 121 (in key); 133 (review). Curran 1932b: 4 (in key). - Curran 1934: 291 (in key). - Foote and Blanc 1963:

369

Stenopa

Figure 400. Right wing, Rhynencina longirostris Johns.

Steyskal (1979) presented a key to the known world genera of Myopitini, the tribe containing all those genera in which cell cup is closed transversely or convexly at its apex and only one pair of orbital bristles is present (White and Clement 1987). Of those genera, only Rhynencina and Urophora occur in the United States and Canada. See the discussion of Urophora for additional comments on the discussion.

status of these two distinctive genera.

Rhynencina longirostris Johnson (Figs. 19, 400; Map 74) Rhynencina longirostris Johnson 1922: 24 (male and female syntypes, MCZ, PDAH; Mt. Alto, Pa.). - Foote 1965a: 659 (in catalog). Aleomyia alpha Phillips 1923: 124 (syntypes male and female, CUI, NMNH; Plummer’s I., Md.). - Curran 1932a: 2 (in key). - Curran 1932b: 12 (in key). Flering 1941a: 128 (in key). - Foote 1965a: 659 (synonymy). Euribia longirostris: Bates 1934a: 295 (taxonomy). - Hering 1941a: 128 (in key). recognition.

distribution.

host.

Not

As

for

the genus.

Map 74.

known.

discussion.

Although no other species have ever been associated with longi¬

rostris in the genus Rhynencina, several Neotropical species described by Steyskal (1979) belong here. A Mesoamerican species appears to be the most closely related to longirostris (Freidberg and Norrbom, in preparation).

Genus Stenopa Loew Stenopa Loew 1873: 234 (type species, Trypeta vulnerata Loew 1873: 232, by original designation); 327 (in key). - Williston 1896: 121 (in key). - Coquillett 1899c: 267 (in key). - Aldrich 1905: 603 (in catalog). - Williston 1908: 286 (in key). - Coquillett 1910: 608 (type data). - Cresson 1914a: 276 (taxonomy). Hendel 1914b: 88 (in key). - V. T. Phillips 1923: 121 (in key); 133 (review). Curran 1932b: 4 (in key). - Curran 1934: 291 (in key). - Foote and Blanc 1963:

370

Systematic Treatment of the Genera

63 (review, Calif.). - Foote 1965a: 671 (in catalog). - Wasbauer 1972: 133 (hosts). - Novak and Foote 1975: 43 (key to species). - Freidberg 1984: 129 (galls). Trypeta (Stenopa): Osten Sacken 1878: 189 (in catalog).

Moderately large flies having a brown to black body with many white, stout setae and with mostly black bristles, yellow legs, and prominently recognition.

banded wings. There are three pairs of frontal bristles, two pairs of orbital bristles, two pairs of black bristles on a subshining, black, inflated scutellum, and the single pair of dorsocentral bristles arise well anterior to the midpoint between the acrostichal and supra-alar bristles. The wing pattern of Stenopa (figs. 401, 402) markedly resembles those of Cecidocbarella and some species of Rhagoletis in having very broad, dark bands. The genus is separable from Rhagoletis in having an apically rounded first flagellomere, stout white postocular and scutal setae, and cells be and c dark. Stenopa differs from Cecidocbarella in having two or three large costal bristles at crossvein h, and in that the bands in the basal and apical halves of the wing are broadly joined in the apex of cell cuat (fig. 401, b).

The species of Stenopa are not commonly encountered, and the genus is of no known economic importance. We have seen a male of an undescribed discussion.

species from Mexico. Stenopa appears to be most closely related to the Neotropical genus Dracontomyia Becker, from which it can be distinguished by the presence of two or three large costal bristles at the humeral crossvein.

Key to Known Species of Stenopa Anal lobe largely dark, with a hyaline spot of variable size within the dark portion (fig. 401, a); outer vertical bristles yellowish to dark brown (fig. 403, a), at least slightly darker than postoculars; posterior margin of scutum and anterior corners of scutellum without clusters of white setulae ..

(Loew)

Anal lobe largely hyaline, dark only along apical border to posterior wing margin (fig. 402, a); outer vertical bristles white (fig. 404, a), concolorous with postoculars; posterolateral margin of scutum and scutellum basolateral to basal bristle with pairs of clusters of white setulae .affinis Quisenberry

Stenopa affinis Quisenberry (Figs. 402, 404; Map 74) Stenopa affinis Quisenberry 1949c: 87 (male holotype, NMNH; Bennett Cr., Colo.). - Byers et al. 1962: 180 (type data). - Foote 1962: 172 (description). - Foote 1965a: 671 (in catalog). - Novak and Foote 1975: 42 (biology, immature stages). — Goeden and Headrick 1990: 641 (biology, immature stages, parasite).

Stenopa

Figures 401, 402. Right wings, Stenopa spp. 401, vulnerata (Lw.).; 402, affinis Quis. Figures 403, 404. Front view of head, Stenopa spp. 403, vulnerata (Lw.).; 404, affinis Quis.

The outer vertical bristles of affinis are distinctly whitish (fig. 404, a) and concolorous with the postoculars, the scutum has several clusters of white setulae, and the area at the base of the anal lobe is extensively hyaline (fig. 402, a). The scutum of affinis has whitish tomentose stripes laterally and posteriorly. RECOGNITION.

distribution.

host.

Map 74.

Senecio multilobatus is the only known host (Goeden and Headrick 1990).

discussion.

Goeden and Headrick (1990) discussed the biology of affinis in

Nevada and decribe and illustrated the immature stages in detail.

Systematic Treatment of the Genera

Stenopa vulnerata (Loew) (Figs. 40, 401, 403; Map 75) Trypeta vulnerata Loew 1873: 232 (male and female syntypes, MCZ; Mass.). Coquillett 1910: 608 (type data). Trypeta (Stenopa) vulnerata: Osten Sacken 1878: 189 (in catalog). Stenopa vulnerata: F. H. Snow 1904: 345 (Ariz.). - Aldrich 1905: 603 (in catalog). Johnson 1909: 113 (distribution). - Hendel 1914b: 88 (type data). - Britton 1920: 203 (Conn.). - V. T. Phillips 1923: 133 (review).-Johnson 1925b: 261 (N. Engl.). — Leonard 1928: 851 (N.Y.). — Curran 1934: 288, 290 (figures of wing, head). — Brimley 1938: 383 (N. Car.). - Strickland 1938: 204 (Alta.). - Quisenberry 1949c: 88 (taxonomy). - Foote 1962: 172 (taxonomy). - Foote and Blanc 1963: 63 (review, Calif.). - Foote 1965a: 671 (in catalog). - Novak et al. 1967: 148

Map 75. Distribution of Stenopa vulnerata.

373

Strauzia (host, gall). - Wasbauer 1972: 133 (host). - Novak and Foote 1975: 43 (biology, immature stages). — Freidberg 1984: 129 (review, mating behavior). The wing patterns of vulnerata and affinis (figs. 401, 402) afford the principal means of distinguishing between them, as most other characters are similar except for the outer vertical bristles, which are dark brown to yellowish and darker than the postocular setae in vulnerata (fig. 403, a) and whitish and concolorous with the postoculars in affinis (fig. 404, a). The anal lobe of vulnerata invariably differs from that of affinis in having a faint to distinct rounded hyaline spot, either medial in position or based on the posterior wing margin (fig. 401, a), surrounded by dark areas; in affinis, the basal half of the lobe is largely hyaline. The whitish scutal stripes described for affinis are scarcely visible, if present at all, and the scutellum is uniformly subshining dark brown. recognition.

distribution.

Map 75.

Senecio aureus appears to be the only recorded host (Novak et al. 1967, Novak and Foote 1975). host.

Novak and Foote (1975) presented a comprehensive account of the biology and immature stages of vulnerata based on their field studies in northeastern Ohio. discussion.

Genus Strauzia Robineau-Desvoidy Strauzia Robineau-Desvoidy 1830: 718 (type species, Strauzia inermis RobineauDesvoidy 1830: 718, by subsequent designation of Coquillett 1910: 609) ( = longipennis (Wiedemann)). — Cresson 1907: 100 (nomenclature). — Coquillett 1910: 609 (type designation). — Cresson 1914a: 276 (taxonomy). — Foote and Blanc 1963: 63 (review, Calif.). - Foote 1965a: 676 (in catalog). - Steyskal 1986: 101 (revision). — Stoltzfus 1988: 117 (revision). Straussia [emendation]: Loew 1873: 243, 327 (taxonomy). — Williston 1896: 121 (in key). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 602 (in catalog). Williston 1908: 286 (in key). - Flendel 1914b: 82 (in key). - V. T. Phillips 1923: 120, 125 (taxonomy). - Curran 1932b: 3 (in key). - Curran 1934: 291 (in key). Trypeta (Straussia): Osten Sacken 1878: 189 (in catalog). The species of Strauzia, which are relatively large yellowish flies, most strongly resemble those of Euleia, Myoleja, Gymnocarena, and Oedicarena, from which this genus is distinguished most easily by the characters given in the key to genera. Strauzia is characterized by the presence of presutural supra-alar bristles, a decided elongation of cell cup, a flat scutellum, the absence of orbital bristles, the anterior oral margin without well-developed setae, and a distinctive wing pattern. recognition.

Systematic Treatment of the Genera The dorsocentral bristles vary in location but are situated no farther anterior than the level of the supra-alar bristles and no farther posterior than a point halfway between the supra-alar and acrostichal bristles. The aculeus is broad, with the sides of the tip finely serrate as in Trypeta and in most other Trypetina. In 1873, Loew recognized Wiedemann’s species longipennis as the single existing species of Strauzia, but he described seven varieties of this amazingly discussion.

variable taxon. All seven were maintained in the literature as varieties until studies by Steyskal (1986) showed that several species were identifiable among about 1,500 specimens he studied from a wide geographic area. He concluded that longipennis itself, with the variety longitudinalis as a synonym, merited species rank, and he elevated each of the varieties arculata, intermedia, and perfecta to full species as well, but he was not able to resolve the status of confluens, typica, and vittigera, three of Loew’s original varieties. We have adopted Steyskal’s conclusions in our treatment of the genus, although we note his statement that his concept of longipennis is highly variable and that it may be a complex of species. Recently, Stoltzfus (1988) further revised Strauzia. He concurred with Steyskal in the status as full species of three of Loew’s original varieties: arculata, intermedia, and perfecta. Based largely on host plant data, he raised two additional Loew vari¬ eties, longitudinalis and vittigera, to full species status and described noctipennis, rugosum, and uvedalia as new species as well. Following an intensive study of this revision and specimens recently loaned to us by Stoltzfus and in the NMNH collec¬ tion, we conclude that the additional taxa recognized as full species by Stoltzfus are not well delimited on solely morphological grounds. Rather than question their validity, we suggest that studies involving biochemistry, genetics, serology, and addi¬ tional field work are needed to establish the true status of the taxa considered to be valid by Stoltzfus. His paper should be consulted for details of any pertinent biolog¬ ical data we have summarized in this handbook. The larvae of all Strauzia whose biology is known bore in the stems of Asteraceae, feeding on the pith. S. longipennis is economically important in reducing the seed production of commercially grown sunflowers. The larvae weaken the stem, making it more susceptible to breakage. The distribution of this genus is restricted to the United States and Canada.

Key to Known Species of Strauzia (Adapted from Steyskal 1986) L Scutellum entirely yellow, without dark markings; remainder of thorax yellow except male with dark markings on anterior margin of scutum; hyaline band between veins r-m and dm-cu present and continuous from costa into cell cuaj (fig. 405, a); postocellar setae much farther apart than width of ocellar triangle (fig. 414, a).perfecta (Loew) Scutellum yellow with lateral corners dark brown to black; thorax often with additional dark markings in addition to those at anterior margin of scutum; hyaline band between veins r-m and dm-cu, if present, usually

375

Strauzia

413 Figures 405-413. Right wings, Strauzia spp. 405, perfecta (Lw.), female; 406, intermedia (Lw.), male; 407, arculata (Lw.), female; 408, arculata (Lw.), male; 409, stoltzfusi Steysk., female; 410,

longipennis (Wied.), female; 411—413, longipennis (Wied.), males.

not entering cell cuaj (fig. 406, a); postocellar setae much closer together (fig. 415, a) .2 2. Fifth tarsomere largely blackish; hyaline band between veins r-m and dm-cu extending anteriorly to costa, posteriorly almost to vein CuAj but not reaching it (fig. 406, a) .intermedia (Loew) Fifth tarsomere yellowish, at most slightly darkened at apex; hyaline band

Systematic Treatment of the Genera

Figures 414, 415. Front view of head, Strauzici spp. 414, perfecta (Lw.).; 415, longipennis (Wied.). Figures 416-418. Ovipositor of Strauzia spp. (redrawn from Steyskal 1986). 416, verbesinae Steysk.; 417, longipennis (Wied.).; 418, gigantei Steysk. Figure 419. Dorsal view of scutum,

Strauzici stoltzfusi Steysk. Figures 420, 421. Side view of head and thorax, Strauzia spp. 420, gigantei Steysk.; 421, longipennis (Wied.).

between veins r-m and dm-cu variable, sometimes absent, if present, sometimes extending to vein CuAa (fig. 409, a) or not reaching anteriorly to costa (figs. 407, b; 408, b) .3 3. Hyaline band between veins r-m and dm-cu present but not extending ante¬ riorly to costa (fig. 407, b); pterostigma 3 times as long as wide at base; hyaline spot in cell br elongated (figs. 407, a).arculata (Loew) Hyaline band between veins r-m and dm-cu, if present, extending anteriorly to costa but sometimes broken posteriorly (fig. 412, a); pterostigma and

377

Strauzia

hyaline spot in cell br not as elongated and slender as described above .4 4. Aculeus tip shaped as in fig. 416, with distinct angulations at base as shown in fig. 416, a .verbesinae Steyskal Aculeus tip with sides evenly rounded at base (figs. 417, 418) .5 5. Oviscape largely blackish; scutum with pattern as in fig. 419; pleura with well-developed black marks; wing of female as in fig. 409 . .stoltzfusi Steyskal Oviscape darkened at most apically; scutum lacking long submesal and lateral stripes; pleura with at most small faint dark marks except some¬ times above hind coxa.6 6. Pleura with distinct dark mark above hind coxa (fig. 420, b); male with frontal setae not enlarged (fig. 420, a); wing pattern as in fig. 410; acu¬ leus tip slightly rounded (fig. 418) .gigantei Steyskal Pleura with mark above hind coxa faint or completely lacking (fig. 421, b); male with frontal setae usually enlarged (fig. 421, a); wing as in figs. 410—413; aculeus tip sharply pointed (fig. 417) . .longipennis (Wiedemann)

Strauzia arculata (Loew) (Figs. 407, 408; Map 76) Trypeta longipennis var. arculata Loew 1873: 242 (male holotype, MCZ; Ill.). Straussia longipennis var. arculata: V. T. Phillips 1923: 127 (review). Strauzia arculata: Steyskal 1986: 107 (revision). - Stoltzfus 1988: 123 (revision). References not strictly applicable to S. arculata: Straussia longipennis var. arculata: Snow 1894: 160 (taxonomy). - G. M. Greene 1910: 430 (biology). Strauzia longipennis var. arculata: Foote 1965a: 676 (in catalog). The wing of arculata (figs. 407, 408) is recognized by the shape of the pterostigma, which is about three times as long as its width at the base, and by the elongated hyaline spot in cell br (figs. 407, a; 408, a), which terminates apically beyond the apex of the pterostigma (both are longer and narrower than in any other RECOGNITION.

Strauzia species). The short transverse hyaline band between veins r-m and dm-cu, which does not attain the costa anteriorly (figs. 407, b; 408, b) or vein CuA! posteriorly (as in fig. 406, a), also is distinctive. The scutum bears a pair of dark spots laterally; the pleura bears coarse, dark setae; and a black spot is present above the hind coxa. The frontal setae of the male are only occasionally enlarged. distribution.

Map 76.

Steyskal (1986) reported arculata to have been commonly reared from Helianthus grosseserratus. The identification of arculata reported by Steyskal from HOSTS.

H. annuus is possibly in error (Stoltzfus 1988).

378

Systematic Treatment of the Genera

Strauzia arculata ★ Strauzia arculata (state record only)

o

Strauzia gigantei

☆

Strauzia gigantei (state record only) Strauzia verbesinae

A

Map 76. Distribution of Strauzia arculata, S. gigantei, and S. verbesinae.

One of Loew’s original varieties of longipennis, arculata was raised to specific rank by Steyskal (1986), a change with which Stoltzfus concurs. The latter discussion.

believes that specimens reported by Steyskal from other than Illinois, Iowa, and Kansas may be noctipennis Stoltzfus.

Strauzia gigantei Steyskal (Figs. 418, 420; Map 76) Strauzia longipennis var. intermedia [misidentification]: Novak et al. 1967: 148 (host, biology). Strauzia gigantei Steyskal 1986: 107 (female holotype, NMNH; 1 mi s. Kent, Por¬ tage County, Ohio). - Stoltzfus 1988: 124 (revision). In most respects, gigantei closely resembles longipennis. It is dis¬ tinguished from the latter by characters of the aculeus (compare figs. 417 and 418), RECOGNITION.

which in gigantei is more rounded at its apex. Host preference, as well as the dark color of the pleural black spot above the hind coxa (fig. 420, b), further distinguish this species. In longipennis, the pleural spot is weak or absent. The male is similar to the female in wing pattern and in the shapes of the wing and head. distribution.

Map 76.

379

Strauzia

host.

Helianthus giganteus is the only known host.

S. gigantei is rare, having been found only at the type locality in Ohio. Stoltzfus (1988) presented interesting biological data. discussion.

Strauzia intermedia (Loew) (Fig. 406; Map 77) Trypeta longipennis var. intermedia Loew 1873: 241 (male holotype, MCZ; type locality unknown). Straussia longipennis var. intermedia: V. T. Phillips 1923: 127 (review).

Map 77. Distribution of Strauzia intermedia.

380

Systematic Treatment of the Genera

Strauzia intermedia: Steyskal 1986: 108 (revision). Strauzia intermedia Steyskal [error]: Stoltzfus 1988: 124 (revision). References not strictly applicable to S. intermedia: Strauzia longipennis var. intermedia: Leonard 1928: 851 (N.Y.). - Foote 1965a: 676 (in catalog). - Wasbauer 1972: 134 (host). S. intermedia is recognized by the largely black fifth tarsomeres, the yellow oviscape (in some specimens with some basal darkening), and the nature of the transverse hyaline band between veins r-m and dm-cu (see couplet 2 of key and recognition.

fig. 406, a). The scutum has at least one pair of short blackish stripes, the pleura have dark marks, and the lateral margins of the abdominal tergites are usually darkened. The frontal setae of the male are usually enlarged and have blunt tips. distribution.

Map 77.

Rudbeckia laciniata, recorded by Steyskal (1986) and Stoltzfus (1988), is the only known host. HOST.

In his revision, Steyskal (1986) raised intermedia to species status. Additional biological and morphological information will be found in Steyskal (1986) and Stoltzfus (1988). discussion.

Strauzia longipennis (Wiedemann) Sunflower maggot, mouche du tournesol (Figs. 55, 60, 410-413, 415, 417, 421; Map 78) Trypeta longipennis Wiedemann 1830: 483 (male and female syntypes, NMW; Nordamerika”). — Coquillett 1910: 609 (type species designation). Strauzia inermis Robineau-Desvoidy 1830: 718 (type lost; Pa.). - Loew 1862c: 59 (synonymy). - Loew 1873: 336 (synonymy). - Osten Sacken 1878: 190 (in catalog). — Aldrich 1905: 602 (in catalog). Strauzia armata Robineau-Desvoidy 1830: 719 (type lost; Pa.). - Osten Sacken 1878: 190 (synonymy). - Aldrich 1905: 602 (in catalog). Trypeta sepentaria Say [manuscript]: T. W. Harris 1835b: 600 (Mass.). Tephritis trimaculata Macquart

1843a:

383

(male syntype(s), probably lost,

MNHN; North America). - Loew 1873: 338 (synonymy). - Osten Sacken 1878: 190 (synonymy). - Aldrich 1905: 602 (in catalog). - Johnson 1925b: 60 (syn¬ onymy). Trypeta cornigera Walker 1849: 1010 (male holotype, BMNH; North America). Osten Sacken 1858: 79 (in catalog). — Loew 1862c: 58 (synonymy). — Loew 1873: 241, 336 (synonymy).-Osten Sacken 1878: 190 (in catalog). - Aldrich 1905:602 (in catalog). - Foote 1964d: 320 (synonymy, type data, taxonomy).

Strauzia

Map 78. Distribution of Strauzia longipenrtis. Records by Stoltzfus for longitudinalis, noctipennis,

rugosum, uvedaliae, and vittigera are included here.

Trypeta cornifera Walker 1849: 1011 (male holotype, BMNH; N. Y.). - Loew 1862c: 58 (synonymy). - Loew 1873: 336 (taxonomy). - Osten Sacken 1878: 190 (in catalog). - Aldrich 1905: 602 (in catalog). - Foote 1964d: 319 (synonymy, taxonomy). Trypeta septenaria [error]: Osten Sacken 1858: 80 (in catalog). — Loew 1862c: 60 (taxonomy). — Loew 1873: 337 (taxonomy). Trypeta armata: Loew 1862c: 58 (synonymy). Trypeta longipenrtis var. longitudinalis Loew 1873: 240 (male lectotype, MCZ; Sharon Springs, N.Y.). - Steyskal 1986: 108 (lectotype designation, synonymy). Trypeta (Straussia) armata: Loew 1873: 335 (taxonomy). Trypeta (Straussia) longipennis var. longitudinalis: Osten Sacken 1877: 345 (dis¬ tribution). Straussia longipennis: Crevecoeur 1906: 95 (Kans.). Straussia longipennis var. longitudinalis: G. M. Greene 1910: 430 (biology). — V. T. Phillips 1923: 127 (review). - Strickland 1938: 204 (Alta.). Trypeta serpentaria [error]: Johnson 1925a: 97 (synonymy). Straussia longipennis form cornigera: Johnson 1925b: 260 (N. Engl.). Straussia longipennis form longitudinalis: Johnson 1925b: 260 (N. Engl.). Straussia longipennis cornigera: Brimley 1938: 383 (N. Car.). Strauzia longipennis: Steyskal 1986: 108 (revision). — Stoltzfus 1988: 124 (revision).

Systematic Treatment of the Genera

References not strictly applicable to S. longipennis: Trypeta longipennis: Osten Sacken 1858: 79 (in catalog). Trypeta longipennis var. typica Loew 1873: 239 (female lectotype, MCZ; Lancaster, Pa.). — Steyskal 1986: 108 (lectotype designation). Trypeta longipennis var. confluens Loew 1873: 241 (male holotype, MCZ; Conn.). Steyskal 1986: 109 (type data). Trypeta longipennis var. vittigera Loew 1873: 241 (female lectotype, MCZ; Nebr.). - Steyskal 1986: 109 (lectotype designation). Straussia longipennis: Loew 1873: 243, 329, 337 (taxonomy). - Keen 1885: 55 (Pa.). - W. A. Snow 1894: 159 (taxonomy). - Williston 1896: 121 (taxonomy). Doane 1899: 178 (taxonomy). - Chagnon 1901: 14 (Canada). - Johnson 1903b: 106 (N. Mex.). - F. H. Snow 1903: 219 (Kans.). - Aldrich 1905: 602 (in catalog). - Washburn 1905: 118 (Minn.). - Gibson 1906: 120 (biology). - Tucker 1907: 104 (Kans., Colo.). - Williston 1908: 282 (figure of adult). - Johnson 1910: 801 (N.J.). - Woodworth 1913: 137 (Calif.). - Hendel 1914b: 82 (type data). - Winn and Beaulieu 1915: 153 (Que.). - Marcovitch 1916: 147 (description, biology). Britton 1920: 203 (Conn.). - Brink 1923: 72 (biology). - V. T. Phillips 1923: 125 (biology). - Johnson 1925a: 97 (Harris collection). - Johnson 1925b: 260 (N. Engl.). - Sturtevant 1925: 215 (seminal receptacles). - Leonard 1928: 851 (N.Y.). - Curran 1930: 77 (N.Y.). - Petch and Maltaise 1932: 56 (Que.). - Curran 1934: 292 (figures of male and female heads). - F. H. Walker 1936: 21 (biology). Strickland 1938: 175 (Alta.). - V. T. Phillips 1946: 77, 122 (description, biology larvae; hosts). - Hennig 1952: 218 (in catalog of immature stages). - Bray and Triplehorn 1953: 22 (plant associations). Trypeta (Straussia) longipennis: Osten Sacken 1877: 345 (distribution). — Osten Sacken 1878: 189 (in catalog). Straussia longipennis var. typica: W. A. Snow 1894: 160 (taxonomy). - G. M. Greene 1910:430 (biology). - V. T. Phillips 1923: 126 (review). Straussia longipennis var. vittigera: W. A. Snow 1894: 160 (taxonomy). — G. M. Greene 1910: 430 (biology). - V. T. Phillips 1923: 127 (review). - Strickland 1938: 204 (Alta.). - Steyskal 1945: 156 (mating behavior). Strauzia longipennis var.: Cresson 1907: 99 (taxonomy). Strauzia longipennis: Cockerell 1914: 195 (host). - Cockerell 1917: 17 (Colo.). Essig 1938: 602 (review).-Forsell 1947: 167 (Wash.).-Westdahl and Barrett 1960: 481 (biology). - Westdahl and Barrett 1962: 11 (damage). - Foote 1965a: 676 (in catalog).-Schulz and Lipp 1969: 99 (biology, damage).-Lipp and Schulz 1970: 27 (damage). - Beirne 1971: 66 (economic importance, biology). - Kamali and Schulz 1971: 85 (artificial diet). - Wasbauer 1972: 133 (hosts). - Muesebeck 1980: 46 (parasite). - Hilgendorf and Goeden 1981: 106 (host). - Steck 1981: 19 (hosts). Straussia longipennis var. confluens: V. T. Phillips 1923: 127 (review). Straussia Straussia Straussia Straussia

longipennis longipennis longipennis longipennis

form confluens: Johnson 1925b: 260 (N. Engl.). form typica: Johnson 1925b: 260 (N. Engl.). confluens: Brimley 1938: 383 (N. Car.). typica: Brimley 1938: 383 (N. Car.).

383

Strauzia

Straussia longipennis vittigera: Brimley 1938: 383 (N. Car.). Strauzia longipennis var. typica: Foote and Blanc 1963: 63 (review, Calif.). — Foote 1965a: 676 (in catalog). - Novak et al. 1967: 148 (host, biology). - Wasbauer 1972: 134 (host). Strauzia longipennis var. vittigera: Foote and Blanc 1963: 63 (review, Calif.). - Foote 1965a: 676 (in catalog). - Novak et al. 1967: 148 (host, biology). - Wasbauer 1972: 134 (hosts). Strauzia longipennis var. confluens: Foote 1965a: 676 (in catalog). — Wasbauer 1972: 134 (hosts). The female wing pattern of longipennis shown in fig. 410 is “typi¬ cal” of this species, and although it resembles that of several other species of Strauzia discussed herein, the characters in the key will assist in distinguishing this species. As can be seen in figs. 411-413, the wing pattern of the male varies tremendously, but this sexual dimorphism is rarely present in the other species of Strauzia as defined by Steyskal (1986). The males of most forms of longipennis have enlarged, blunt-tipped recognition.

frontal bristles (fig. 421, a), and the wings are generally narrower and more sharply pointed than in females or in either sex of any of the other species of Strauzia. The dark pleural spot above the hind coxa is absent or poorly developed (fig. 421, b), the other pleural spots are generally small and faint, no paired dark stripes are present on the scutum, and the fifth tarsomeres are largely yellow, as is the oviscape. Because longipennis is so variable, one should rely on the use of the key rather than on any distinctive characteristic of this species for its identification. distribution.

Map 78. Distribution records by Stoltzfus (1988) for longitudi¬

nals, noctipennis, rugosum, uvedaliae, and vittigera are also included. hosts.

Smallantbus uvedalia, Ageratina altissima, and three species of Helian-

thus are hosts of the material studied by Steyskal (1986). S. longipennis probably infests other species of Heliantbus as well. Since its revision by Loew (1873), Strauzia has been recognized as containing only one species, longipennis, with seven varieties. Steyskal (1986) re¬ stricted this concept of longipennis to “typical1 longipennis, with which he syndiscussion.

onymized four of Loew’s varieties. In his recent revision of Strauzia, Stoltzfus (1988) regarded longipennis as com¬ prising individuals reared only from Heliantbus annuus', this concept includes Loew’s variety typica. Stoltzfus also raised Loew s varieties longitudinalis and vit¬ tigera to full species status. He described three host-specific species: S. noctipennis, reared from Heliantbus grosseserratus; S. rugosum, from Eupatorium rugosum; and S. uvedaliae, from Smallantbus uvedalia. As stated in the discussion of the genus, none of these species is morphologically distinct because of interspecific variation in color and wing pattern. Furthermore, we have no proof at this time that those taxa, based as they are mainly on host preference, are deserving of full species status. Despite his careful work, Steyskal admits that longipennis, although not in any

384

Systematic Treatment of the Genera sense a dumping ground, remains a composite of various populations requiring further investigation, and the actions by Stoltzfus appear to emphasize this conclu¬ sion. Therefore, only those literature references definitely assignable to longipennis in the sense of Steyskal, whose conclusions we follow (although somewhat ar¬ bitrarily), are included in the synonymy. This leaves a large body of literature which is listed separately above, not being strictly assignable to longipennis in our sense. As further work is done on the longipennis complex, many of these references may become associated with it or with the separate taxa that Stoltzfus has recognized.

Strauzia noctipennis Stoltzfus Strauzia noctipennis Stoltzfus 1988: 124 (male holotype, NMNH; Ames, Iowa). Not mapped (see caption, Map 78). Alberta south to Indiana, Wisconsin and New Mexico. distribution.

host. Heliantbus grosseserratus is the only known host. discussion. Stoltzfus (1988) included figures of the surstyli, aculeus tip, dis-

tiphallus, and three different kinds of male wing pattern. For the reasons described above, we have not been able to include this species in our key.

Strauzia perfecta (Loew) (Figs. 405, 414; Map 79) Trypeta longipennis var. perfecta Loew 1873: 239 (male and female syntypes, MCZ; type locality not known). Straussia longipennis var. perfecta: W. A. Snow 1894: 159 (taxonomy). - V. T. Phillips 1923: 126 (review). — Steyskal 1972d: 130 (taxonomy). Strauzia longipennis var. perfecta: Novak et al. 1967: 148 (host, biology). Strauzia perfecta: Steyskal 1986: 109 (revision). - Stoltzfus 1988: 125 (revision). References not strictly applicable to S. perfecta: Straussia longipennis var. perfecta: G. H. Greene 1910: 430 (biology). - Leonard 1928: 851 (N.Y.). Straussia longipennis perfecta: Brimley 1938: 383 (N. Car.). Strauzia longipennis var. perfecta: Foote 1965a: 676 (in catalog). - Wasbauer 1972: 134 (host). recognition. S. perfecta is recognized by being almost entirely yellow, lacking

the usual dark markings at the corners of the scutellum. The female lacks the normal dark spots at the anterior margin of the scutum, but a very broad, black apical ring is present on the oviscape. The additional characters stated in couplet 1 of the key to species further distinguish perfecta from all other Strauzia species.

385

Strauzia

distribution.

HOST.

Map 79.

According to Steyskal (1986), perfecta appears to be restricted to Ambrosia

trifida. This plant also was mentioned as a host by Stoltzfus (1988). discussion.

Formerly one of Loew’s seven varieties, perfecta is fairly common

throughout its geographic range.

Strauzia rugosum Stoltzfus Strauzia rugosum Stoltzfus 1988: 125 (female holotype, NMNH; 2 mi. south Wadsworth, Ohio).

386

Systematic Treatment of the Genera distribution.

Not mapped (see caption, Map 78). Ohio, Virginia, West Vir¬

ginia. host.

Larvae were collected by Stoltzfus (1988) from Eupatorium rugosum.

Stoltzfus (1988) included figures of the aculeus tip and female wing. For the reasons described above, we are not able to include rugosum in our key to species. discussion.

Strauzia stoltzfusi Steyskal (Figs. 409, 419; Map 79) Strauzia stoltzfusi Steyskal 1986: 110 (female holotype, NMNH; St. Anthony Park,

Ramsay County, Minn.). - Stoltzfus 1988: 125 (revision). recognition.

S. stoltzfusi shows much more extensive body darkening than any

other Strauzia, although the fifth tarsomeres are largely yellowish. The abdominal tergites bear conspicuous broad, ill-defined, brown crossbands; the oviscape is al¬ most entirely black; the pleura have well-developed blackened areas; and the scutum is rather heavily marked as in fig. 419. The wing pattern of both sexes (see fig. 409) resembles that of longipennis, but the frontal bristles of the male are not enlarged or bluntly tipped. distribution.

HOST.

Not

Map 79.

known.

An extremely rare species, stoltzfusi was described from only 2 specimens among the 1,500 Strauzia specimens studied by Steyskal (1986). Mor¬ phological traits were discussed briefly by Stoltzfus (1988). discussion.

Strauzia uvedaliae Stoltzfus Strauzia uvedaliae Stoltzfus 1988: 125 (male holotype, NMNH; 1 mi. south Elkton

Va.). distribution.

Not mapped (see caption, Map 78). Maryland, Virginia, North

Carolina. host.

The only known host is Smallanthus uvedalia.

Stoltzfus (1988) figured the head, aculeus tip, and male wing pat¬ tern. For the reasons described above, we have not included uvedaliae in our key. discussion.

387

Tepbritis

Strauzia verbesinae Steyskal (Fig. 416; Map 76) Strauzia verbesinae Steyskal 1986: 110 (female holotype, NMNH; 5 mi e. Harrison¬ burg, Rockingham County, Va.). — Stoltzfus 1988: 125 (revision). In many respects, especially in wing pattern, verbesinae resembles longipennis (as in fig. 410), from which it is distinguished primarily by the characters of the aculeus tip (compare figs. 416, a and 417, a). The oviscape is yellow with only a slight apical darkening, the thorax has dark markings only above the wing base on the lateral corners of the scutellum, on the anterior margin of the scutum, and on the lower lateral corners of the mediotergite. The posterior two pairs of frontal bristles of recognition.

the males are enlarged and have blunt tips. distribution.

host.

Map 76.

Verbesina occidentalis is the only host recorded by Steyskal (1986) and

Stoltzfus (1988). S. verbesinae is a rare species. Steyskal (1986) encountered only about 30 specimens among the 1,500 Strauzia he studied. Stoltzfus (1988) presented discussion.

brief biological observations.

Genus Tephritis Latreille Tepbritis Latreille 1804: 196 (type species, Musca arnicae Linnaeus 1758: 600, by designation of Cresson 1914a: 278). - Loew 1862b: 49 (taxonomy). - Loew (part) 1873: 295, 328 (taxonomy, in key). - Williston (part) 1896: 123 (in key). Coquillett 1899c: 268 (in key). - Aldrich (part) 1905: 611 (in catalog). - Cresson 1907: 99 (taxonomy). - Williston 1908: 287 (in key). - Coquillett 1910: 613 (type data). - Cresson 1914a: 277, 278 (type designation). - Curran (part) 1932b: 4 (in key). - Curran (part) 1934: 291 (in key). - Hendel 1935: 53 (taxonomy). - Hering 1944b: 17 (key to species, world). - Foote 1959a: 16 (taxonomy). - Foote 1960e: 71 (revision). - Foote 1960f: 146 (taxonomy). - Foote 1960g: 3 (taxonomy). Foote and Blanc 1963:64 (review, Calif.). — Foote 1965a: 667 (in catalog). — Wasbauer 1972: 134 (hosts). — Foote and Freidberg 1980. 32 (type data). — Freidberg 1984: 129 (galls).-Jenkins 1985: 1 (spp. attacking Baccharis). -Jenkins and Turner 1989: 674 (spp. attacking Baccbaris). - Maddison and Bartlett 1989: 32 (zoogeography). Euaresta Loew: Loew (part) 1873: 322, 328 (taxonomy, in key). — Aldrich (part) 1905: 612 (in catalog). - V. T. Phillips (part) 1923: 123, 145 (in key, review e. U.S.). - Curran (part) 1932b: 4 (synonymy).

387

Tepbritis

Strauzia verbesinae Steyskal (Fig. 416; Map 76) Strauzia verbesinae Steyskal 1986: 110 (female holotype, NMNH; 5 mi e. Harrison¬ burg, Rockingham County, Va.). — Stoltzfus 1988: 125 (revision). In many respects, especially in wing pattern, verbesinae resembles longipennis (as in fig. 410), from which it is distinguished primarily by the characters of the aculeus tip (compare figs. 416, a and 417, a). The oviscape is yellow with only a slight apical darkening, the thorax has dark markings only above the wing base on the lateral corners of the scutellum, on the anterior margin of the scutum, and on the lower lateral corners of the mediotergite. The posterior two pairs of frontal bristles of recognition.

the males are enlarged and have blunt tips. distribution.

host.

Map 76.

Verbesina occidentalis is the only host recorded by Steyskal (1986) and

Stoltzfus (1988). S. verbesinae is a rare species. Steyskal (1986) encountered only about 30 specimens among the 1,500 Strauzia he studied. Stoltzfus (1988) presented discussion.

brief biological observations.

Genus Tephritis Latreille Tepbritis Latreille 1804: 196 (type species, Musca arnicae Linnaeus 1758: 600, by designation of Cresson 1914a: 278). - Loew 1862b: 49 (taxonomy). - Loew (part) 1873: 295, 328 (taxonomy, in key). - Williston (part) 1896: 123 (in key). Coquillett 1899c: 268 (in key). - Aldrich (part) 1905: 611 (in catalog). - Cresson 1907: 99 (taxonomy). - Williston 1908: 287 (in key). - Coquillett 1910: 613 (type data). - Cresson 1914a: 277, 278 (type designation). - Curran (part) 1932b: 4 (in key). - Curran (part) 1934: 291 (in key). - Hendel 1935: 53 (taxonomy). - Hering 1944b: 17 (key to species, world). - Foote 1959a: 16 (taxonomy). - Foote 1960e: 71 (revision). - Foote 1960f: 146 (taxonomy). - Foote 1960g: 3 (taxonomy). Foote and Blanc 1963:64 (review, Calif.). — Foote 1965a: 667 (in catalog). — Wasbauer 1972: 134 (hosts). — Foote and Freidberg 1980. 32 (type data). — Freidberg 1984: 129 (galls).-Jenkins 1985: 1 (spp. attacking Baccharis). -Jenkins and Turner 1989: 674 (spp. attacking Baccbaris). - Maddison and Bartlett 1989: 32 (zoogeography). Euaresta Loew: Loew (part) 1873: 322, 328 (taxonomy, in key). — Aldrich (part) 1905: 612 (in catalog). - V. T. Phillips (part) 1923: 123, 145 (in key, review e. U.S.). - Curran (part) 1932b: 4 (synonymy).

Systematic Treatment of the Genera Trypeta (Tephritis): Osten Sacken 1878: 193 (in catalog). Trypeta (Euaresta): Osten Sacken (part) 1878: 194 (in catalog). Urellia Robineau-Desvoidy: Aldrich (part) 1905: 613 (in catalog). - Williston (part) 1908: 287 (in key). Euribia Meigen: Hendel (part) 1914b: 96 (in key). Trupanea (Tepbritoides) Benjamin 1934: 58 (type species, Euaresta subpura Johnson 1909: 114, by original designation). The genus Tephritis may be recognized by the following combina¬ tion of characters: head with distinct fronto-facial angle; nongeniculate proboscis; RECOGNITION.

lower facial margin prominent; two pairs of frontal bristles (except in baccharis, which sometimes has one or two additional pairs of weak setae); two pairs of orbital bristles, the posterior pair usually white or yellowish and reclinate; two pairs of scutellar bristles; one pair of dorsocentral bristles, situated very close to the tranverse suture; the hind femur with one or two preapical anterodorsal bristles, never situated side by side; wing pattern sometimes extensively darkened but usually reticulate; pterostigma dark centrally without a central hyaline spot. These characters place the species of Tephritis in a large assemblage of small fruit flies that breed in the flower heads and stems of a wide variety of Asteraceae and a few other plant families. From all small species of tephritids having reticulate wings, Tephritis species may invaria¬ bly be recognized by the presence of two pairs of frontal and two pairs of scutellar bristles, the number and arrangement of the minute anterodorsal bristles on the hind femur, and an extension of the dark area in the pterostigma posteriorly into the proximal portions of at least cells r2+3 and r4+5 (fig. 437, a). The genera most likely to be confused with Tephritis are easily distinguished from it by a number of distinc¬ tive characters. Euaresta males have a swollen fore femur and distinctive genitalia and both sexes usually possess a bulla and lack a distinct fronto-facial angle; Tru¬ panea has only one pair of scutellar bristles; Dioxyna, Gonioxya, and Paroxyna possess squarish heads with distinctly geniculate probosces; and Neotephritis and other similar genera (see couplet 46, key to genera) have three pairs of frontal bristles. Jenkins and Turner (1989) also suggested that the shape of the distiphallus is diagnostic for Tephritis, at least in the New World. The generic limits of Tephritis have been the object of much discus¬ sion in the literature. Because the very large number of similar, unnamed Neotropical discussion.

taxa have never been studied to any appreciable extent, and because other taxonomic characters in the known species have yet to be thoroughly evaluated, we maintain the somewhat arbitrary classification in which U.S. and Canadian Tephritis includes all those species with two pairs of frontal bristles (except baccharis, which may have up to four) and two pairs of scutellar bristles, with a nongeniculate proboscis, and without expanded, ridged, Euaresta-type male genitalia. This results in the inclusion of several species with rather aberrant wing patterns, at least two of which closely resemble those of Trupanea, a situation discussed at some length by Foote (1959a). The U.S. and Canadian species of Tephritis have been reared from a variety of composite genera, mainly in the Astereae, Anthemideae, and Senecioneae. In some

389

Tepbritis species, the generations alternate between gall formation and flower head feeding. In several species reared from Baccharis, these generations differ in color and were termed light and dark morphs by Jenkins (1985) and Jenkins and Turner (1989) in several species of Tepbritis reared from Baccharis. These studies, and recent biolog¬ ical investigations reported by Goeden (see applicable synonymies) and under way by him, indicate that the recognition of species in this genus may not be as straight¬ forward as our presentation in this handbook may suggest. The results of future biological studies may alter our concepts at the species level.

Key to U.S. and Canadian Species of Tephritis 1. Light areas of wing yellowed, not consisting of white hyaline spots that contrast strongly with dark areas; wing pattern as in fig. 422 . .baccharis (Coquillett) Light areas of wing comprising whitish hyaline spots that contrast strongly with the dark areas .2 2. Wing pattern as in Trupanea; cell dm hyaline except for 2 dark transverse rays across distal half (fig. 423, b, c); dark bar from pterostigma to vein r-m without any hyaline interruptions or spots (fig. 423, a).3 Wing pattern otherwise; cell dm with hyaline spots variously fused; dark bar from pterostigma to posterior wing margin with a few hyaline spots (fig. 425, d) .4 3. Both dark rays in cell dm extending across vein CuA,, the distalmost one to posterior wing margin (fig. 423, b), the proximal one to center of cell cua, (fig. 423, c), where it is associated with a dark streak basad (fig. 423, d) .stigmatica (Coquillett) Neither of the rays in cell dm extending into cell cuaA (fig. 424, b). .labecula Foote 4. An uninterrupted hyaline crescent at wing apex consisting of the hyaline apices of cells r2+3, r4 + 5, and m (fig. 425, c) .rufipennis Doane This hyaline area always interrupted by a dark mark lying upon apices of veins R2+3 or R4f5, or both (fig. 429, d) .5 5. Thorax predominantly bright yellow tomentose in dry specimens; cell c without distinct brownish spots (fig. 426, a); preapical brownish area entire and not broken by many small hyaline spots (fig. 426, b) . .subpura (Johnson) Combination of characters not as above.6 6. Preapical brown area of wing interrupted by numerous round subhyaline spots (fig. 428, c); dark pattern in cell r1 immediately posterior to pterostigma usually with at least 1 yellowish or hyaline spot (fig. 429, b); if no such spot present, anterior arm of Y-shaped mark at wing apex broken or conspicuously narrower than posterior arm (fig. 429, d) .7 Preapical brown area of wing usually with at most a few round subhyaline spots, its appearance never obscured by them (fig. 430, b); dark pattern in cell rj immediately posterior to pterostigma unmarked (fig. 430, a); Y-shaped mark in wing apex never as above.10

a

Figures 422-431. Right wings, Tephritis spp. 422, baccharis (Coq.); 423, stigmatica (Coq.); 424, labecula Foote; 425, rufipennis Doane; 426, subpura (Johns.); 427, arizonaensis Quis.; 428, Califor¬ nia Doane; 429, dilacerata (Lw.); 430, signatipennis Foote; 431, angustipennis (Lw.).

390

391

Tephritis

7. Basal half of cell dm with few or no dark markings (fig. 427, b) .8 Basal half of cell dm with extensive dark markings.9 8. Basal half of cell dm almost entirely hyaline (fig. 427, b); male sternite 5

with small posteromedial patch of short, stout, black setae. .arizonaensis Quisenberry Basal half of cell dm with a dark central spot (as in fig. 428, e); male sternite 5 without posteromedial patch of setae.palmeri Jenkins 9. Basal V3 of wing almost hyaline (fig. 429, a); pterostigma with hyaline area medially (fig. 429, b).dilacerata (Loew) Basal V3 of wing darkened (fig. 428); pterostigma dark medially, sometimes entirely dark (fig. 428, a).californica Doane 10. Dark brown of wing pattern extending to posterior margin along entire length of wing (fig. 430, c) .signatipennis Foote Anal lobe never darkened at wing margin (fig. 431, c).11 11. Dark pattern in cell m never attaining posterior margin of that cell (fig. 431, b).angustipennis (Loew) Dark pattern in cell m always extending close to, or touching, posterior margin of that cell (fig. 432, d) .12 12. Apical margin of second hyaline spot in cell rj never extending apicad beyond anterior extension of vein dm-cu (fig. 433, b).13 Apical margin of second hyaline spot in cell rj extending well beyond ante¬ rior extension of vein dm-cu (fig. 435,'c).14 13. Vein CuA! broadly bordered with brown beyond its junction with vein dmcu, at least in cell m (fig. 432, e); cell be darkened (fig. 432, a). .webbii Doane Vein CuA, narrowly bordered with brown beyond its junction with vein dm-cu (fig. 433, c); cell be largely hyaline (fig. 433, a) . .pura (Loew) 14. At least 3 hyaline spots present in cell r1 beyond apex of pterostigma (fig. 434, a).michiganensis Quisenberry Never more than 2 hyaline spots in cell rt beyond apex of pterostigma (fig. 435, b) .15 15. Dark band from pterostigma to vein R4+5 situated at right angles to hori¬ zontal axis of wing (fig. 436, a) .16 Dark band from pterostigma extending obliquely to cover vein r-m (fig. 437, a) .17 16. Apex of cell br largely dark, although hyaline spots may be present (fig. 435, a); Y-shaped mark in apex of cell r4+5 rarely with broken arms (fig. 435, d); hind tibiae with row of distinct blackish anterodorsal setae (fig. 439, a) .leavittensis Blanc Apex of cell br largely hyaline (fig. 436, b); Y-shaped mark in apex of cell r4 + 5 often incomplete (fig. 436, c); hind tibia lacking a distinct row of prominent setae .candidipennis Foote 17. Femora primarily light brown to yellow; if dark gray dusting present, never so heavy as to obscure yellow ground color; frons yellow; wing length usually under 3.5 mm; wing pattern as in fig. 437 ... .araneosa (Coquillett) Femora, especially those of hind legs, dark tomentose; frons reddish browntomentose; wing length usually exceeding 3.5 mm; wing pattern as in fig. 43 g .ovatipennis Foote

Figures 432-438. Right wings, Tephritis spp. 432, webbii Doane; 433, pura (Lw.); 434, michiganensts Quis.; 435, leavittensis Blanc; 436, candidipennis Foote; 437, araneosa (Coq )■ 438 ovatipennts Foote. Figure 439. Anterior view, hind leg, Tephritis leavittensis Blanc

392 '

393

Tephritis

Tephritis angustipennis Tephritis candidipennis ▲ Tephritis dilacerata

•

♦

Map 80. Distribution of Tephritis angustipennis, T. candidipennis, and T. dilacerata.

Tephritis angustipennis (Loew) (Fig. 431; Map 80) Trypeta angustipennis Loew 1844: 382 (male and female syntypes, PZMHU; Scan¬ dinavia). - Loew 1873: 293 (review). Tephritis angustipennis: Loew 1873: 296, 330 (taxonomy). — Aldrich 1905: 611 (in catalog). - Hendel 1927: 182 (review, world). - Hering 1944b: 31 (in key). Quisenberry 1950: 11 (taxonomy). - Quisenberry 1951: 67 (review, N.A.). Foote 1960e: 78 (review, N.A.). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 134 (hosts). - Foote 1984: 127 (in catalog).

Systematic Treatment of the Genera

Trypeta (Tepbritis) angustipennis: Osten Sacken 1878: 193 (in catalog). Euaresta angustipennis: V. T. Phillips 1923: 147 (taxonomy). - Johnson 1925b: 264 (N. Engl.). - Johnson 1927: 218 (Mt. Desert). - Leonard 1928: 853 (N.Y.). Strickland 1938: 204 (Alta.).

Tepbritis angustipennis [error]: Foote 1960e: 80 (taxonomy). recognition. The wing pattern of

angustipennis resembles that of araneosa, but

in the former, the dark ray occupying the pterostigma descends almost vertically from that cell (fig. 431, a) and is not contiguous with vein r-m, and the dark markings in cell m never attain the posterior margin of that cell (fig. 431, b). The legs of angustipennis are yellow except for the blackened basal halves of the hind femora; the ground color of the thorax and abdomen is black, the tomentum distinctly gray; the scutellum is entirely dark; and the oviscape is 1.5-1.75 times as long as the terminal abdominal tergite.

distribution. Map 80. T.

angustipennis also occurs in northern Europe and

western areas of the former USSR.

hosts.

Achillea ptarmica and Aster occidentalis var. intermedins were the only

hosts recorded by Wasbauer (1972).

Tephritis araneosa (Coquillett) (Figs. 80, 90, 437; Map 81)

Trypeta (Euaresta) araneosa Coquillett 1894: 74 (female syntypes, NMNH; Los Angeles, Calif.).

Euaresta aldrichii: Coquillett 1899c: 266 (taxonomy). Euaresta araneosa: Coquillett 1899c: 266 (synonymy, taxonomy). — Doane 1900: 48 (taxonomy). - C. F. Baker 1904: 31 (Calif., Nev.). - Aldrich 1905: 612 (in catalog). - Cresson 1907: 105 (taxonomy). - Tucker 1908: 303 (Oreg.). - Cole and Lovett 1921: 327 (Oreg.). - Forsell 1947: 167 (Wash.).

Urellia aldrichii Doane 1899: 192 (female lectotype, WSU; Brookings, S. Dak.). Coquillett 1899c: 266 (taxonomy). - C. F. Adams 1904: 450 (in key). - Aldrich 1905: 613 (in catalog). - Foote 1966b: 121 (lectotype designation). — Zack 1984: 33 (type data).

Urellia pacifica Doane 1899: 192 (female lectotype, WSU; Corvallis, Oreg.). Coquillett 1899c: 266 (synonymy). - Doane 1900: 48 (taxonomy). - C. F. Adams 1904: 450 (in key). - Aldrich 1905: 614 (in catalog). - Cole and Lovett 1921: 327 (in list, Oreg.). - Ballou 1925: 1 (description of gall). - Foote 1966b: 124 (lec¬ totype designation). - Zack 1984: 33 (type data).

Tephritis aldrichii: Cole and Lovett 1921: 327 (Oreg.). - Janes and Thomas 1932: 103 (Utah). - Ouellet 1941: 130 (Que.). - Hering 1944b: 31 (in key, world). Trypanea pacifica: Sweet 1930: 123, 130 (Calif.).

Tephritis pacifica: Knowlton and Harmston 1937: 145 (Utah). - Hering 1944b: 25 (in key, world). - Quisenberry 1951: 65 (synonymy). - Foxlee 1957: 36 (B.C.).

395

Tephritis

Tepbritis aldricbii Hine [error]: Strickland 1938: 204 (Alta.). Tephritis araneosa: Hering 1944b: 31 (in key, world). - Quisenberry 1950: 11 (taxonomy). - Quisenberry 1951: 65 (review). - Foote 1960e: 84 (review). - Foote and Blanc 1963: 64 (review, Calif.). - Foote 1965a: 668 (in catalog). - Lavigne 1965: 100 (host). - Wasbauer 1972: 134 (hosts). - Goeden and Benjamin 1985: 392 (fungal parasite).

Euaresta pacifica: V. T. Phillips 1946: 114 (hosts). Trupanea pacifica: V. T. Phillips 1946: 125 (host). recognition. T. araneosa, the most commonly encountered Tephritis in Amer¬ ica north of Mexico, is distinguished by the characters given in the key to species. It may be separated from four similar species (candidiperinis, leavittensis, ovatipennis,

396

Systematic Treatment of the Genera signatipennis) by the following combination of characters: dark ray connecting pterostigma with vein r-m lying at an angle to the horizontal axis of the wing (fig. 437, a); dark markings in cell a never extending to posterior wing margin (fig. 437, b); femora primarily yellow to light brown; the oviscape only slightly longer or shorter than, or equal to, the terminal abdominal tergite; and anterodorsal setae lacking on the hind tibia. See the discussions of the other aforementioned species for additional comments. DISTRIBUTION.

Map 81.

Wasbauer (1972) recorded the following plant genera as hosts for this species: Arnica, Artemisia, Chrysothamnus, Erigeron, Grindelia, Poa, and Tanacetum. hosts.

As presented here, araneosa belongs to a complex of closely related species, the precise identification of which may never be attained without extensive DISCUSSION.

biological studies. Foote (1960e) described three of these (candidipennis, ovatipennis, signatipennis) to distinguish those larger species with longer oviscapes, and Blanc (Foote and Blanc 1979) described another, leavittensis, on the basis of addi¬ tional characters. Other segregates will almost certainly be discovered as this com¬ plex is studied further.

Tephritis arizonaensis Quisenberry (Fig. 427; Map 82) Tephritis arizonaensis Quisenberry 1951: 62 (female holotype, UKL; Tumacacori Mts., Ariz.). - Byers et al. 1962: 180 (type data). - Jenkins 1985: 28 (revision, spp. attacking Baccharis). — Jenkins and Turner 1989: 676 (revision, spp. attacking Baccharis). Tephritis arizonensis [error]: Foote 1960e: 76 (review, N. A.). - Foote 1962: 172 (taxonomy). - Foote and Blanc 1963: 67 (review, Calif.). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 134 (hosts). Somewhat resembling californica and palmeri, arizonaensis differs in being smaller, in having less contrasting wing markings, in bearing a distinct, recognition.

dark, apical Y-shaped mark without the presence of a diagonal hyaline area in the apices of cells r2+3 and r4+5 (fig. 427, a), and in cell a being hyaline along its posterior margin (fig. 427, c). In most specimens of arizonaensis, the dark band connecting the pterostigma with vein r-m is punctuated by relatively large hyaline spots, and cell dm is largely hyaline (fig. 427, b). The ground color of the thorax is black with heavy yellow tomentum and the ovscape is yellow and about 2.0 times as long as the terminal abdominal tergite. Males can be distinguished from those of all other Tephritis species by the patch of small setae posteromedially on tergite 5. distribution.

Map 82.

397

Tephritis

Map 82. Distribution of Tephritis arizonaensis, T. baccharis, T. leavittensis, T. rufipennis, and T. signatipennis.

Jenkins (1985) and Jenkins and Turner (1989) recorded arizonaensis as having been reared from terminal galls, stem tip mines, and female flowers of Bac¬ charis sarothroides. The species has been “taken on” B. sergiloides. hosts.

Jenkins (1985) and Jenkins and Turner (1989) described the light and dark morphs of this species. discussion.

Tephritis baccharis (Coquillett) (Fig. 422; Map 82) Trypeta (Tephritis) baccharis: Coquillett 1894: 73 (female lectotype, NMNH; Los Angeles County, Calif.). — Quisenberry 1951: 59 (taxonomy). — Jenkins 1985: 34 (lectotype designation). - Jenkins and Turner 1989: 677 (lectotype designation). Icterica fasciata Adams 1904: 449 (female lectotype, UKL: Bill Williams Fork, Ariz.). - F. H. Snow 1904: 345 (Ariz.). — Byers et al. 1962: 180 (type data). — Foote 1962: 174 (lectotype designation). — Steyskal and Foote 1963: 166 (synonymy). Trypeta baccharis: Aldrich 1905: 604 (in catalog). — Thompson 1907: 71 (biology). — Bates 1934a: 295 (taxonomy). — Essig 1938: 602 (review). - V. T. Phillips 1946: 126 (host). — Essig 1958: 602 (description of gall). — Foote 1960b: 254 (taxonomy, nomenclature). Tephritis baccharis: Aldrich 1907: 6 (in catalog). - Bates 1934a: 295 (taxonomy). V. T. Phillips 1946: 122 (host). — Stoltzfus 1977: 373 (taxonomy).—Jenkins 1985:

Systematic Treatment of the Genera 33 (revision, spp. attacking Baccharis). - Jenkins and Turner 1989: 677 (revision, spp. attacking Baccharis). - Goeden and Headrick 1991b: 86 (life history). Eutreta baccharis: Quisenberry 1951: 59 (generic placement). - Foote 1962: 174 (taxonomy). - Foote and Blanc 1963: 28 (review, Calif.). - Foote 1965a: 661 (in catalog). - Wasbauer 1972: 116 (hosts). Like subpura, baccharis is one of the few Tephritis species in America north of Mexico having the ground color of the body yellow rather than recognition.

brown to black, although the body tomentum tends to be brown. The frontal bristles are small, variable in length, and (unlike most North American Tephritis) are present in 2-4 pairs. The dark parts of the wing pattern (fig. 422) are distinctly brownish rather than dark brown to black, and the lighter areas are suffused with brown rather than being distinctly hyaline. The wing pattern resembles those of rufipennis (fig. 425) and dilacerata (fig. 429) in consisting essentially of two transverse dark bands, but the characters given above and in the key to species will distinguish baccharis from all other Tephritis species in our area. The oviscape of baccharis is entirely light yellow and almost 2.0 times as long as the terminal abdominal tergite. distribution.

Map 82.

Jenkins (1985) and Jenkins and Turner (1989) reported baccharis from Baccharis salicifolia (as B. glutinosa) and B. viminea. It is known to be a monophagous, obligate gall former. HOSTS.

At first glance, baccharis does not appear to belong in Tephritis as we have defined the genus, but the bristles and other characters place it here without discussion.

doubt. It was described and discussed in detail by Jenkins (1985) and Jenkins and Turner (1989).

Tephritis californica Doane (Fig. 428; Map 83) Tephritis californica Doane 1899: 190 (female holotype, WSU; Palo Alto, Calif.). Coquillett 1899c: 266 (synonymy). - Doane 1900: 48 (taxonomy). - Aldrich 1905: 611 (in catalog). - Quisenberry 1951: 64 (review, N.A.). - Foote 1960e: 76 (review, N.A.). - Foote and Blanc 1963: 67 (review, Calif.). - Foote 1965a: 668 (in catalog). - Foote 1966b: 122 (type data). - Wasbauer 1972: 134 (hosts). - Zack 1984. 33 (type data). — Goeden and Benjamin 1985: 392 (fungal parasite). — Jenkins 1985. 38 (revision, spp. attacking Baccharis). —Jenkins and Turner 1989: 678 (revision, spp. attacking Baccharis). Trypeta californica: Woodworth 1913: 137 (Calif.). T. californica is distinctive among most species of Tephritis in America north of Mexico in having an oblique hyaline area in the apices of cells r7 +3 recognition.

399

Tephritis

and r4 + s (fig. 428, d). This hyaline area is usually interrupted to some extent by a dark spot at the extreme apex of vein R4+5, and in contrast to the apical crescent in rufipennis, it does not continue into the apex of cell m and lies at a distinct angle to the horizontal axis of the wing rather than perpendicular to it. In californica, the apical dark Y-shaped mark is usually obscured by other markings. The oviscape usually bears an hourglass mark centrally and is about 1.8 times as long as the terminal abdominal tergite. T. palmeri closely resembles californica in many mor¬ phological characters; see the recognition and discussion sections of the former species for characters separating the two. distribution.

Map 83.

Baccbaris pilularis and B. p. consanguinea were recorded as hosts of californica by Jenkins (1985) and Jenkins and Turner (1989). As in related species, the light morph probably emerges from stems and the dark morph from flowers. A species of Senecio and B. emoryi, also reported as hosts by Wasbauer (1972), have hosts.

not been confirmed. discussion.

The existence of dark morphs of californica was discussed in con¬

siderable detail by Jenkins (1985) and Jenkins and Turner (1989). Because of the morphological similarities of californica and palmeri, they are considered to be sibling species by those two authors.

400

Systematic Treatment of the Genera

Tephritis candidipennis Foote (Fig. 436; Map 80) Tephritis araneosa (Coquillett): Quisenberry (part) 1951: 66 (figure of wing). Tephritis candidipennis Foote 1960e: 82 (female holotype, NMNH; Fieldbrook, Calif). - Foote and Blanc 1963: 68 (review, Calif.). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 135 (hosts). - Arnaud 1979: 332 (type data). One of the members of the araneosa complex, candidipennis is distinguished from araneosa by its longer oviscape, which is about 1.5 times as long as the terminal abdominal tergite and is dark brown to black with a yellow border RECOGNITION.

dorsally; by the darkened dorsal and ventral surfaces of the middle and hind femora; and by the enlarged and coalesced hyaline areas distad of and below the pterostigma (fig. 436, b), giving the dark band connecting the pterostigma and vein r-m the appearance of descending at right angles to the horizontal axis of the wing (fig. 436, a). The hyaline areas of the wing are more extensive than in any other of our Tephritis species, and the Y-shaped mark at the apex of cell r4+5 is never complete; these are characters which distinguish it from all other species similar to araneosa. distribution.

Map 80.

One species each of Ambrosia and Arnica and two species of Tanacetum were recorded as hosts by Wasbauer (1972). HOSTS.

T. candidipennis araneosa complex.

discussion. and the

is discussed further in our treatment of

araneosa

Tephritis dilacerata (Loew) (Figs. 94, 429; Map 80) Trypeta dilacerata Loew 1846: 509 (male and female syntypes, PZMHU; Germany [various regions] and Poznan, Poland). Tephritis dilacerata: Goeden et al. 1974: 491 (status of studies, host). - Berube 1978a: 69 (description of immature stages, biology). - Berube 1978b- 331 (host specificity).

- P.

Harris 1979: 242 (cost of biological control studies). - Peschken

1979: 455 (biology, hosts). - Shorthouse 1980: 1534 (host, biology). - Bennett 1982: 30, 35 (releases). - Peschken 1984b: 206 (releases, establishment). The wing pattern of this introduced Tephritis species (fig. 429) is distinctive in having a large central hyaline area (fig. 429, a) and an apical hyaline recognition.

crescent interrupted by black markings at the apices of veins R4+5 and M (fig. 429 d), and by a distinct transverse hyaline spot or bar extending through the center of t e pterostigma (fig. 429, b). In these respects the wing pattern is somewhat similar to that of rufipenms (fig. 425), but in the latter, no dark interruption of the apical crescent is present, and the pterostigma is totally dark. The hyaline markings in

Tepbritis dilacerata are more diffuse and mosdy fused rather than appearing as individual hyaline spots. The wing pattern of dilacerata somewhat resembles that of baccbaris (fig. 422), but the dark areas of the wing of the latter are almost completely brown rather than dark gray to black. Map 80. This species is native to northern and central Europe and extends into western parts of the former USSR. distribution.

Sow thistles belonging to the genus Sonchus are the native hosts of di¬ lacerata in the Old World. hosts.

T. dilacerata was introduced in 1979, 1980, and 1981 into eastern and western Canada for the biological control of perennial sow thistle, Sonchus arvensis. As of 1981, it had not established significant populations in North America discussion.

because of its difficulty in overwintering.

Tephritis labecula Foote (Fig. 424; Map 84) Trypanea stigmatica (Coquillett): Malloch (part) 1942: 8 (taxonomy). Tepbritis labecula Foote 1959a: 13 (female holotype, CMP; Grand County, Utah). Foote 1960e: 77 (review). — Foote and Blanc 1963: 69 (review, Calif.). — Foote 1965a: 668 (in catalog).

Map 84. Distribution of Tepbritis labecula and T. stigmatica.

Systematic Treatment of the Genera

The wing pattern of labecula (fig. 424) strikingly resembles that of Trupanea femoralis (fig. 447). The former species is readily distinguished from the recognition.

latter by the presence of two pairs of scutellar bristles, by the presence in most specimens of a dark spot in the base of cell cua,, by the narrower ray connecting the pterostigma with vein r-m, and by the rounded posterior termination of the hyaline spot immediately distad of the pterostigma (fig. 424, a). Although labecula and stigmatica (fig. 423) are the only two species of Tephritis in America north of Mexico having Trupanea-like wing patterns, stigmatica may be distinguished easily from labecula by the lack of a dark bar lying upon vein CuA, and by the extension of the proximal ray through cell dm across vein CuA, into cell cua, (fig. 423, c). T. labecula also has a much narrower ray connecting the pterostigma with vein r-m and a less heavily dark wing pattern. distribution.

Map 84.

T. labecula has been reared from the terminal shoots of Chrysothamnus nauseosus bololeucus. HOST.

Foote (1959a) discussed labecula in detail and commented on its relationships to other species of Tepbritis. discussion.

Tephritis leavittensis Blanc (Figs. 435, 439; Map 82) Tephritis leavittensis Blanc in Foote and Blanc 1979: 173 (female holotype, NMNHLeavitt Lake, Mono County, Calif.). A relatively dark species, leavittensis belongs to the araneosa complex wh'ch contains araneosa, candidipennis, ovatipennis, and signatipennis, all of which it closely resembles in one way or another. T. leavittensis differs from all these RECOGNITION.

species in bearing an anterodorsal row of dark setae on the hind tibia (fig. 439, a). The oviscape of leavittensis is nearly black, very wide, and about 2.0 times as long as the last abdominal segment. The wing pattern resembles those of candidipennis and araneosa, but candidipennis can be distinguished from leavittensis by the much more extensive hyaline areas and the lack of a complete Y-shaped mark at the apex of cell r4+5 (bg- 436, c). T. araneosa differs from leavittensis by the dark band from the pterostigma extending obliquely to cover vein r-m (fig. 437, a) and by the absence of a large hyaline spot anterobasad of vein r-m. distribution. Map 82. Not known. F. L. Blanc swept this species from Arnica diversifolia and reared females from an unidentified California composite. H°ST.

See our discussion of araneosa for additional details concerning the araneosa complex and its members. discussion.

403

Tephritis

Tephritis michiganensis Quisenberry (Fig. 434; Map 85) Tephritis michiganensis Quisenberry 1951: 68 (male holotype, UKL, Birch R., Man.). - Foote 1960e: 81 (review). - Byers et al. 1962: 181 (type data). - Foote 1962: 176 (type data). - Foote 1965a: 668 (in catalog). In wing pattern (fig. 434), michiganensis is similar to pura and it may prove to be indistinguishable from that species when larger samples of the two taxa are studied. The thorax and abdomen of michiganensis are generally more heavily tomentose than those of pura, and the dark scutellum and oviscape exhibit no lateral or apical brownish yellow color, the latter being about 1.2 times as long as recognition.

Systematic Treatment of the Genera

the terminal abdominal tergite. The legs are unmarked yellow-brown, and the body bristles are dark and robust. distribution.

host.

Not

Map 85.

known.

Tephritis ovatipennis Foote (Fig. 438; Map 86) Tephritis ovatipennis Foote 1960e: 81 (female holotype, NMNH; Halfmoon Bay, Calif.). - Foote and Blanc 1963: 69 (review, Calif.). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 135 (hosts). - Arnaud 1979: 332 (type data). The characters given in the key to species distinguish ovatipennis a equately. In araneosa, candidipennis, leavittensis, and ovatipennis, the posterior RECOGNITION.

margin of cell a is hyaline, a character which distinguishes it from signatipennis, another member of the araneosa complex in which this cell is darkened along the posterior wing margin. T. ovatipennis is distinguished from candidipennis by the generally more extensive dark wing markings, and from leavittensis and araneosa by the ovate shape of the hyaline spot in cell r4+5 anterior to vein dm-cu (fig. 438, a). In those two species, this spot is large and broadly joins the spot immediately adjacent to it in cell r2+3. distribution.

Map

86.

The only hosts known for ovatipennis belong to the genera Coretbroeyne and Erigeron (Wasbauer 1972). HOSTS.

T. ovatipennis is one of five species in the araneosa complex, which is discussed in more detail in our treatment of araneosa. discussion.

Tephritis palmeri Jenkins (Map 86) Tephritis palmerii Jenkins 1985: 45 (female holotype, NMNH; Langtry, Tex.) Tephritis new sp.: W. A. Palmer 1987: 192 (host). Tephritis palmeri: Jenkins and Turner 1989: 681 (revision, spp. attacking Baccbaris). T. palmer, closely resembles californica; some morphological characters distinguishing these two tend to overlap. In the former species, the dark areas of the wing pattern are usually darker than in californica (see fig. 428) the recognition.

proximal half of the wing is lighter, the dark areas attaining the posterior margin on y near the apex of vein CuA, or in fragmented spots, and the veins are predomi-

405

Tephritis

Map 86. Distribution of Tephritis ovatipennis, T. palmeri, and T. pura.

nantly pale yellow. In californica, the basal half of the wing is darker and attains the posterior margin at several points, and the veins are distinctly brown. A distinct, broad, crescent-shaped mark is usually present on the frons above the lunule in palmeri, whereas in californica this mark is usually not evident. distribution.

HOSTS.

Map

T. palmeri

86.

has been reared from

flowers and terminal stem galls on

Baccharis halimifolia

B. neglecta.

and from female

Systematic Treatment of the Genera

discussion.

T. palmeri and californica were considered by Jenkins (1985) and

Jenkins and Turner (1989) as being sibling species, each of which give rise to seasonal light and dark morphs. These authors indicate that the morphological differences between the two are consistent, their morphs show fewer differences in key features within than between species, each appear to be monophagous or stenophagous, they attack different species of host plants, and the distribution of the flies is allopatric or at most narrowly sympatric. The previously cited papers by Jenkins and Jenkins and Turner should be consulted for additional information concerning the sympatry and morphs of these two species.

Tephritis pura (Loew) (Fig. 433; Map 86) Trypeta tribulus Say [nomen nudum] in Harris 1835b: 600 (Mass.). - Osten Sacken 1858: 80 (in catalog). - Loew 1862c: 60 (taxonomy). - Loew 1873: 338 (taxon¬ omy). -Johnson 1925a: 97 (synonymy). Trypeta pura Loew 1873: 320 (female holotype, MCZ; Mass.). Euaresta pura: Loew 1873: 322, 330 (taxonomy). - Doane 1899: 192 (distribution). - Aldrich 1905: 613 (in catalog). - Cresson 1907: 105 (taxonomy). - Johnson 1909: 114 (distribution). - Britton 1920: 204 (Conn.). - V. T. Phillips 1923: 147 (review). - Johnson 1925a: 97 (Harris Collection). - Johnson 1925b: 264 (N. Engl.). - Johnson 1927: 218 (Mt. Desert). - Leonard 1928: 853 (N.Y.). - Johnson 1930: 151 (Mass.). - Brimley 1938: 384 (N. Car.). - Quisenberry 1950: 11 (taxonomy). Trypeta (Euaresta) pura: Osten Sacken 1878: 194 (in catalog). Tephritis pura: Strickland 1938: 204 (Alta.). - Hering 1944b: 25 (in key, world). Quisenberry 1951: 71 (review). - Foote 1959a: 17 (taxonomy). - Foote 1960e: 80 (review). - Foote 1965a: 668 (in catalog). In wing pattern (fig. 433), pura resembles michiganensis and webbii, but the two former species can be distinguished from webbii (fig. 432) by the recognition.

much more extensive hyaline aspect of their wings, the narrower arms of the apical Y-shaped marks, and the lack of darkening in cell be. From michiganensis, pura is distinguished by the characters given in the key to species and by the oviscape, which is dark basally with lateral brownish markings. The brownish yellow legs are un¬ marked, and the oviscape is about 1.3 times as long as the terminal abdominal tergite. distribution.

HOST.

Not

Map

86.

known.

Tephritis pura (Loew) would be a secondary junior homonym of Tephritis pura Boheman, 1864 if the latter name, placed by Foote (1984) in the list of discussion.

unrecognized Palearctic species, were to be considered a valid species of Tephritis. Dr.

407

Tephritis P. I. Persson (personal communication) kindly examined the type series in the NRS and discovered that they are Paroxyna plantaginis (Haliday). Tephritis pura (Loew) thus remains the valid name for our species.

Tephritis rufipennis Doane (Fig. 425; Map 82) Tephritis rufipennis Doane 1899: 190 (male lectotype, WSU; Santa Cruz Mts., Calif.). — Aldrich 1905: 612 (in catalog). — Hering 1944b: 18 (in key, world). — Quisenberry 1950: 11 (taxonomy). - Quisenberry 1951: 61 (review). - Foote 1960e: 75 (review). - Foote and Blanc 1963: 70 (review, Calif.). - Foote 1965a: 668 (in catalog). — Foote 1966b: 124 (type designation). - Zack 1984: 33 (type data). - Jenkins 1985: 53 (revision). - Jenkins and Turner 1989: 683 (revision). Euaresta rufipennis: C. F. Baker 1904: 31 (Calif.). - Woodworth 1913: 137 (Calif.). Straussia rufipennis: Woodworth 1913: 137 (Calif.). T rufipennis is light brown with distinctive wing markings (fig. 425) consisting essentially of two transverse dark bands punctuated by numerous small, scattered hyaline spots. These two dark areas are almost completely separated by an extensive central hyaline area extending from the anterior to the posterior borders of the wing (fig. 425, b), sometimes narrowly broken by a dark area along vein M. This pattern somewhat resembles that of baccharis (fig. 422), but the dark areas of the wing of the latter are distinctly brown rather than black; the hyaline areas, except for some spots, are suffused with brown, and the apical light area does not resemble a terminal crescent as in rufipennis (fig. 425, c). The wing of dilacerata (fig. 429) also is similar, but the hyaline crescent in the latter is interrupted by dark marks at the apices of veins R4+5 and M (fig. 429, d), and a distinct transverse hyaline spot or bar is present through the center of the pterostigma (fig. 429, b). recognition.

distribution.

Map 82.

Foote and Blanc (1963) reported a specimen swept from Baccharis pilularis consanguinea, but no actual rearing records are known. Jenkins (1985) and Jenkins and Turner (1989) indicated that rufipennis has been collected in areas close to host.

abundant growths of Baccharis.

Tephritis signatipennis Foote (Fig. 430; Map 82) Tephritis araneosa (Coquillett): Quisenberry (part) 1951: pi. 1, fig. 7 (figure of wing). Tephritis signatipennis Foote 1960e: 77 (female holotype, NMNH; Dardanelles, Tolumne County, Calif.). - Foote and Blanc 1963: 71 (review, Calif.). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 135 (host). - Arnaud 1979: 332 (type data). - Lavigne 1982: 50 (host). - Pemberton et al. 1985: 793 (hosts).

Systematic Treatment of the Genera

One of five species quite similar to araneosa, signatipennis may be recognized by the characters given in the key. It is distinguished from araneosa by the recognition.

darkened posterior margin of cell a (fig. 430, c); the presence in most specimens of a third hyaline spot near the apex of cell r^ and by the longer and darker oviscape, which is almost black and about 1.5 times as long as the terminal abdominal tergite. The darkened posterior margin of the wing in cell a distinguishes this species from candidipennis, leavittensis, and ovatipennis, the other members of the araneosa com¬ plex. distribution.

host.

Map 82.

Mach aeranther a canescens was reported as a host by Wasbauer (1972).

discussion.

See our discussion of araneosa for additional details.

Tephritis stigmatica (Coquillett) (Fig. 423; Map 84) Urellia stigmatica Coquillett 1899c: 266 (male and female syntypes, NMNH; Colo.). - C. F. Adams 1904: 450 (in key). - C. F. Baker 1904: 31 (Nev.). - Aldrich 1905: 614 (in catalog). - Cresson 1907: 106 (N. Mex.). Trypanea stagmatica [error]: Sweet 1930: 123 (plant associations). Trypanea stigmatica: Sweet 1930: 131 (plant associations). - Curran 1932b: 5 (taxonomy). - Malloch (part) 1942: 8 (review). Tephritis stigmatica: Knowlton and Harmston 1937: 145 (Utah). - Quisenberry (review). Foote 1959a: 15 (taxonomy). — Foote 1960e: 76 (review)._ Foote and Blanc 1963: 71 (review, Calif.). — Foote 1965a: 668 (in catalog). — Tauber and Toschi 1965b: 73 (life history, mating behavior). - Novak et al. 1967: 148 (biology, hosts). - Bateman 1972: 504 (biology). - Wasbauer 1972: 135 (hosts). — Goeden 1988: 37 (biology, host). T. stigmatica and labecula are the only two species of Tephritis in America north of Mexico having distinctly Trupanea-like wing patterns (figs. 423, 424). This pattern in stigmatica cannot easily be confused with those of North RECOGNITION.

American Trupanea species, however, and the presence of two pairs of scutellar bristles will definitely place the species in Tephritis. From labecula, stigmatica is distinguished by the much more heavily dark wing pattern, including the much wider dark ray connecting the pterostigma with vein r-m (fig. 423, a), the extension of the two dark rays in cell dm across vein CuAj (fig. 423, b, c), and the presence of an illdefined dark clouding in the base of cell cuaj. distribution.

Map 84.

Several plant species belonging to the genera Arnica, Aster, and Hetero¬ theca have been reported as hosts of stigmatica. Novak et al. (1967) reported it from HOSTS.

409

Tephritis Senecio crassulus, S. douglasii, S. integerrimus, S. pseudaureus, and S. triangularis. Goeden (1988) described the biology and reported that it forms galls as well as feeds in flowers of Senecio douglasii. discussion. Foote (1959a) discussed the relationships of stigmatica and labecula

to similar species of Trupanea.

Tephritis subpura (Johnson) (Figs. 95, 426; Map 83) Euaresta subpura Johnson 1909: 114 (male and female syntypes, MCZ: Wildwood, Anglesea, N.J.). - Johnson 1910: 803 (N.J.). - V. T. Phillips 1923: 147 (review). Trupanea (Tepbritoides) subpura: F. H. Benjamin 1934: 59 (review, Fla.). Tephritis subpura: Hering 1944b: 25 (in key, world). - Quisenberry 1950: 11 (tax¬ onomy). - Quisenberry 1951: 62 (review). - Foote 1959a: 16 (taxonomy). - Foote 1960e: 75 (review). - Foote 1965a: 668 (in catalog). - Wasbauer 1972: 35 (hosts). — Jenkins 1985: 55 (revision, spp. attacking Baccharis). - W. A. Palmer 1987: 192 (host). - Palmer and Bennett 1988: 222 (host). - Jenkins and Turner 1989: 683 (revision, spp. attacking Baccharis). Trupanea subpura: V. T. Phillips 1946: 88, 126 (description, biology larva; hosts). Trypanea subpura: Flennig 1952: 212 (catalog of immature stages). recognition. The ground color and relatively heavy tomentum of the head,

body, and legs of subpura are predominantly yellow without any distinctive dark markings. Most of the major body bristles are light brown; the oviscape, about 1.2 times as long as the preceding abdominal tergite, is slightly darkened at its apex. As indicated in the key to species, the wing of subpura is distinguished by the lack of any darkening in cell c (fig. 426, a), and the preapical brown area is not punctuated by small hyaline spots (fig. 426, b). distribution. Map 83. hosts. T. subpura was reared from terminal stems and flowers of Baccharis

halimifolia from North Carolina to Georgia by Palmer (1987) and Palmer and Bennett (1988) and from that host and from B. glomulifera by Jenkins (1985) and Jenkins and Turner (1989). DISCUSSION. Jenkins (1985) and Jenkins and Turner (1989) discussed in detail

the presence of dark morphs of subpura taken during certain times of the year.

Tephritis webbii Doane (Fig. 432; Map 85) Tephritis webbii Doane 1899: 189 (lectotype, sex not known, WSU; Collins, Ida.). Coquillett 1899c: 266 (taxonomy). - Aldrich 1905: 612 (in catalog). - Hering

410

Systematic Treatment of the Genera 1944b: 21 (in key, world). - Foote 1966b: 126 (lectotype designation) - Zack 1984: 33 (type data). Euaresta webbii: Coquillett 1899c: 266 (taxonomy). - C. F. Baker 1904: 31 (Nev.) Cresson 1907: 105 (description of adult). - V. T. Phillips 1923: 147 (review). Tephritis webbii: Quisenberry 1950: 11 (taxonomy). - Quisenberry 1951: 69 (re¬ view). - Foote 1960e: 80 (review). - Foote and Blanc 1963: 72 (review, Calif.). Foote 1965a: 668 (in catalog). - Wasbauer 1972: 135 (host). T. webbii and pura are similar in that the apical margin of the apicalmost hyaline spot in cell never extends apicad of an anterior extension of vein dm-cu, and the dark ray connecting the pterostigma with vein r-m is very wide recognition.

and lacks hyaline spots centrally (fig. 432, b). For the most part, the hyaline spots in the wing of webbii are relatively small, resulting in a very dark wing pattern. The apical arms of the Y-shaped mark at the wing apex are greatly expanded (fig. 432, c). Most body bristles are black and relatively robust, the tomentum of the thorax and abdomen is brownish yellow, and the legs are brownish yellow with ill-defined darkening on at least the fore femora. The oviscape is very wide basally, about 2.0 times as long as the terminal abdominal tergite, and is black except for some yellow¬ ing laterally at or near the apex. distribution.

host.

Map 85.

A Solidago sp. is the only host plant reported by Wasbauer (1972).

Genus Tomoplagia Coquillett Plagiotoma Loew 1873: 252 (type species, Trypeta obliqua Say 1830: 186, by designation of Coquillett 1910: 591 [preoccupied Dujardin, 1841]). - Williston 1896: 121 (in key). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 605 (in catalog). - Williston 1908: 286 (in key). - Coquillett 1910: 591, 615 (type desig¬ nation). Trypeta (Plagiotoma): Osten Sacken 1878: 190 (in catalog). Tomoplagia Coquillett 1910: 591, 615 (type species, Trypeta obliqua Say 1830: 186, automatic). - Cresson 1914a: 276 (taxonomy). - Flendel 1914b: 88 (in key). Bezzi 1920: 6 (in key). - V. T. Phillips 1923: 121, 132 (in key; review, e. U.S.)'. Curran 1932b: 4 (in key). - F. H. Benjamin 1934: 32 (review, Fla.). - Curran 1934: 291 (in key).-Aczel 1955a: 321 (revision, New World). - Aczel 1955b: 139 (morphology of adults). - Foote and Blanc 1963: 72 (review, Calif.). - Foote 1965a: 678 (in catalog). - Wasbauer 1972: 135 (hosts). - Freidberg 1984: 129 (galls).

Small to medium-sized flies with yellow bodies marked with dis¬ tinct black spots on thorax and abdomen; yellow to light brown body bristles and RECOGNITION.

411

Tomoplagia setae; and yellow, unmarked legs. These flies have three pairs of frontal bristles; two pairs of orbitals; two pairs of scutellars, the apical pair of which are nearly as long as the basal pair; and one pair of dorsocentral bristles situated closer to the transverse suture than to a transverse line through the supra-alar bristles. Both of the species discussed herein have a pair of round black spots at the lateral ends of the posterior three or four abdominal tergites, forming a lateral row of spots on each side of the abdominal tergum. The wings of both species (figs. 440, 441) are distinctive among all U.S. tephritids in bearing yellow and light brown bands that lie at about a 45° angle to the longitudinal axis of the wing, somewhat similar in general appearance to the wing pattern of Peronyma. A total of 45 species has been described from the New World, most of which are included in a revision of the genus by Aczel (1955a). Aczel (1955b) details their morphology. The genus is entirely New World in distribution. Neither of discussion.

the two species treated here are economically important. Tomoplagia has previously been placed in the Trypetinae, but the poorly differenti¬ ated scapular setae, the moderately swollen postocular setae, the dense scutal tomentum, the location of the dorsocentral bristles near the transverse suture, the shape of the male genitalia, and the presence of only two spermathecae indicate that it belongs in the Tephritinae. Few species have been reared, but these breed in flower heads or galls of species of Asteraceae (Liabeae, Mutisieae, or Vernonieae), which also sup¬ ports this classification. Although Tomoplagia species are frequently collected in McPhail traps, the few records of fleshy fruits as hosts (see Aczel 1955a) are doubt¬ ful. Goeden and Headrick (1991a) described the presence of bumplike “verrucae” on the third-instar larva and puparium of cressoni. According to the figure presented by Schwitzgebel and Wilbur (1943), these also appear to be present in obliqua.

Key to U.S. Species of Tomoplagia Apex of scutellum with small, dark spot (figs. 442, a; 443, a); pleuron with only 1 dark spot, situated immediately in front of haltere (fig. 443, b); wing as in fig. 440; southwestern U.S.cressoni Aczel Apex of scutellum without dark spot; pleuron with 3 dark spots (fig. 444, a); wing as in fig. 441; Arizona east to New York and Florida . .obliqua (Say)

Tomoplagia cressoni Aczel (Figs. 86, 440, 442, 443; Map 87) Trypeta (Plagiotoma) obliqua Say: Osten Sacken (part) 1878: 190 (in catalog). Plagiotoma obliqua: Wulp (part) 1899: 405 (in catalog). Tomoplagia obliqua: F. H. Snow 1904: 345 (Ariz.). - Cresson (part) 1907: 100 (taxonomy). - Hendel (part) 1914c: 35 (review, Neotropical Region). - V. T. Phillips (part) 1923: 133 (review). — F. H. Benjamin (part) 1934: 33 (taxonomy). Tomoplagia cressoni Aczel 1955a: 347 (male holotype, NMNH; Ontario, Calif.). -

412

Systematic Treatment of the Genera

Figures 440, 441. Right wings, Tomoplagia spp. 440, cressoni Aczel; 441, obliqua (Say). Figure 442. Dorsal view of scutellum, Tomoplagia cressoni Aczel. Figures 443, 444. Lateral view of thorax Tomoplagia spp. 443, cressoni Aczel; 444, obliqua (Say).

Aczel 1955b: 139, 169 (description, in key). - Foote and Blanc 1963: 72 (review, Calif.). - Wasbauer 1964: 8 (review). - Foote 1965a: 678 (in catalog). - Wasbauer 1972: 135 (hosts). - Goeden and Ricker 1989: 329 (host). - Goeden and Headrick 1991a: 549 (biology, hosts, immature stages, parasites). The wing patterns and general morphology of cressoni and obliq¬ ua are so similar that they are of little use in species identification. T. cressoni is RECOGNITION.

recognized by the presence of only a single rounded black spot on the pleuron immediately anterior to the haltere (fig. 443, b) and a very small black mark at the apex of the scutellum. These markings are in contrast to those of obliqua, which displays three dark pleural markings (fig. 444, a) and no scutellar dark spot whatever. distribution.

Map 87.

413

Tomoplagia

Map 87. Distribution of Tomoplagia cressoni, T. obliqua, and Toxotrypana curvicauda.

This species has been reared from flower heads of Perezia microcepbala and Trixis californica (Mutisieae) (Foote and Blanc 1963, Goeden and Ricker 1989, Goeden and Headrick 1991a). A record of Prunus persica (Aczel 1955a) appears to hosts.

be doubtful (Wasbauer 1972).

Tomoplagia obliqua (Say) (Figs. 441, 444; Map 87) Trypeta obliqua Say 1830: 186 (types probably lost; Ind.). — Osten Sacken 1858: 79 (in catalog). - LeConte 1859: 370 (taxonomy). - Loew 1862c: 59 (taxonomy). Osten Sacken 1862: 97, 99 (description of adult). - Loew 1873: 251 (review). Beach 1895: 94 (parasite). - Coquillett 1910: 591 (type species designation). Plagiotoma obliqua: Loew 1873: 252, 329, 337 (taxonomy). — W. A. Snow 1894: 162 (distribution). - Doane 1899: 179 (distribution). - Wulp (part) 1899: 405 (in catalog). - Johnson 1900b: 687 (N.J.). - F. H. Snow 1903: 219 (Kans.). - Aldrich 1905: 605 (in catalog). - Johnson 1910: 801 (N.J.). Trypeta (Plagiotoma) obliqua: Osten Sacken (part) 1878: 190 (in catalog). Tomoplagia obliqua: Coquillett 1910: 591 (type species designation). - Hendel 1914b: 88 (type data). - Hendel 1914c: 35 (review, Neotropical Region). - V. T. Phillips (part) 1923: 133 (review). - F. H. Benjamin (part) 1934: 33 (review Fla.). - Brimley 1938: 383 (N. Car.). - Schwitzgebel and Wilbur 1943: 4 (biology). V. T. Phillips 1946: 124 (host). - Aczel 1949: 241 (distribution). - Aczel 1955b:

414

Systematic Treatment of the Genera

139, 167 (description of adult, in key). - Steyskal 1957: 94 (eye color). - Foote 1965a: 678 (in catalog). - Novak et al. 1967: 148 (host, biology). - Wasbauer 1972: 136 (hosts). T. obliqua may be distinguished easily from cressoni by the nature of the black spotting of the thorax. Although both species possess a dark mark on the recognition.

thoracic pleuron immediately anterior to the haltere (figs. 443, b; 444, a), obliqua also bears two additional spots, one on the katepisternum and one on the katepimeron (fig. 444, a); these two dark spots are entirely absent in cressoni. The dark marking at the apex of the scutellum of cressoni is not present in obliqua. distribution.

Map 87.

An unknown species of Aster and at least six species of Vernonia were recorded as hosts by Wasbauer (1972). The former record is questionable. HOSTS.

Genus Toxotrypana Gerstaecker Toxotrypana Gerstaecker 1860: 191 (type species, Toxotrypana curvicauda Gerstaecker 1860: 194, by monotypy). - Loew 1873: 27, 34, 36 (taxonomy). Osten Sacken 1878: 181 (in catalog). - W. A. Snow 1895: 117 (taxonomy). Williston 1896: 120 (in key). - Aldrich 1905: 600 (in catalog). - Coquillett 1910: 615 (type data). - Cresson 1914a: 275 (taxonomy). - Hendel 1914b: 74 (in key). F. H. Benjamin 1934: 10 (review, Fla.). - Foote 1965a: 659 (in catalog). Wasbauer 1972: 136 (hosts). - Foote 1980: 48 (review). - Burditt and Baranowski 1982: 85 (review). - Norrbom 1985: 57 (taxonomy). - Flancock 1986b: 275 (classification). Mikimyia Bigot 1884: xxix (type species, Mikimyia furcifera Bigot 1884: xxix, by monotypy) [= curvicauda Gerstaecker]). - Coquillett 1910: 570 (type data). Toxytrypana [error]: Williston 1908: 283 (in key). Toxotrypanea [emendation]: Curran 1932b: 2 (in key). - Curran 1934: 293 (in key). Members of Toxotrypana are large brownish flies with the follow¬ ing combination of characters that distinguishes them from members of all other genera occurring in America north of Mexico: female with a greatly elongated RECOGNITION.

oviscape, downwardly curved except in a few species (figs. 15, a; 16, a); the abdomen

Figure 445. Right wing, Toxotrypana curvicauda Gerst.

415

Toxotrypana of both sexes relatively slender and constricted basally (fig. 15); frontal bristles weak or absent; antenna longer than the face with the arista pubescent; scutum with a medial longitudinal furrow; scutellar bristles situated apicad of the middle of the scutellum; wing (fig. 445) long, narrow, and hyaline with a broad, light brown costal band and vein R2+3 distinctly sinuate, usually with at least one small anteriorly directed accessory vein (fig. 445, a). The head and thoracic bristles in this genus are reduced in size, especially those on the thorax. In some of these characters, Toxo¬ trypana remotely resembles Dacus and Bactrocera, but the long oviscape and the other characters given in the key to genera will serve to distinguish it adequately.

Toxotrypana is restricted to the New World, where a number of described and undescribed species occur south to Argentina. T. curvicauda is the only one of these found in the region covered by this handbook, although at least one discussion.

other species with a close resemblance occurs in Mexico. This genus has long been thought to be closely related to Dacus, but morphological studies by Norrbom (1985) and Hancock (1986b) and biochemical studies by Sarma et al. (1987) have shown that the true affinities of Toxotrypana actually lie with Anastrepha rather than within the subfamily Dacinae.

Toxotrypana curvicauda Gerstaecker Papaya fruit fly (Figs. 15, 16, 445; Map 87) Toxotrypana curvicauda Gerstaecker 1860: 194 (female syntypes, ZMHU; St. John, Virgin Is.). - W. A. Snow 1894: 159 (taxonomy). - Aldrich 1905: 600 (in catalog). - Coquillett 1910: 570 (type data). - Hendel 1914c: 10 (review). - Knab and Yothers 1914: 447 (review). - Anon. 1915: 60 (biology). - Mason 1922a: 1 (biology). - Mason 1922b: 6 (review). - C. T. Greene 1929: 491 (description larva, pupa). - Bates 1933b: 160 (hosts). - Bates 1934b: 1 (morphology of larva). - F. H. Benjamin 1934: 11 (review, Fla.). - Curran 1934: 286 (illustration of female). - A. C. Baker et al. 1944: 142 (review, biology, hosts). - McClanahan 1946: 29 (biology). - Merrill 1946: 39 (host). - V. T. Phillips 1946: 81, 124 (description, biology larva; host). - Aczel 1949: 180 (in catalog). - Butcher 1952: 196 (host). - Hennig 1952: 209 (in catalog immature stages). - Peterson 1953: 333 (figures of larva). - Hardy 1955: 426, 437 (taxonomy). - Christenson and Foote 1960: 171 (biology). - Foote 1965a: 659 (in catalog). - Brogdon and Wolfenbarger 1968: 1 (review). - Weems 1969a: 1 (review). - Wasbauer 1972: 136 (hosts). - Wolfenbarger and Walker 1974: 384 (biology). - Wharton and Marsh 1978: 149 (parasite). - Berg 1979: 15 (in key to larvae). - Landolt and Hendricks 1983: 413 (reproductive behavior). Landolt 1984a. 285 (mating). Landolt 1984b: 240 (biology). - Landolt 1984c: 215 (behavior). - Sharp and Landolt 1984: 176 (biology). - Landolt 1985: 354 (biology). - Landolt et al. 1985: 751 (pheromones). - Norrbom 1985: 57 (taxonomy). - Heppner 1986: 1 (description of larva).

416

Systematic Treatment of the Genera

Mikimyia furcifera Bigot 1884: xxix (male holotype, UMO; Brazil [probably er¬ roneous]). - W. A. Snow 1894: 159 (synonymy). - Coquillett 1910: 570 (type data). recognition.

As

for

the genus.

Map 87. T. curvicauda is widespread in the New World, occur¬ ring in the Bahamas, the West Indies, and in the United States and Mexico south to Colombia and Venezuela. distribution.

T. curvicauda is the most destructive pest known to commercial growers of papaya in the New World; both cultivated and wild fruits are attacked. Several hosts.

species of Gonolobus (Asclepiadaceae), including that reported as “talayote” by Baker et al. (1944), are hosts in Mexico and Guatemala (Castrejon 1987; A. L. Norrbom, unpublished data), as is Jacaratia mexicana (Castrejon 1987). Bates (1933b) reported a host belonging to Apocynaceae, probably a species of Tabernaemontana. It has also been reported to attack mango, especially when populations are unusually high.

discussion.

T. curvicauda was introduced into Florida about 1905 (Weems

1969a). Heppner (1986) described and illustrated the larva in considerable detail. The synonymy given for this species presents the source of a considerable amount of information about its biology.

Genus Trupanea Schrank Trupanea Guettard 1762: 171 (unavailable name; author not binominal). Trupanea Schrank 1795: 147 (type species, Trupanea radiata Schrank 1795: 147, by monotypy (= stellata (Fuessly)). - Coquillett 1899c: 259 (nomenclature)5. Coquillett 1910: 618 (type data). - Cresson 1914a: 276, 278 (taxonomy). - F. H. Benjamin (part) 1934: 52 (taxonomy). - Curran 1934: 293 (in key). - Foote 1958: 290 (taxonomy). - Foote 1959a: 16 (taxonomy). - Foote 1960e: 72 (taxonomy). Foote 1960f: 146 (taxonomy). - Foote 1960g: 1 (revision). - Foote and Blanc 1963: 73 (review, Calif.). - Munro 1964a: 4 (taxonomy, Afrotropical). - Foote 1965a: 666 (in catalog). - Wasbauer 1972: 136 (hosts). - Freidberg 1984: 129 (galls). — Goeden 1992: 107 (hosts). Urellm Robineau-Desvoidy 1830: 774 (type species, U re Ilia calcitrapae RobineauDesvoidy 1830: 774, by subsequent designation of Coquillett 1910- 618) ( = stellata (Fuessly)). - Loew (part) 1873: 324, 328 (taxonomy). - Williston (part) 1896: 123 (in key). - Coquillett 1899c: 268 (in key). - C. F. Adams (part) 1904: 450 (in key). - Aldrich (part) 1905: 613 (in catalog). - Williston (part) 1908: 287 (in key). - Curran 1932b: 5 (synonymy). Trypeta (Urellia): Osten Sacken (part) 1878: 194 (in catalog).

417

Trupanea Trypanea [emendation]: Cresson 1914a: 279 (nomenclature). - Hendel 1914b: 97 (in key). - V. T. Phillips (part) 1923: 123, 147 (in key, review). - Curran 1932b: 3, 5 (in key). - Malloch (part) 1942: 1 (revision). The genus Trupanea comprises small flies with yellowish head and legs and grayish or brownish tomentum covering the thorax, scutellum, and abdomen. The wing patterns of most species are characteristic in that the hyaline areas are very light to white, contrasting strongly with the usually very dark brown to almost black markings which consist of a prominent dark area in the anterior distal quarter of the disk, from which rather narrow dark rays extend to the wing border. All Nearctic species have three pairs of frontal, two pairs of orbital, and one pair of scutellar bristles. The dorsocentral bristles are situated very close to the transverse suture and are distantly removed from the supra-alar bristles. Sexual dimorphism is evident in a number of species; the wing pattern of males generally have less extensive dark markings, and many of the characteristic dark rays are interrupted or missing, and males of several species have outstanding dark anterodorsal setae on the middle femur. In the Nearctic region, Trupanea is the only genus in which males have a large basal spinelike sclerite on the distiphallus. Although the genus is quite distinctive and should not be mistaken for any other occurring in the United States and Canada, several of its species may be confused with a few belonging to Tephritis. In such cases, recognition.

the presence of only one pair of scutellar bristles always indicates Trupanea. Species of Trupanea are difficult to identify because of the large amount of variation in wing pattern, the only characters that have been studied to discussion.

any extent in the U.S. and Canadian species of this genus; the user should approach identification at the species level with this in mind. Separate keys to males and females have been provided because of the relatively large amount of sexual dimor¬ phism present in Trupanea, a situation not common to other tephritid genera in the United States and Canada. The genus Trupanea occurs in all the major zoogeographical regions of the world; nearly 200 species are known worldwide, and possibly as many more remain to be described. Goeden (1992) discussed the host relationships of a number of species that occur in California. The following key, based almost exclusively on wing pattern, will assist the user in grouping species and using the more detailed keys to species which follows it. How¬ ever, the groupings shown in the following key are not intended to represent formal “species groups” in the taxonomic sense.

Key to Species Groups of U.S. and Canadian Trupanea Based on Wing Pattern 1. Dark band connecting pterostigma with vein r-m, if present, as wide or nearly as wide as pterostigma (fig. 447, a) ..... .femoralis, nigricornis, bisetosa

418

Systematic Treatment of the Genera

Dark band connecting pterostigma with vein r-m distinctly narrower than length of pterostigma (fig. 453, a), sometimes broken or absent.2 2. Two dark rays extending into or through cell dm from vein M (fig. 453, b, c) ..

Only 1 dark ray extending into cell dm from vein M (fig. 457, b) .4 3. Distal dark ray in cell dm crossing vein CuAj (fig. 453, b) . .dacetoptera, arizonensis, maculigera Distal dark ray in cell dm ending at vein CuA1? never extending beyond that vein (fig. 457, b).wheeleri, mevarna, ageratae, conjuncta, texana 4. Hyaline spot at extreme base of cell r4+5 present and distinct (fig. 468, d) .jonesi, pseudovicina, radifera, imperfecta, signata, eclipta Hyaline spot at extreme base of cell r4+5 indistinct and without sharp margins, clouded, or absent (fig. 471, c) . .actinobola, californica, vicina, viciniformis

Key to U.S. and Canadian Species of Trupanea Females 1. Unbroken dark brown area present in anterior apical quarter of wing (fig. 446, a); apex of cell r4+5 entirely black (fig. 446, b) . .conjuncta (Adams) Wing with preapical dark area in apical third or half of cell r2 + 3 and basal half of cell r4+5 with narrow dark rays diverging from it to wing margins (fig. 447); apex of cell r445 largely hyaline (fig. 447, c) .2 2. Vein r-m connected to pterostigma by a dark ray as wide or nearly as wide as length of pterostigma (fig. 447, a) .3 Dark ray connecting pterostigma with vein r-m distinctly narrower than length of pterostigma (fig. 453, a), sometimes partially or completely absent (fig. 464, a).. 3. Two dark rays partially or entirely crossing cell dm (fig. 447, d); basal V2 or more of vein CuAj infuscated (fig. 447, e) .femoralis (Thomson) Only 1 dark ray, if any, crossing cell dm, sometimes incomplete (fig. 451, c); vein CuAj with a dark line or spot near its center (fig. 452, b), or entirely devoid of markings (fig. 451, d) .4 4. Apicalmost hyaline spot in cell r2+3 usually extending broadly across vein r4+5> giving the appearance of a distinct stem for the apical Y-shaped dark mark (fig. 449, a); width of stem about equal to width of posterior apical arm .nigricornis (Coquillett) Apicalmost hyaline spot in cell r2+3 usually not extending broadly across vein R4+5, tending to obliterate the appearance of a distinct slender stem for the apical Y-shaped mark (fig. 451, a); width of stem about 2 times as wide as posterior arm.bisetosa (Coquillett) 5. Two dark rays extending through cell dm (fig. 455, a), the proximal one sometimes incomplete in posterior half of cell and not attaining vein CuAj, or broken in center of cell; in addition, sometimes a spot present near the middle of that cell and another spot in line with it on vein CuAj

a

b

a

b

Figures 446-455. Right wings, Trupanea spp. 446, conjuncta (Adams), female; 447, femoralis (Thoms.), female; 448, femoralis (Thoms.), male; 449, nigricornis (Coq.), female; 450, nigricornis (Coq.), male; 451, bisetosa (Coq.), female; 452, bisetosa (Coq.)., male; 453, dacetoptera Phil., female; 454, arizonensis Mall., female; 455, maculigera Foote, female.

419

Figures 456-465. Right wings, Trupanea spp. 456, ageratae Benj., male; 457, wheeleri Curr female; 458, mevarna (Walk.), female; 459, texana Mall., female; 460, imperfecta (Coq.), female 461, imperfecta (Coq.), male; 462, radifera (Coq.), female; 463, radifera (Coq.), male; 464, eclipta Benj., remale; 465, jonesi Curr., female.

420

421

Trupanea

(fig. 454, a), or with other markings within cell proximal to distal ray

.6 One dark ray extending through cell dm (fig. 460, b) or entirely lacking

.11 . Distal ray through cell dm continuing to hind margin of wing (fig. 453, b); if broken in cell cual5 its continuation always marked by a dark spot at hind wing margin .7 Distal ray through cell dm extending only to vein CuA1? never beyond (fig. 457, b) .9

6

7. Proximal ray present in cell dm terminating at middle of cell (fig. 453, c); vein CuAj never with a dark spot in line with it . .dacetoptera Phillips Proximal ray extending completely across cell dm; if broken in that cell, always at least a dark spot present on vein CuAj in line with it (fig. 459, c) .8 . Distance along vein CuAj between proximal and distal rays in cell dm greater than distance between distal ray and vein dm-cu (fig. 454, b); dark mark on vein CuAj situated at middle of that vein (fig. 454, a) .. .arizonensis Malloch Distance along vein CuAj between proximal and distal rays equal to or less than the distance between distal ray and vein dm-cu (fig. 455, b); dark marks on vein CuAj distinctly apicad of middle of that vein (fig. 455, a) .maculigera Foote

8

9. Elongate dark streak extending from base to middle of vein CuAa (fig. 457, c), or beyond .wheeled Curran (part) Base of vein CuA! entirely without infuscation (fig. 458, d), but a dark spot may be present near middle of that vein (fig. 459, c) .10 10. Marginal light spot at apex of vein R2+3 centered in surrounding dark area (fig. 458, b); hyaline spot immediately distad of pterostigma narrowly rounded apically (fig. 458, a).mevarna (Walker) (part) This marginal light spot closer to tip of wing in surrounding dark area (fig. 459, b); hyaline spot distad of pterostigma broadly rounded apically (fig. 459? a).texana Malloch 11. Hyaline spot immediately anterior to vein dm-cu quadrate and attaining vein R4+5 along most of its width (fig. 460, a) imperfecta (Coquillett) Hyaline spot immediately anterior to vein dm-cu rounded anteriorly and rarely exceeding posterior 2/3 of cell r4+5 (fig. 462, c).12 12. Apex of cell br usually completely hyaline except for an occasional very narrow band of infuscation bordering vein r-m (fig. 462, b); dark infusca¬ tion on vein CuAj extending nearly to extreme base of that vein (fig. 465, b) .^ Apex of cell br definitely infuscated in addition to narrow band along vein r-m (fig. 471, a); dark infuscation on vein CuAl5 if present, confined to center of that vein (fig. 471, d) .19 13. Dark ray from pterostigma to vein r-m distinctly broken in either cell r1 or cell r2+3 (fig. 462, a), or completely absent ..14 Dark ray from pterostigma to vein r-m complete in cells and r2+3 (fig. 465, a), although it may be broken within the pterostigma (fig. 469, a)

422

Systematic Treatment of the Genera

14. Dark ray extending from pterostigma to vein r-m broken in cell r, and complete in cell r2 + 3 (fig. 462, a, b), or completely absent . .radifera (Coquillett) Dark ray extending from pterostigma to vein r-m complete in cell r, and broken in cell r2+3 (fig. 464, b) .eclipta Benjamin 15. Infuscation at center of vein CuAj at least 3 times as long as wide, never extending to extreme base of that vein (fig. 465, b).jonesi Curran Infuscation at center of vein CuAt subquadrate, triangular (fig. 471, d), or completely absent (fig. 464, d) .16 16. Marginal hyaline spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray about as wide as that covering vein dm-cu (fig. 467, c); western U.S.californica Malloch Marginal hyaline spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray 2 times or more as wide as that covering vein dm-cu (fig. 468, c) .17 17. Proximal light spot in cell r4+5 distinct, broadly based on vein M and extending about 2/3 of way across cell r4+5 (fig. 468, d); infuscation at center of vein CuAj triangular (fig. 468, e) .pseudovicina Hering Proximal light spot in cell r4+5 lacking (fig. 471, c), indistinct, or extending entirely across cell (fig. 469, c), never broadly based on vein M_18 18. In profile, head distinctly higher than long (fig. 473) . ..signata Foote (part) In profile, head at least as high as long or longer (fig. 474). .actinobola (Loew) (part) 19. Apex of cell br usually with dark, even infuscation similar to that in base of cell r4+5 (figs. 471, a; 472, a) .20 Apex of cell br usually with infuscation either in distinct spot surrounded at least in part apically by a lighter hyaline area, or broad and lightly areolate, its proximal border never sharply delimited (fig. 469, b).21 20. Proximal border of dark area in apex of cell br sharply delimited (fig. 471, a); dark subapical mark present through cell dm to vein CuA^ usually a dark mark on vein CuA1 near center of cell dm (fig. 471, d) . • ..vicina (Wulp) Proximal borders of dark area in apex of cell br and bar through pterostig¬ ma to vein r-m shaded gradually into the adjoining hyaline areas (fig. 472, a); cell dm without subapical bar; no dark marking present at center of vein CuAj.viciniformis Foote 21. Vein CuAt with elongated infuscated streak from its base to its midpoint or somewhat beyond (fig. 457, c) .wheeleri Curran (part) Infuscation on vein CuA| in form of a spot (fig. 466, d) or entirely lacking (fig. 462, d).22 22. Brownish species with the hyaline area immediately distad of pterostigma narrowly rounded apically (fig. 458, a) .mevarna (Walker) (part) Grayish species with the hyaline area at apex of pterostigma rounded broad¬ ly posteriorly (fig. 467, a) or ending broadly along vein R4+5 (fig. 470, a) .23 23. Infuscation in apex of cell br, if present, with distinct margins (fig. 469, b) .. • ..signata Foote (part) Infuscation in apex of cell br, if present, indistinctly margined .24

Figures 466-472. Right wings, Trupanea spp. 466, jonesi Curr., male; 467, californica Mall., female; 468, pseudovicina Her., female; 469, signata Foote, female; 470, actinobola (Lw.), female; 471, vicina (Wulp), female; 472, viciniformis Foote, female. Figures 473, 474. Side view of head, Trupanea spp. 473, signata Foote; 474, actinobola (Lw.).

423

424

Systematic Treatment of the Genera

24. Marginal hyaline spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray about as wide as that covering vein dm-cu (fig. 467, c); western U.S.californica Malloch (part) Marginal hyaline spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray 2 times or more as wide as that covering vein dm-cu (fig. 470, c); entire U.S.actinobola (Loew) (part)

Males 1. An unbroken dark brown area present in anterior apical quarter of wing disk (fig. 446, a); apex of cell r4+5 entirely black (fig. 446, b) .

.conjuncta (Adams) Wing with a preapical dark area in apical half or third of cell r4 + 5 with rays diverging from it to margins (fig. 447); apex of cell r4+5 largely hyaline (fig. 447, c) .. 2. Dark ray connecting pterostigma with vein r-m as wide or nearly as wide as length of pterostigma (fig. 448, a).3 Dark ray connecting pterostigma with vein r-m much narrower than length of pterostigma (fig. 453, a), sometimes broken or completely absent (fig. 463, a) . 3 3. Two dark rays partially (fig. 448, b) or entirely (fig. 447, d) crossing cell dm; vein CuAj infuscated along most of its length (fig. 448, c)

.femoralis (Thomson) Only 1 dark ray crossing cell dm (fig. 450, b), sometimes incomplete; vein CuAj with dark spot near center or entirely devoid of markings .... 4 4. First flagellomere dark brown to black; apical Y-shaped mark incomplete (fig. 450, a); vein CuAj usually with dark spot at center (fig. 450, c)

.nigricomis (Coquillett) First flagellomere yellow, about the same color as pedicel; apical Y-shaped mark complete (fig. 452, a); vein CuAj with a dark spot at center (fig. 452, b) or devoid of markings .bisetosa (Coquillett) 5. Two dark rays extending through cell dm (fig. 455, a), the proximal one sometimes incomplete in posterior half of that cell and not attaining vein CuAi..

One dark ray extending partially or entirely through cell dm (fig. 457, b) or entirely lacking; in addition, sometimes a spot present in middle of that cell and a dark spot in line with it on vein CuAj (fig. 454, a) .12 6. Distal ray through cell dm continuing to hind margin of wing (fig. 453, b); if

broken in cell cual5 its continuation always marked by a dark spot at hind margin of wing . 7 Distal ray through cell dm extending only to vein CuAi and never bevond (fig. 457, b) .... 9 7. Proximal ray in cell dm ending at middle of cell (fig. 453, c); vein CuAl never with a dark spot in line with it.dacetoptera Phillips Proximal ray extending completely across cell dm, or if broken in that cell, always at least a dark spot present on vein CuAi in line with it (fie 459, c) ... 8. Distance along vein CuAj between proximal and distal rays in cell dm

425

Trupanea

greater than distance between distal ray and vein dm-cu (fig. 454, b); dark mark on vein CuAj situated at middle of that vein (fig. 454, a) .. .arizonensis Malloch Distance along vein CuAj between proximal and distal rays in cell dm equal to or less than the distance between the distal ray and vein dm-cu (fig. 455, b): dark mark on vein CuAj distinctly apicad of middle of that vein .maculigera Foote 9. Elongate dark streak extending from base to middle of vein CuAl (fig. 457, b) or beyond .wheeleri Curran (part) Base of vein CuAj entirely without infuscation (fig. 458, d), but a dark spot may be present at middle of that vein (fig. 459, c).10 10. Hyaline riorly Hyaline riorly

area immediately distad of pterostigma narrowly rounded poste¬ (fig. 458, a).mevarna (Walker) (part) area immediately distad of pterostigma broadly rounded poste¬ (fig. 456, a) or ending broadly on vein R4+5 (fig. 457, a)_11

11. Marginal hyaline spot near apex of vein R2+3 situated slightly apicad of center of dark area surrounding it (fig. 456, b) _ageratae Benjamin Marginal hyaline spot near apex of vein R2-)-3 situated distinctly closer to apical than to proximal margin of dark area surrounding it (fig. 459, b) .texana Malloch 12. Apex of cell br completely hyaline except for an occasional narrow band of infuscation bordering vein r-m (fig. 462, b) .13 Apex of cell br more broadly infuscated in addition to narrow band border¬ ing vein r-m (fig. 471, a).20 13. Apical Y-shaped mark incomplete or absent (fig. 461, b).14 Apical Y-shaped mark usually complete or nearly so (fig. 464, c) -16 14. Dark ray extending from pterostigma to vein r-m complete in cells and r2+3 (fig- 461, a).imperfecta (Coquillett) Dark ray extending from pterostigma to vein r-m usually distinctly broken or completely absent in cells rj and r2+3 (figs. 463, a; 466, a).15 15. Light spot immediately anterior to vein dm-cu large and occupying at least half of width of cell r4+5 (fig. 462, c); dark area in apex of cell rl barely longer than adjacent hyaline spot at apex of vein R2+3 (fig. 463, b); vein CuAt without infuscation or with a dark spot (fig. 463, d). .radifera (Coquillett) Light spot immediately anterior to vein dm-cu small and occupying dis¬ tinctly less than posterior half of cell r4+5 (fig. 466, c); dark area in apex of cell r, more than twice as long as adjacent hyaline spot (fig. 466, b); vein CuA, with an elongate infuscation at its central 1/3 (fig. 466, d) .jonesi Curran 16. Dark ray from pterostigma to vein r-m broken or absent in cell r2+3 (fig. 464, a); usually no spot on vein CuAj (fig. 464, d) . .eclipta Benjamin Dark ray from pterostigma to vein r-m complete in cells and r2+3 (fig. 467, d); usually a dark spot present on vein CuAt (fig. 471, d)-17 17. Proximal light spot in cell r4+5 broadly based on vein M and extending about 2/3 of way across cell r4+5 (fig. 468, d) ... pseudovicina Hering Proximal light spot in cell r4+5 lacking (fig. 471, c), indistinct, or extending entirely across cell (fig. 469, c), never broadly based on vein M-18

Systematic Treatment of the Genera

18. Marginal hyaline spot near apex of vein R2+3 situated very close to distal margin of dark area surrounding it (fig. 467, b, c) . .californica Malloch (part) Marginal hyaline spot near apex of vein R2 + 3 situated far from distal mar¬ gin of dark area surrounding it (fig. 468, b) .19 19. In profile, head distinctly higher than long (fig. 473) . .signata Foote (part) In profile, head subquadrate or longer than high (fig. 474) .actinobola (Loew) (part) 20. Apex of cell br with dark, even infuscation similar to that in base of cell r4 + (figs. 471, a; 472, a) .21 Apex of cell br with infuscation either in a distinct spot surrounded at least in part apically by a lighter or hyaline area, or broad and lightly areolate, the proximal border sharply delimited (fig. 469, b).22 21. Proximal border of dark area in apex of cell br sharply delimited (fig. 471, a); dark subapical mark present through cell dm to vein CuA3; usually a dark mark on vein CuAj near center of cell dm.vicina (Wulp) Proximal borders of dark area in apex of cell br and bar through pterostigma to vein r-m shading gradually into the adjoining hyaline areas (fig. 472, a); cell dm without subapical bar (fig. 472, b); no dark marking present at center of vein CuA1 .viciniformis Foote 22. Vein CuAj with elongate infuscated streak extending from at or near its base to its midpoint or slightly beyond (fig. 457, c) . .wheeleri Curran (part) Infuscation on vein CuA1 in the form of a spot (fig. 460, c), or entirely lacking . 23 23. Brownish species with the hyaline area at apex of pterostigma narrowly rounded apically (fig. 458, a).mevarna (Walker) (part) Grayish species with hyaline area at apex of pterostigma broadly rounded posteriorly (fig. 471, b) or ending broadly along vein R4+5 .24 24. Infuscation in apex of cell br with distinct margin (fig. 469, b); in profile, head distinctly higher than long (fig. 473) .signata (Foote) (part) Infuscation in apex of cell br indistinctly margined; in profile, head subqua¬ drate or longer than high (fig. 474) . 25 25. Marginal hyaline spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray about as wide as that covering vein dm-cu (fig. 467, c); western U.S.californica Malloch (part) Marginal spot at apex of vein R2+3 separated from next distal hyaline area by a dark ray nearly 2 times as wide as that covering vein dm-cu (fig. 470, c); entire U.S.actinobola (Loew) (part) Trupanea actinobola (Loew) (Figs. 470, 474; Map 88) Trypeta actinobola Loew 1873: 326 (male lectotype, MCZ; Tex.). — Foote 1960g: 24 (lectotype designation). Urellia actinobola: Loew 1873: 326, 330 (taxonomy). — Coquillett 1899c:266 syn¬ onymy). - F. H. Snow 1903: 219 (Kans.). - C. F. Adams 1904: 450 (in key). Aldrich 1905: 613 (in catalog). - Tucker 1907: 105 (Kans., Colo.). Trypeta (Urellia) actinobola: Osten Sacken 1878: 194 (in catalog).

427

Trupanea

Tephritis daphne (Wiedemann) [misidentification]: Sweet 1930: 123 (plant associa¬ tions). - Knowlton and Cutler (part) 1932:

111 (Utah). - Knowlton and

Harmston 1937: 145 (Utah). Trypanea actinobola: Curran 1932b: 6 (in key). — Brimley 1938: 384 (N. Car.). — Malloch 1942: 16 (review). - Henmg 1952: 212 (in catalog immature stages). Trupanea (Trupanea) actinobola: F. H. Benjamin 1934: 56 (review, Fla.). Trupanea actinobola: V. T. Phillips 1946: 85 (description, biology larva; host). Foote 1960g: 24 (revision). - Foote and Blanc 1963: 75 (review, Calif.). - Munro 1964: 73 (taxonomy). - Foote 1965a: 667 (in catalog). - Stegmaier 1968b: 48 (biology, hosts). - Stegmaier 1968c: 95 (biology). - Wasbauer 1972: 136 (hosts). Goeden 1985: 565 (hosts). - Goeden and Ricker 1989: 329 (host). - Goeden and Headrick 1991b: 553 (host). — Goeden 1992: 108 (hosts). Trupanea actinibola [error]: V. T. Phillips 1946: 125 (hosts). recognition. The wing pattern of actinobola resembles those of several other species of Trupanea in that the dark ray connecting the pterostigma with vein r-m is narrower than the length of the pterostigma, only one dark ray extends posteriorly from vein A4 into cell dm, the apex of cell br is without dark markings, and the extreme base of cell r4+5 lacks a well-defined hyaline area. It may be distinguished from all those species by the shape of the head in lateral view; in the former, the head is distinctly subquadrate or longer than high, in the latter higher than long. The small dark spot at the center of vein CuA^ is indistinct or lacking, and the hyaline area at the extreme base of cell r4+5 is variable in extent. T. actinobola is distin¬ guished from californica, which it most resembles, in the position of the hyaline spot

428

Systematic Treatment of the Genera at the apex of vein R2+3 (fig. 470, b), which is located distinctly more basad in the dark spot surrounding it than in the latter species. It is a relatively variable species in wing pattern. There appears to be little or no sexual dimorphism, and males resem¬ ble females in every important respect. distribution.

HOSTS.

Map

88.

At least 20 plant species belonging to the following eight genera were

listed by Wasbauer (1972): Actinospermum, Aster, Coreopsis, Erigeron, Gnaphaliurn, blieracium, Sitilias, and Solidago. T. actinobola has been reared from Trixis californica in southern California (Goeden and Ricker 1989, Goeden and Headrick 1991b). Additional plant genera added by Goeden (1992) are Ampbipappus, Brickellia, Haplopappus, and Trichoptilium. It is possible that biological studies will demonstrate the existence of several discrete populations of actinobola that are not now separable mor¬ DISCUSSION.

phologically. This species may well extend into Mexico and even farther south in the New World.

Trupanea ageratae Benjamin (Fig. 456; Map 89) Trupanea (Trupanea) ageratae Benjamin 1934: 56 (male holotype, NMNH; Ferry Landing, No Name Key, Fla.). Trypanea ageratae: Malloch 1942: 8 (revision). - V. T. Phillips 1946: 25 (host). Trupanea ageratae: Foote 1960g: 14 (revision). - Foote 1965a: 667 (in catalog). Wasbauer 1972: 136 (host). In wing pattern, ageratae most closely resembles mevarna, wheeleri, and texana in that the dark ray connecting the pterostigma with vein r-m is RECOGNITION.

narrower than the length of the pterostigma, and two dark rays extend into the distal half of cell dm posteriorly from vein M, neither of which cross vein CuAj into the adjacent cell. T. ageratae differs from all three of the above-named species in having a broadly rounded hyaline triangle apicad of the pterostigma (fig. 456, a), and the hyaline spot at the apex of vein R2+3 is situated slightly apicad of the center of the dark area surrounding it (fig. 456, b). The female of this species is not known, thus it is omitted in the key to females. distribution.

host.

(1934).

Map 89.

T. ageratae was reared from flowers of Ageratum littorale by Beniamin J

T. ageratae was originally described from a single male, as far as known the only specimen ever having been collected or described. discussion.

Trupanea arizonensis Malloch (Fig. 454; Map 90) Trypanea arizonensis Malloch 1942: 15 (female holotype, NMNH; Tucson, Ariz.). Trupanea arizonensis: Foote 1960g: 12 (revision). - Foote and Blanc 1963: 75 (review, Calif.). — Foote 1965a: 667 (in catalog). In wing pattern, arizonensis resembles dacetoptera and maculigera in that the dark band connecting the pterostigma with cell r-m is narrower than the length of the pterostigma, and two narrow brown bands are present in the distal half of cell dm, the distalmost one extending to the posterior wing margin. It is distinguished from both those species by the proximal ray through cell dm, which extends proximally at an angle such that the distance between this ray and the distal one is greater than between the distal ray and the dark marking covering vein dm-cu (fig. 454, b). T. arizonensis is also unique in that the dark spot near the center of vein CuAa appears as a continuation of the proximal dark ray (fig. 454, a); it extends posteriorly into cell cua1? although it never attains the hind margin of the wing. The wing pattern of males is like those of females in every important respect. recognition.

distribution.

host.

Not

Map 90.

known.

430

Systematic Treatment of the Genera

•

Trupanea arizonensis

o

Trupanea maculigera

A

Trupanea vicmiformis

Map 90. Distribution of Trupanea arizonensis, T. maculigera, and T. viciniformis.

Trupanea bisetosa (Coquillett) (Figs. 451, 452; Map 91) Urellia bisetosa Coquillett 1899c: 266 (female lectotype, NMNH; Las Cruces, N. Mex.). - C. F. Adams 1904: 450 (in key). - F. H. Snow 1904: 345 (Ariz.). Aldrich 1905: 613 (in catalog). - Tucker 1907: 105 (Colo.). - Foote 1960g: 11 (lectotype designation, by assumption of holotype). Trypanea bisetosa: Curran 1932b: 6 (in key). - Janes and Thomas 1932: 103 (Utah). - Knowlton and Harmston 1937: 145 (Utah). - Malloch 1942: 12 (revision). Aczel 1949: 300 (in catalog). Tephritis bisetosa: Janes and Thomas 1932: 103 (Utah). Tepbritis daphne (Wiedemann) [misidentification]: Knowlton and Harmston (part) 1937: 145 (Utah). Trupanea bisetosa: Foote 1960g: 11 (revision). - Foote and Blanc 1963: 76 (review, Calif.). - Foote 1965a: 667 (in catalog). - Novak et al. 1967: 148 (host). - Goeden 1971: 47 (host). - Wasbauer 1972: 137 (hosts). - Goeden and Riker 1976a: 49 (hosts). — Hilgendorf and Goeden 1981: 106 (host). — Cavender and Goeden 1982: 400 (hosts, biology, parasites, description of larva). - Cavender and Goeden 1983: 275 (taxonomy, biology). - Goeden 1985: 569 (hosts). Only three Trupanea species in the United States and Canada possess a dark band distinctly wider than the length of the pterostigma connecting recognition.

the pterostigma with vein r-m. One of these (femoralis) is easily distinguished from

431

Trupanea

/ Map 91. Distribution of Trupanea bisetosa and T. nigricornis. Females of bisetosa identified before 7 July 1982 are not shown.

bisetosa and nigricornis by the presence of at least traces of two dark rays in cell dm and an elongated dark area covering vein CuAj from its base to the proximal ray in cell dm. The wing pattern of female bisetosa differs from that of nigricornis in that the apical Y-shaped mark of the wing is connected to the next proximal dark area by a band about two times as wide as either arm of the apical mark (fig. 451, a). This character also is found in males; in that sex, the first flagellomere is very light brown to yellow. The apical Y-shaped mark in males of bisetosa is rarely broken, and there is almost never a dark spot at the center of vein CuAj. DISTRIBUTION.

Map 91.

Wasbauer (1972) cited six plant species, each belonging to a different genus, as hosts for bisetosa. Cavender and Goeden (1982, 1983) indicated that, compared with nigricornis, the host range of this species is quite limited, and that the records listed by Wasbauer (1972) are suspect. The only species they confirmed as hosts for bisetosa are Geraea canescens, Heliatbus annuus, H. ciliaris, and H. niveus HOSTS.

ssp. tephrodes. Until the publication of investigations by Cavender and Goeden (1983), which made possible the separation of females of bisetosa and nigricornis, some misidentifications were made. For example, one female listed by Foote and Blanc (1963, p. 77) has been re-examined and identified as nigricornis. Undoubtedly discussion.

other cases of misidentification remain to be resolved.

432

Systematic Treatment of the Genera

Trupanea californica Malloch (Fig. 467; Map 92) Trypanea californica Malloch 1942: 17 (male holotype, NMNH; Emerald R. Tahoe, Calif.). Trypanea microsetulosa Malloch 1942: 17 (male holotype, NMNH; Lakeside, Tahoe, Calif.). - Foxlee 1957: 36 (B.C.). Trupanea californica: Foote 1960g: 21 (revision). - Foote and Blanc 1963: 77 (re¬ view, Calif.). - Foote 1965a: 667 (in catalog). - Goeden 1971: 47 (host). Wasbauer 1972: 47 (host). - Goeden 1985: 565, 569 (hosts). - Goeden 1986: 327 (host). — Headrick and Goeden 1991: 559 (life history, hosts).

433

Trupanea

T. californica is one of several species in which the dark ray con¬ necting the pterostigma with vein r-m is narrower than the length of the pterostigma; only one dark ray, if any, descends from vein M posteriorly into cell dm; no dark markings are present in the apex of cell br; and the hyaline spot at the extreme base of cell r4+5 is attenuated or lacking. It is distinguished from all those species by the position of the hyaline spot at the apex of vein R2-1-3; that spot lies quite close to the apical margin of the dark area surrounding it (fig. 467, b, c). recognition.

distribution.

Map 92.

T. californica appears to be restricted to species of Anapbalis and Gna¬ pbalium, according to Wasbauer (1972). Goeden (1985) reported this species from Gnapbalium bicolor and G. californicum. Headrick and Goeden (1991) studied the life history of californica on Gnapbalium beneolens in California and reported G. luteo-album, G. microcepbalum, G. palustre, G. ramissimum, G. cbilense, and Anapbalis margaritacia as additional hosts. hosts.

Trupanea conjuncta (Adams) (Fig. 446; Map 93) Urellia conjuncta Adams 1904: 451 (female holotype, UKL; Bill Williams Fork, Ariz.). — F. H. Snow 1904: 345 (Ariz.). — Byers et al. 1962: 181 (type data). Trypanea conjuncta: Malloch 1942: 12 (revision). Trupanea conjuncta: Foote 1960g: 8 (revision). — Foote 1962: 173 (taxonomy). — Foote and Blanc 1963: 78 (review, Calif.). - Foote 1965a: 667 (in catalog). Goeden 1983: 400 (host). - Goeden 1987b: 284 (biology). — Goeden and Ricker 1989: 329 (host). — Goeden and Headrick 1991b: 553 (host). The unusual wing pattern of conjuncta distinguishes it from all other U.S. and Canadian Trupanea (fig. 446). The anterior distal quarter of the disk is completely occupied by an unbroken dark area which also fills the apex of cell r4+5 (fig. 446, b). The shading proximad of vein r-m is comparatively extensive. In side view, the heads of both sexes are almost as long as high, the fore femora of the males are distinctly expanded, and the oviscape is about 1.5 times as long as the two recognition.

preceding abdominal tergites. distribution.

Map 93.

Goeden (1983) reared more than 300 adults of conjuncta from Trixis californica in southern California, and that plant was again reported as a host by HOST.

Goeden and Ricker (1989) and Goeden and Headrick (1991b). discussion.

Goeden (1987b) discussed at length the biology, seasonal history,

and parasites of conjuncta.

Systematic Treatment of the Genera

Trupanea dacetoptera Phillips (Fig. 453; Map 92) Trypanea dacetoptera Phillips 1923: 148 (female holotype, CUI; Karner, N.Y.). Johnson 1925b: 265 (N. Engl.). - Leonard 1928: 853 (N.Y.).-Johnson 1930: 151 (Nantucket). - Curran 1932b: 6 (in key). - Brimley 1938: 384 (N. Car.). Malloch 1942: 14 (revision). - V. T. Phillips 1946: 125 (hosts). - Aczel 1949: 118 (in catalog). Trupanea (Trupanea) dacetoptera: F. H. Benjamin 1934: 54 (review, Fla.). Trupanea dacetoptera: Ouellet 1941: 130 (Que.). - Foote 1960g: 11 (revision). Foote 1965a: 667 (in catalog). - Wasbauer 1972: 137 (hosts). recognition. T. dacetoptera resembles arizonensis and maculigera in having a

narrow dark band connecting the pterostigma with vein r-m and two narrow brown bands in the distal half of cell dm, the distalmost one extending to the posterior wing margin (fig. 453, b). It may be distinguished from both those species by the proximal ray through cell dm (fig. 453, c), which ends short of vein CuAj without any central dark spot on that vein in line with it. In many females, the distal ray in cell dm is fused with that covering vein dm-cu, leaving only a small hyaline spot in the apex of cell dm. The male wing pattern closely resembles that of the female, but the position of the hyaline spot at the apex of vein R2+3 is more variable, and the infuscation in the apex of cell br also varies to a greater extent in the male than in the female.

435

Trupanea distribution.

Map 92.

Gnaphalium obtusifolium and Heterotheca microcephala were cited as hosts by Wasbauer (1972). Benjamin (1934) reared this species from “tender tips of nonflowering plants” of H. microcephala. hosts.

Trupanea eclipta Benjamin (Fig. 464; Map 89) Trupanea (Trupanea) eclipta Benjamin 1934: 57 (male holotype, NMNH; Orlando, Fla.). Trypanea eclipta: Malloch 1942: 9 (revision). — V. T. Phillips 1946: 125 (host). — Hennig 1952: 212 (in catalog immature stages). Trupanea eclipta: Foote 1960g: 19 (revision). — Foote 1965a: 667 (in catalog). Wasbauer 1972: 137 (host). In wing pattern, eclipta shares with radifera and jonesi a narrow pterostigmal band; only one dark ray progressing posteriorly from vein M into cell dm; little or no dark marking in the apex of cell br; and a large, well-defined hyaline spot at the extreme base of cell r4+5. The ray connecting the pterostigma with vein r-m in eclipta always is complete in cell r1 but is broken in cell r2+3 (fig. 464, b). The opposite condition exists in radifera, in which this ray is complete in cell r2+3 but is broken in cell rj. We have seen almost no variation in this character in the specimens we have examined. The wing pattern in males of this species does not differ mate¬ recognition.

rially from that in females. DISTRIBUTION.

Map 89. We have also seen specimens from the Greater and

Lesser Antilles, Mexico, Belize, and Guatemala. host.

Benjamin (1934) reported only Eclipta alba as a host for T. eclipta; the

larvae “feed in the flowers.” Specimens in the NMNH from Dominica and the Virgin Islands were reared from flowers of E. prostrata and a Wedelia species.

Trupanea femoralis (Thomson) (Figs. 447, 448; Map 93) Trypeta femoralis Thomson 1869: 582 (male syntype, NRS; Calif.). — Curran 1932b: 5 (taxonomy). Urellia femoralis: Loew 1873: 336, 339 (taxonomy). - C. F. Adams 1904: 450 (in key). - Aldrich 1905: 613 (in catalog). - Woodworth 1913: 37 (Calif.). Trypeta (Urellia) femoralis: Osten Sacken 1878: 195 (in catalog). JJrellia occidentals Adams 1904: 452 (female lectotype, UKL; Stanford University, Palo Alto, Calif.). - Foote 1960g: 9 (lectotype designation, synonymy). - Byers et al. 1962: 181 (type data). - Foote 1962: 177 (type data). Trypanea femoralis: Malloch 1942: 10 (revision).

Systematic Treatment of the Genera Trupanea femoralis: Foote 1959a: 16 (taxonomy). - Foote 1960g: 8 (revision). Foote and Blanc 1963: 79 (review, Calif.). - Foote 1965a: 667 (in catalog). Wasbauer 1972: 137 (hosts). - Goeden 1985: 565, 569 (hosts). - Goeden 1986: 327 (host). — Goeden 1992: 108 (hosts). The wing pattern of femoralis resembles those of bisetosa and nigricornis in that the dark band connecting the pterostigma with vein r-m is as wide recognition.

in at least part of its length as the length of the pterostigma (fig. 447, a). These are the only three Trupanea species in the United States and Canada possessing this charac¬ ter. T. femoralis may be distinguished from the other two species by the presence of two dark rays, or parts thereof, extending posteriorly from vein M into cell dm (fig.

447, d) and an elongate darkening from the base to past the middle of vein CuAt (fig! 447, e), usually connecting with the proximal dark ray in cell d-m. The male wing pattern is more widely hyaline; usually only remnants of the apical Y-shaped mark are present, and the discal rays are always incomplete, although they are readily detectable. The middle femur of the male bears a patch of dark, erect setae on its anterior surface. The female of femoralis also resembles that of Tephritis labecula, but the single pair of scutellar bristles, the rounded apex of the hyaline triangle at the apex of the pterostigma, and the fact that the male wing pattern differs considerably from the female distinguishes these two species.

distribution. Map 93. hosts.

Seven plant species in the following five genera were reported as hosts by

Wasbauer (1972): Aster, Coretbrogyne, Haplopappus, Hemizonia, and Heterotheca. Goeden (1985, 1986) added five plant species in Lepidospartum and Peucephyllum. Goeden (1992) reared femoralis from the following additional genera in California: Baccharis, Brickellia, Eriophyllum, Holocarpa, Lessingia, Machaeranthera, Monolopia, Perezia, and Viguiera.

Trupanea imperfecta (Coquillett) (Figs. 460, 461; Map 94) Urellia imperfecta Coquillett 1902b: 181 (male holotype, NMNH- Williams Ariz 1 - C. F. Adams 1904: 450 (in key). - Aldrich 1905: 614 (in catalog) ’ Trypanea tmperfecta: Curran 1932b: 5 (in key). - Malloch 1942: 10 (revision) Trupanea imperfecta: Strickland 1946: 166 (Alta.). - Foote 1960g: 18 (revision). Foote and Blanc 1963: 80 (review, Calif.). - Foote 1965a: 667 (in catalog) ®S0b"1W2: 137 , ^ usuallv present and apicad of that spot (fig. 515, a) ... .HthurmaLl Foo« Hyaline incisions in cell m broader than intervening blackish areas (fig. 516, c); hyaline spot posterior to apical spot in cell r2+3 absent or contiguous with that spot (fig. 517) . 4 4. Femora and tibiae, especially of hind legs, largely blackish; wing pattern as in fig. 516) .munda (Coquillett) Legs usually wholly yellowish; if black color is present, occurring largely on basal portions of femora only . 5 5. Cell Tj with 3 hyaline incisions apicad of pterostigma (fig. 517, a); anterior arm of Y-shaped mark in apex of cell r4+5 as long as posterior arm (fig. 517, c) ...californica (Coquillett) Cell r1 with only 2 hyaline incisions apicad of pterostigma (fig. 518, a); anterior arm of Y-shaped mark at apex of cell r4+5 shorter than posterior arm (fig- 518, e).dodsoni Foote

477

Valentibulla

d

f

Figures 513—518. Right wings, Valentibulla spp. 513, steyskali Foote; 514, mundulata Foote; 515,

tburmanae Foote; 516, munda (Coq.); 517, californica (Coq.); 518, dodsoni Foote.

Valentibulla californica (Coquillett) (Fig. 517; Map 104)

Trypeta (Euaresta) californica Coquillett 1894: 73 (holotype female, NMNH; Los Angeles County, Calif.). - Quisenberry 1950: 10 (taxonomy).

Euaresta californica: Aldrich 1905: 613 (in catalog). - Woodworth 1913: 37 (Calif.). Valentibulla californica: Foote and Blanc 1959: 150 (revision). - Foote and Blanc 1963: 91 (review, Calif.). - Sanders 1964: 24 (Channel Is., Calif.). - Foote 1965a: 670 (in catalog). - Novak et al. 1967: 148 (host, biology). - Wasbauer 1972: 141 (host). — Wangberg 1978a: 472 (biology).

recognition. The wing of californica resembles most closely that of mundulata in that the hyaline spot in cell r4+5 occupies not much more than one-fourth of the apex of that cell (fig. 517, b). However, the wing of the former species is com¬ paratively narrow, in some specimens the hyaline spot in cell cuat at the apex of vein

A1 + CuA2 is somewhat enlarged (fig. 517, d), and the hind femora and tibiae are largely yellow. These two species may be further distinguished in that at least the posterior half of the scutum in mundulata is shining black and in californica it is densely tomentose.

Systematic Treatment of the Genera

distribution. Map 104.

V. californica also has been found in Baja California

Norte.

host.

V. californica has been reared from stem swellings on Chrysothamnus

nauseosus.

Valentibulla dodsoni Foote (Fig. 518; Map 104)

Valentibulla dodsoni Foote in Blanc and Foote 1987: 435 (female holotype, NMNH; Jemez Valley, Sandoval County, N. Mex.). - Dodson 1987b: 494 (biology).

479

Valentibulla

recognition. The wing pattern of

dodsoni is the most distinctive in the genus. It

differs from those of all other species in having no hyaline incision in the pterostigma, only two hyaline spots in cell r1 distad of the pterostigma (fig. 518, a), two hyaline spots at the apex of cell r2+3 in the female (fig. 518, c) and one in the male, a long, relatively narrow hyaline incision anteriorly situated in cell r4+5 (fig. 518, b), and an extensive unbroken hyaline area in most of the anal lobe (fig. 518, f). In many females, a subhyaline arc extends into cell r4+5 from the apical hyaline spot in cell r2+3 (%• -518, b, c); this marking is not present in any other Valentibulla species. distribution. Map 104.

V. dodsoni was reared from Chrysothamnus nauseosus ssp. bigelovii by Dodson (1987b). host.

Valentibulla munda (Coquillett) (Figs. 46, 516; Map 104)

Euaresta munda Coquillett 1899c: 265 (female holotype, NMNH; Elko, Nev.). Aldrich 1905: 613 (in catalog). — Quisenberry 1950: 10 (taxonomy).

Tephrella euarestoides Bates 1935: 106 (female holotype, MCZ; Ridgeway, Colo.). Valentibulla munda: Foote 1965a: 670 (in catalog). - Novak et al. 1967: 148 (host, gall). — Wasbauer 1972: 141 (hosts). — Steyskal and Foote (part) 1977: 153 (syn¬ onymy, taxonomy). recognition. In

munda, the hind femora and tibiae are almost entirely black

with yellowish color only at their apices. The apical hyaline incision in cell r4+5 of the wing resembles that of steyskali in being quite wide and deep, occupying in both species at least the apical third of that cell (fig. 516, a). In munda, however, the apical hyaline spot in cell m is much wider at the wing margin (fig. 516, b) than in steyskali (fig. 513, c), and the scutellum and apical half of the scutum are tomentose rather than shining black. distribution. Map 104.

host. Not known.

Valentibulla (Foote and Blanc 1959), we misidentified this species; our description of munda in that paper is discussion. In our original description and discussion of

actually that of steyskali.

Valentibulla mundulata Foote (Fig. 514; Map 104)

Valentibulla mundulata Foote in Foote and Blanc 1979: 175 (female holotype, UCD; Mountain Meadow Ranch at head of Coffee Cr., Trinity County, Calif.).

Systematic Treatment of the Genera

mundulata and steyskali, at least the apical half of the scutum is shining black rather than tomentose. V. mundulata RECOGNITION. In only two species of the genus,

contrasts with steyskali in having vein R2+3 gradually curving to the costa (fig. 514, a) rather than presenting a more abrupt bend, and it has a shorter and narrower hyaline incision in the apex of cell r4+5 than in steyskali (fig. 514, b). The wing of

mundulata resembles that of califovnica in the size of the apical hyaline spot in cell r4+5 (fig- 514, b), but the wing of the latter is comparatively narrow and the hyaline spot in cell cuaj at the apex of vein At + CuA2 is sometimes enlarged (fig. 517, d). Furthermore, the hind femora and tibiae of mundulata are largely black, and the scutum of californica is entirely tomentose.

distribution. Map 104.

host. Not known.

Valentibulla steyskali Foote (Fig. 513; Map 104)

Valentibulla munda (Coquillett) [misidentification]: Foote and Blanc 1959: 152 (re¬ vision).

Valentibulla steyskali Foote in Steyskal and Foote 1977: 153 (female holotype, NMNH; 18 mi e. Boise, Ida.) (new name for munda Coquillett of Foote and Blanc). - Wangberg 1978a: 472 (biology). - Foote and Blanc 1979: 176 (taxon¬ omy). - Blanc and Foote 1987: 435 (taxonomy, partial key to species).

recognition.

V. steyskali and mundulata are the only two members of the genus

in which at least the posterior half of the scutum is shining black. In steyskali, vein

R2+3 is bent forward rather sharply (fig. 513, a), and the hyaline spot in the apex of cell r4+5 is wide at the wing tip and occupies the apical V3 of cell r4+5 (fig. 513, b). The bend of vein R2+3 in mundulata is not pronounced (fig. 514, a), and the hyaline spot in the apex of cell r4+5 is narrower and shorter (fig. 514, b). The apical hyaline spot in cell r4+5 of steyskali resembles that in munda in size and shape, but in the latter species, the apical hyaline incision in cell m is much wider at the wing margin (fig. 516, b).

distribution. Map 104.

host.

V. steyskali has been reared from galls on Chrysotbamnus nauseosus.

discussion.

V. steyskali is the species we misidentified as munda (Coquillett) in

our original description and discussion of this genus (Foote and Blanc 1959).

481

Valentibulla

Valentibulla thurmanae Foote (Figs. 84, 515; Map 104)

Valentibulla thurmanae Foote in Foote and Blanc 1959: 154 (female holotype, CAS; Mt. Tamaulpais, Calif.). — Foote and Blanc 1963: 91 (review, Calif.). — Foote 1965a: 670 (in catalog). — Arnaud 1979: 332 (type data).

recognition. Among

Valentibulla species, thurmanae is rather distinctive in

having no hyaline wedge in the pterostigma, the hyaline spot at the apex of cell r4+5 does not penetrate that cell very deeply (fig. 515, b), and the three relatively narrow hyaline spots at the margin of cell m are separated by dark areas wider than the hyaline spots (fig. 515, c). A hyaline spot is usually present in cell r2+3 posterior to the apical spot in cell r^ this spot lies apicad of the former rather than directly behind it (fig. 515, a).

distribution. Map 104.

host. Not known.

Genus Xanthaciura Hendel Aciura Robineau-Desvoidy: Loew (part) 1873: 268, 328 (taxonomy). — Williston 1896: 122 (in key). — Coquillett (part) 1899c: 268 (in key). — Aldrich (part) 1905: 607 (in catalog). — Williston 1908: 286 (in key). - V. T. Phillips (part) 1923: 121, 130 (in key, revision).

Trypeta (Aciura): Osten Sacken 1878: 191 (in catalog). Xanthaciura Hendel 1914b: 86 (type species, Trypeta chrysura Thomson 1869: 580, by original designation). - Bates 1933a: 55 (taxonomy). - Curran 1932b: 2, 10 (in key, taxonomy). - F. H. Benjamin 1934: 43 (review, Fla.). - Curran 1934: 293 (in key). - Aczel 1950: 111 (revision). - Aczel 1952c: 245 (revision). - Foote and Blanc 1963: 92 (review, Calif). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 141 (hosts). - Blanc and Foote 1987: 437 (taxonomy, partial key to species).

Aciura (Eucosmoptera) Phillips 1923: 131 (type species, Acuira (Eucosmoptera) tetraspina Phillips 1923: 132, by subsequent designation of Bates 1933a: 55). - F. H. Benjamin 1934: 43 (review). - Foote 1965a: 671 (in catalog).

Aciura (Xanthaciura): V. T. Phillips 1923: 131 (in key to subgenera). Eucosmoptera: Curran (part) 1932b: 4, 12 (in key, key to species). - Curran 1934: 291 (in key). Chrysaciura Aczel 1953: 188 (type species, Chrysaciura bipuncta Aczel, by original designation. New synonymy.

481

Valentibulla

Valentibulla thurmanae Foote (Figs. 84, 515; Map 104)

Valentibulla thurmanae Foote in Foote and Blanc 1959: 154 (female holotype, CAS; Mt. Tamaulpais, Calif.). — Foote and Blanc 1963: 91 (review, Calif.). — Foote 1965a: 670 (in catalog). — Arnaud 1979: 332 (type data).

recognition. Among

Valentibulla species, thurmanae is rather distinctive in

having no hyaline wedge in the pterostigma, the hyaline spot at the apex of cell r4+5 does not penetrate that cell very deeply (fig. 515, b), and the three relatively narrow hyaline spots at the margin of cell m are separated by dark areas wider than the hyaline spots (fig. 515, c). A hyaline spot is usually present in cell r2+3 posterior to the apical spot in cell r^ this spot lies apicad of the former rather than directly behind it (fig. 515, a).

distribution. Map 104.

host. Not known.

Genus Xanthaciura Hendel Aciura Robineau-Desvoidy: Loew (part) 1873: 268, 328 (taxonomy). — Williston 1896: 122 (in key). — Coquillett (part) 1899c: 268 (in key). — Aldrich (part) 1905: 607 (in catalog). — Williston 1908: 286 (in key). - V. T. Phillips (part) 1923: 121, 130 (in key, revision).

Trypeta (Aciura): Osten Sacken 1878: 191 (in catalog). Xanthaciura Hendel 1914b: 86 (type species, Trypeta chrysura Thomson 1869: 580, by original designation). - Bates 1933a: 55 (taxonomy). - Curran 1932b: 2, 10 (in key, taxonomy). - F. H. Benjamin 1934: 43 (review, Fla.). - Curran 1934: 293 (in key). - Aczel 1950: 111 (revision). - Aczel 1952c: 245 (revision). - Foote and Blanc 1963: 92 (review, Calif). - Foote 1965a: 670 (in catalog). - Wasbauer 1972: 141 (hosts). - Blanc and Foote 1987: 437 (taxonomy, partial key to species).

Aciura (Eucosmoptera) Phillips 1923: 131 (type species, Acuira (Eucosmoptera) tetraspina Phillips 1923: 132, by subsequent designation of Bates 1933a: 55). - F. H. Benjamin 1934: 43 (review). - Foote 1965a: 671 (in catalog).

Aciura (Xanthaciura): V. T. Phillips 1923: 131 (in key to subgenera). Eucosmoptera: Curran (part) 1932b: 4, 12 (in key, key to species). - Curran 1934: 291 (in key). Chrysaciura Aczel 1953: 188 (type species, Chrysaciura bipuncta Aczel, by original designation. New synonymy.

Systematic Treatment of the Genera

recognition.

Xantkaciura is one of the most easily recognized tephritid genera

in North America north of Mexico. The frons is very narrow, its width at the vertex being less than the width of one eye (fig. 77); it bears three pairs of frontal bristles and two pairs of orbitals, the anterior pair of which are comparatively long. The first flagellomere is at least two times as long as wide, its apex attaining the anterior oral margin. One pair of dorsocentral bristles is present, situated close to the transverse suture; there are one or two pairs of scutellar bristles; the legs in the U.S. and Canadian species are entirely yellow and unmarked; and the abdominal tergites are either entirely yellow or the posterior two or three tergites are dark brown, contrast¬ ing distinctly with the anterior yellow ones. The wing pattern (figs. 519-522) is similar in the four species in the United States and Canada and differs from most other American tephritids occurring north of Mexico; it is mostly dark brownish black with two large, sharply pointed hyaline triangles in cell

immediately distad

of the pterostigma and has a number of hyaline incisions or a solid hyaline area along the posterior border; the apical one-fourth of the wing and the middle (except for 23 hyaline spots in cells br and r4+5), are entirely dark. discussion. The genus

Xanthaciura is endemic to the New World; about 12

species occur south of the United States in addition to those treated here. Benjamin (1934) presented a description of the genus and discussed its nomenclature in detail. Aczel (1950, 1952c) reviewed all but one of the known species. As discussed in the introduction, Xanthaciura bipuncta (Aczel), new combination, the type species of

Cbrysaciura, is closely related to the Xanthaciura species having vein R2+3 short, particularly speciosa Hendel and excelsa Aczel. It differs only slightly in wing pattern and in the predominantly yellow color of the pattern. We therefore consider Chrysaciura to be a junior synonym of Xanthaciura.

Key to U.S. Species of Xanthaciura 1. Scutellum brown; Scutellum yellow;

with 2 pairs of bristles (fig. 523); katepisternum and oviscape wing pattern as in fig. 519 .tetraspina (Phillips) with only 1 pair of bristles (fig. 524); katepisternum brown or oviscape yellow . 2

2. Vein R2+3 bent sharply forward beyond second hyaline costal spot (fig. 520, b), distance between apices of Rj and R2+3 equal to or less than distance between apices of R2+3 and R4+5; ocellar bristles even with or posterior to anterior ocellus; cell br dark, without a rounded hyaline spot (fig. 520, ...connexionis Benjamin Vein R2+3 ending well beyond second hyaline mark in cell rl5 distance between apices of Rj and R2+3 1.5—2.0 times as long as distance between apices of R2+3 and R4+5; ocellar bristles anterior to anterior ocellus; cell br usually with a small or indistinct hyaline spot (fig. 521, a).3 3. Sides of thorax sharply bicolored, blackish brown above, yellow ventrally (fig. 525); only the apicalmost hyaline mark in cell cuaj crossing forward over vein CuAj into posterior apical corner of cell dm (fig. 521, c) .insecta (Loew)

Xanthaciura

a

b

Figures 519-522. Right wings, Xanthaciura spp. 519, tetraspina (Phil.); 520, connexionis Benj.; 521, insecta (Lw.); 522, chrysura (Thoms.). Figures 523, 524. Dorsal view, scutellum, Xanthaciura spp. 523, tetraspina (Phil.); 524, insecta (Lw.). Figure 525. Side view, thorax, Xanthaciura insecta (Lw.).

Thorax entirely blackish brown in ground color; the two apical hyaline marks in cell cuaj fused, both crossing forward over vein CuAj or these areas fused to fill posterior apical half of cell dm (fig. 522, a) . .chrysura (Thomson)

Xanthaciura chrysura (Thomson) (Fig. 522; Map 105)

Trypeta chrysura Thomson 1869: 580 (male and female syntypes, NRS; Rio de Janeiro, Guanabara, Brazil).

Xanthaciura chrysura: Aczel 1950: 132 (revision). - Aczel 1952c: 253 (revision). Blanc and Foote 1987: 437 (taxonomy). RECOGNITION. As indicated in the key to species, the wing pattern of

chrysura

differs from those of the other three Xanthaciura species occurring north of Mexico in that both of the marginal hyaline areas in the apex of cell cuaj extend anteriorly into cell dm, or they are completely coalesced with one large spot there (fig. 522, a).

Systematic Treatment of the Genera

Xanthaciura chrysura A Xanthaciura connexionis • Xanthaciura insecta ° Xanthaciura tetraspina A

Map 105. Distribution of Xanthaciura chrysura, X. connexionis, X. insecta, and X. tetraspina.

The pattern displaying one large spot is more typical of specimens from Florida. The frons is evenly brown; the entire thorax, including the scutellum, is dark brown to black and subshining; the abdomen is brownish except for the two yellow anterior tergites; and the oviscape is yellow (except for the apex) and about two times as long as the posterior tergite.

distribution. Map 105. X. chrysura has been recorded from Mexico south to

Argentina and Brazil.

host. Not known.

485

Xanthaciura

Aczel (1952c) reported chrysura from Florida, and we have seen specimens collected earlier from that state. discussion.

Xanthaciura connexionis Benjamin (Fig. 520; Map 105) Xanthaciura sp. n.: Malloch 1933: 269 (review). Xanthaciura brevinervis [nomen nudum] Malloch 1933: 269 (taxonomy, nomencla¬ ture). Xanthaciura connexionis Benjamin 1934: 45 (male holotype, NMNH; 3 mi s. Flor¬ ida City, Fla.). - V. T. Phillips 1946: 128 (hosts). - Aczel 1949: 254 (in catalog). Aczel 1950: 118 (revision). - Aczel 1952c: 254 (revision). - Flennig 1952: 213 (in catalog immature stages). - Foote 1965a: 671 (in catalog). - Wasbauer 1972: 141 (hosts). X. connexionis may be distinguished from the other members of the genus occurring north of Mexico by the sharply bent vein R2+3 (fig. 520, b), the apex of which meets the costa quite close to the second hyaline spot in cell q. It is the only Xanthaciura species north of Mexico with the ocellar bristles not anterior to the anterior ocellus. The thorax is bicolored as in insecta (i.e., yellow ventrally), the oviscape is yellow except for its apex, and there is never a central hyaline spot in cell br (fig. 520, a). recognition.

Map 105. X. connexionis also occurs in Costa Rica, Nicaragua, Guatemala, Mexico, and many islands of the West Indies. distribution.

This species has been reared from flowers of several species in the genera Ageratum, Bidens, and Eupatorium. A. Freidberg (personal communication) has reared a few specimens (NMNFI) from flowers of Chromolaena odorata in Do¬ minica. hosts.

Benjamin (1934) described and illustrated the adult and immature stages of this species. Hardy and Delfinado (1980) reviewed its introduction into Hawaii in attempts to control the weeds Eupatorium glandulosum and E. riparium. It is not known whether this species is established there. discussion.

Xanthaciura insecta (Loew) (Figs. 77, 521, 524, 525; Map 105) Trypeta insecta Loew 1862c: 72 (female syntype, MCZ; Cuba). — Loew 1873: 268 (review). Aciura insecta: Loew 1873: 269, 329 (taxonomy). — Johnson 1895b: 337 (Fla.). — Doane 1899: 182 (distribution).-Johnson 1904a: 79 (Bermuda). - Aldrich 1905: 607 (in catalog). - Johnson 1913a: 452 (Bermuda). - Johnson 1913b: 84 (Fla.). Ogilvie 1928a: 44 (Bermuda).

Systematic Treatment of the Genera Trypeta (Aciura) insecta: Osten Sacken 1878: 191 (in catalog). Xanthaciura insecta: Hendel 1914c: 46 (review). - Malloch 1933: 267 (review). F. H. Benjamin 1934: 44 (review, Fla.). - Curran 1934: 292 (figs, head, wing). Needham 1946: 343 (biology). - V. T. Phillips 1946: 94, 128 (description, biology larva; host). - Aczel 1949: 255 (in catalog). - Aczel 1952c: 255, 263 (revision). — Flennig 1952: 21, 213 (in catalog immature stages). - Foote 1965a: 671 (in catalog). - Steyskal 1972a: 87 (host). - Wasbauer 1972: 141 (hosts). - Blanc and Foote 1987: 437 (taxonomy). recognition.

Most specimens of insecta have a bright yellow frons; the scutum

and scutellum are brown and subshining; the pleuron is bicolored (fig. 525), the dorsal half concolorous with the scutum and the ventral half broadly yellow and concolorous with the legs; the abdomen is bicolored; and the oviscape is short and also bicolored with at least the basal half to two-thirds yellow and the remainder brownish. This species is distinguished from chrysura by the color of the pleuron and by having only the narrow apical hyaline incision in cell cuaj (fig. 521, c) crossing into cell dm; from connexionis by the absence of a sharp bend in vein R2+3 (fig. 521, b), the location of the ocellar bristles, and usually a light spot, sometimes indistinct, in cell br (fig. 521, a); and from tetraspina by the presence of only one pair of scutellar bristles (fig. 524) and the color of the pleuron and oviscape. Map 105. Originally described from Cuba, insecta also occurs in Mexico, Bermuda, the Greater Antilles, and the Bahama Islands. distribution.

Ageratum littorale and a number of species of Bidens were recorded by Wasbauer (1972). A. L. Norrbom (unpublished data) has reared it from flowers of hosts.

Dahlia coccinea in Mexico and Guatemala and from flowers of Bidens squarrosa in Venezuela. Benjamin (1934) described and illustrated the adult and immature stages in detail. The third instar and puparium bear a distinctive black posterior plate. discussion.

Xanthaciura tetraspina (Phillips) (Figs. 519, 523; Map 105) Aciura (Eucosmoptera) tetraspina Phillips 1923: 132 (male and female syntypes, CUI; Columbia, Mo.). Eucosmoptera tetraspina: Curran 1932b: 12 (in key). — Bates 1933a: 55 (type spe¬ cies designation). - Curran 1934: 288 (figure of wing). Xanthaciura tetraspina: M^alloch 1933: 270 (review). — F. H. Benjamin 1934: 46 (review, Fla.). - V. T. Phillips 1946: 128 (host). - Hennig 1952: 214 (in catalog of immature stages). - Aczel 1952c: 253, 274 (revision). - Foote 1965a: 671 (in catalog). - Novak et al. 1967: 148 (host, biology). - Wasbauer 1972: 142 (hosts).

487

Xanthomyia

X. tetraspina is the only species occurring in the United States or Canada having an entirely brown oviscape and two pairs of scutellar bristles; the posterior pair are less than half as long as the anterior pair (fig. 523) but are nevertheless distinct. The frons is velvety brown, the scutum and scutellum are dark brownish gray and subshining, the pleural sclerites are uniformly dark, and the abdominal tergites are either entirely yellow or the apical two or three tergites are brown. The oviscape is shining dark brown and is about 1.25 times as long as the terminal abdominal tergite. recognition.

Map 105. X. tetraspina is known to occur in Mexico and the Bahamas Islands. Aczel (1952c) reported it from Brazil. distribution.

Wasbauer (1972) indicated that Ageratum houstonianum and three spe¬ cies of Eupatorium are hosts of this species. hosts.

Benjamin (1934) described and illustrated the adult, larva, and puparium of this species in some detail. discussion.

Genus Xanthomyia Phillips Tepbritis (part): Loew 1873: 307 (taxonomy); 328 (in key). - Aldrich 1905: 611 (in catalog). Trypeta (Tepbritis) (part): Osten Sacken 1878: 193 (in catalog). Eutreta (part): Williston 1908: 287 (in key). Xanthomyia Phillips 1923: 140 (type species, Trypeta platyptera Loew 1873: 306, by original designation). - Curran 1932b: 3 (in key). - Curran 1934: 289 (in key). — Quisenberry 1949b: 50 (taxonomy). — Foote 1964c: 140 (review). — Foote 1965a: 633 (in catalog). Small flies with reticulate wing pattern (figs. 526, 527); a short but pointed 1st flagellomere; two pairs of frontal bristles; two pairs of orbital bristles, the anterior pair very strong; a very wide frons; and a dark mark on the parafacial between the face and the anterior margin of the compound eye. Xanthomyia resembles Eurosta, Eutreta, Xenochaeta, and Jamesomyia in many morphological characters. From the two first-named genera, Xanthomyia is distin¬ guished by the pattern of the hyaline spots in the wing and from Xenochaeta and jamesomyia by the presence of only two pairs of frontal bristles and a very broad wing. The distinctive apical sclerite on the distophallus will also separate recognition.

Xanthomyia from all other New World genera. Xanthomyia is tentatively included in the tribe Eutretini, mainly because it has a parafacial spot. As noted in the classification section, this tribe is discussion.

488

Systematic Treatment of the Genera probably not a natural one; the relationships of Xantbomyia are not well under¬ stood. Noeeta (Paranoeeta), from the eastern Palearctic Region, is the most closely related taxon. Xantbomyia apparently does not occur south of the United States.

Key to U.S. and Canadian Species of Xanthomyia Wing less than 2 times as long as wide; costal margin of cell r2 + 3 with 2 or 3 large hyaline spots broadly bordering costa (fig. 526, a) . platyptera (Loew) Wing more than 2 times as long as wide; costal margin of cell r2 + 3 with 3 or 4 small (rarely 2) rounded hyaline spots narrowly or not at all bordering costa (fig. 527, a) .. (Doane)

Xanthomyia nora (Doane) (Fig. 527; Map 106) Tepbritis nora: Coquillett 1899c: 264 (synonymy). Eutreta nora Doane 1899: 184 (female holotype, WSU; Moscow Mt., Ida.). - Doane 1900: 48 (taxonomy). - Aldrich 1907: 6 (in catalog). - Foote 1966b: 124 (type data). — Zack 1984: 32 (type data). Xantbomyia nora: Foote 1964c: 140 (taxonomy). - Foote 1965a: 663 (in catalog). X. nora is a dark-winged species, and the hyaline spots are much smaller and less numerous than in platyptera, especially in cells r1 and r2+3, where recognition.

these spots only narrowly border the costa or do not touch it at all (fig. 527, a). In addition, the wing is narrower, the first flagellomere in nora tends to be more deeply emarginate, and the scutellum does not bear the dorsal marking that is so prominent in platyptera. distribution.

host.

Not

Map 106.

known.

Wings of specimens from the more northern parts of the range of nora are more extensively dark and with smaller hyaline spots than in specimens discussion.

Figures 526, 527. Right wings, Xantbomyia spp. 526, platyptera (Lw.); 527, nora (Doane).

Xanthomyia

Xanthomyia nora Xanthomyia platyptera

I

.

Map 106. Distribution of Xanthomyia nora and X. platyptera.

from farther south, but to date no dividing line between two distinctively different populations has been found. Additional material must be made available before the status of these two populations can be determined.

Xanthomyia platyptera (Loew) (Figs. 72, 75, 526; Map 106) Trypeta platyptera Loew 1873: 306 (female holotype, MCZ; Conn.). Tephritis platyptera: Loew 1873: 307, 330 (taxonomy). - Coquillett 1899c: 264 (synonymy). — Doane 1900: 48 (taxonomy). — Cockerell 1902: 346 (N. Mex). —

490

Systematic Treatment of the Genera

Aldrich 1905: 612 (in catalog). - Daecke 1906: 312 (N.J.). - Johnson 1910: 803 (N.J.). - Cresson 1914a: 278 (taxonomy). - Britton 1920: 204 (Conn.). Trypeta (Tepbritis) platyptera: Osten Sacken 1878: 193 (in catalog). Campiglossa platyptera: Cresson 1914a: 278 (taxonomy). Xanthomyia platyptera: V. T. Phillips 1923: 140 (review). - Johnson 1925b: 263 (N. Engl.). - Leonard 1928: 852 (N.Y.). - Curran 1934: 288 (figure of wing). Brimley 1938: 384 (N. Car.). - Ouellet 1941: 130 (Que.). - Foote 1964c: 140 (taxonomy). - Foote 1965a: 664 (in catalog). In contrast to that of nora, the wing pattern of platyptera exhibits more extensive hyaline areas, especially in cells r1 and r2+3, where the large hyaline spots broadly border the costa (fig. 526, a). This area in nora is occupied by several recognition.

smaller, rounded hyaline spots that hardly touch the costa. In addition, the wing is broader, the anterior border of the first flagellomere is less deeply concave, and the scutellum bears a prominent, characteristically shaped dark mark. distribution.

host.

Not

Map 106.

known.

discussion.

See the discussion of nora for comments that apply to this species.

Genus Xenochaeta Snow Xenochaeta Snow 1894: 166 (type species, Xenochaeta dichromata Snow 1894: 166, by original designation). - Williston 1896: 122 (in key). - Coquillett 1899c: 268 (in key). - Aldrich 1905: 610 (in catalog). - Williston 1908: 287 (in key). Coquillett 1910: 620 (type data). - Hendel 1914b: 82, 85 (in key). - V. T. Phillips 1923: 120, 121 (in key). - Curran 1932b: 2 (in key). - Curran 1934: 287 (in key). - Foote 1960h: 109 (key to species, taxonomy). - Foote and Blanc 1963: 92 (review, Calif.). - Foote 1965a: 664 (in catalog). - Wasbauer 1972: 142 (hosts). RECOGNITION.

Small flies with reticulate wings, black body bristles, and a yellow

abdomen that usually shows a distinct pattern of dark spots. The frons is very wide with three pairs of frontal bristles, the two anterior pairs close together, the posterior pair inside of and close to the anterior orbitals; two pairs of orbital bristles; scutellum flat with two pairs of nearly equally sized bristles; one pair of dorsocentral bristles, close to a transverse line between the supra-alars; and a dark mark on the parafacial between the face and the anterior margin of the compound eye. Many of these characters apply as well to Acidogona, Icterica, Eurosta, Eutreta, Jamesomyia, Paracantha, and Xanthomyia. From the first three of these genera, Xenochaeta is distinguished by the very different type of wing pattern (figs. 528, 529) and from Icterica and Acidogona by the much smaller size of its species. Xanthomyia and

491

Xenochaeta

Jamesomyia are heavily tomentose genera with extremely stout head hairs; the for¬ mer genus carries only two pairs of frontal bristles and has a very broad wing with the hyaline spots disposed in a different way. Xenochaeta differs from Eutreta and Paracantha by characters given in the key to genera. Foote (1960h) discussed Xenochaeta in some detail. The species are quite distinctive but are rarely encountered. We have not recognized this genus as occurring south of Mexico. Xenochaeta is most closely related to Acidogona, jamesomyia, and several Palearctic genera. discussion.

Key to Known Species of Xenochaeta Wing pattern consisting of small, discrete hyaline spots (fig. 528); 1 small dark midventral spot on each femur; dark spots fused, or nearly so, on all but the basal abdominal tergite .aurantiaca (Doane) Wing pattern consisting of large, diffuse hyaline spots (fig. 529); hind femur with 1 dark midventral spot, remaining femora entirely yellow; dark spots not fused on any but the terminal abdominal tergite . .dichromata Snow

Xenochaeta aurantiaca (Doane) (Fig. 528; Map 107) Tephritis aurantiaca: Coquillett 1899c: 264 (taxonomy). Eutreta aurantiaca Doane 1899: 185 (female holotype, WSU; Whidby I., Wash.). — Coquillett 1899c: 264 (taxonomy). — Foote 1966b: 122 (type data). - Zack 1984: 32 (type data). Xenochaeta aurantiaca: Foote 1960h: 109 (review). — Foote 1965a: 664 (in catalog). X. aurantiaca may be distinguished from dichromata by the much smaller, more numerous hyaline wing spots (fig. 528); the presence of a single small, dark midventral spot on each femur; and the fusion of the dark abdominal spots on recognition.

all but the basal abdominal tergite. distribution.

host.

Not

Map 107.

known.

Figures 528, 529. Right wings, Xenochaeta spp. 528, aurantiaca (Doane); 529, dichromata Snow.

Systematic Treatment of the Genera

o

Xenochaeta aurantiaca

• Xenochaeta dichromata ▲ Zonosemata electa A

Zonosemata vittigera

Map 107. Distribution of Xenochaeta aurantiaca, X. dichromata, Zonosemata electa, and Z. vittigera.

D. aurantiaca and dichromata are rarely collected. Only females of aurantiaca are known; all known specimens of dichromata are males except for one recorded female. Further studies may show these two species to be conspecific. discussion.

Xenochaeta dichromata Snow (Figs. 74, 529; Map 107) Xenochaeta dichromata Snow 1894: 166 (male holotype, UKL; Mt. Hood, Oreg.). Aldrich 1905: 610 (in catalog). — Coquillett 1910: 620 (type data). — Hendel 1914b: 82, 85 (type data). - Cole and Lovett 1921: 326 (Oreg.). - V. T. Phillips 1923: 123 (type data). - Foote 1960h: 109 (taxonomy). — Byers et al. 1962: 181

493

Zonosemata

(type data). - Foote 1962: 173 (type data). - Foote and Blanc 1963: 92 (review, Calif.). — Foote 1965a: 664 (in catalog). — Novak et al. 1967: 147 (host, biology). — Wasbauer 1972: 142 (host). X. dicbromata is similar to aurantiaca structurally but differs in having large, diffuse hyaline spots on the wing (fig. 529), only the hind femur with a dark midventral spot, and the dark abdominal markings not fused except on the terminal tergite. RECOGNITION.

distribution.

host.

Map 107.

Novak et al. (1967) listed Hieracium cynoglossoides as a host plant.

The comments in the discussion of aurantiaca apply to this species. Novak et al. (1967) discussed several aspects of the biology of dicbromata. discussion.

Genus Zonosemata Benjamin Spilograpba Loew [misidentification]: Loew 1873: 237, 244 (taxonomy). Williston (part) 1896: 121 (in key). - Aldrich (part) 1905: 604 (in catalog). - F. FI. Benjamin 1934: 18 (literature). Trypeta (Spilograpba): Osten Sacken (part) 1878: 190 (in catalog). Zonosema Loew [misidentification]: Coquillett (part) 1899c: 267 (in key). - Cresson 1914a: 276 (taxonomy). — Hendel 1914b: 85 (in key). - V. T. Phillips (part) 1923: 120, 127 (in key, revision). - Curran (part) 1932b: 3 (in key). - Curran (part) 1934: 289 (in key). Zonosemata Benjamin 1934: 17 (type species, Trypeta electa Say 1830: 185, by original designation). — Curran (part) 1934: 289 (in key).— Stone 1951: 47 (taxon¬ omy). — Foote 1960d: 148 (taxonomy). - Foote 1960h: 114 (taxonomy). — Bush 1965: 307 (revision, cytology). — Foote 1965a: 675 (in catalog). — Wasbauer 1972: 142 (hosts). — Steyskal 1974: 234 (taxonomy). - Steyskal 1975b: 231 (larvae). Fiernandez 1989: 205 (key, n. sp.). - Norrbom 1990: 53 (taxonomy, hosts). The species of Zonosemata are predominantly yellowish flies of moderate size with cream-colored and dark brown to black body markings and wing bands (figs. 530, 531). The dorsocentral bristles are located behind the halfway point between the acrostichal and supra-alar bristles; four pairs of frontal bristles are usually present; the posterior orbital bristles are reclinate; cell cup is drawn out into a prominent apical elongation; and the first flagellomere is distinctly pointed. In the last-named respect, Zonosemata resembles Rbagoletis, but except in R. basiola, the intercalary band (figs. 530, a; 531, a) is absent in all Nearctic species of the latter genus (see discussion of Rbagoletis). R. basiola also is a predominantly yellow spe¬ cies, but the black body markings in that species are minimal. The characters given in recognition.

493

Zonosemata

(type data). - Foote 1962: 173 (type data). - Foote and Blanc 1963: 92 (review, Calif.). — Foote 1965a: 664 (in catalog). — Novak et al. 1967: 147 (host, biology). — Wasbauer 1972: 142 (host). X. dicbromata is similar to aurantiaca structurally but differs in having large, diffuse hyaline spots on the wing (fig. 529), only the hind femur with a dark midventral spot, and the dark abdominal markings not fused except on the terminal tergite. RECOGNITION.

distribution.

host.

Map 107.

Novak et al. (1967) listed Hieracium cynoglossoides as a host plant.

The comments in the discussion of aurantiaca apply to this species. Novak et al. (1967) discussed several aspects of the biology of dicbromata. discussion.

Genus Zonosemata Benjamin Spilograpba Loew [misidentification]: Loew 1873: 237, 244 (taxonomy). Williston (part) 1896: 121 (in key). - Aldrich (part) 1905: 604 (in catalog). - F. FI. Benjamin 1934: 18 (literature). Trypeta (Spilograpba): Osten Sacken (part) 1878: 190 (in catalog). Zonosema Loew [misidentification]: Coquillett (part) 1899c: 267 (in key). - Cresson 1914a: 276 (taxonomy). — Hendel 1914b: 85 (in key). - V. T. Phillips (part) 1923: 120, 127 (in key, revision). - Curran (part) 1932b: 3 (in key). - Curran (part) 1934: 289 (in key). Zonosemata Benjamin 1934: 17 (type species, Trypeta electa Say 1830: 185, by original designation). — Curran (part) 1934: 289 (in key).— Stone 1951: 47 (taxon¬ omy). — Foote 1960d: 148 (taxonomy). - Foote 1960h: 114 (taxonomy). — Bush 1965: 307 (revision, cytology). — Foote 1965a: 675 (in catalog). — Wasbauer 1972: 142 (hosts). — Steyskal 1974: 234 (taxonomy). - Steyskal 1975b: 231 (larvae). Fiernandez 1989: 205 (key, n. sp.). - Norrbom 1990: 53 (taxonomy, hosts). The species of Zonosemata are predominantly yellowish flies of moderate size with cream-colored and dark brown to black body markings and wing bands (figs. 530, 531). The dorsocentral bristles are located behind the halfway point between the acrostichal and supra-alar bristles; four pairs of frontal bristles are usually present; the posterior orbital bristles are reclinate; cell cup is drawn out into a prominent apical elongation; and the first flagellomere is distinctly pointed. In the last-named respect, Zonosemata resembles Rbagoletis, but except in R. basiola, the intercalary band (figs. 530, a; 531, a) is absent in all Nearctic species of the latter genus (see discussion of Rbagoletis). R. basiola also is a predominantly yellow spe¬ cies, but the black body markings in that species are minimal. The characters given in recognition.

Systematic Treatment of the Genera the key to genera further distinguish these two genera. In wing pattern, Zonosemata also resembles Oedicarena, but in the latter genus, the first flagellomere is rounded, the dark body markings are not as prominent or well defined, and the scutum is microtrichose except for small bare spots at the inner ends of the transverse suture and at the bases of the dorsocentral bristles. The scutum of Zonosemata lacks microtrichia. Characters of the male surstylus and distiphallus and female terminalia and spermathecae readily distinguish Oedicarena, Rhagoletis, and Zonosemata. Bush (1965) has revised this genus. At least five Neotropical species are known in addition to the two that occur in the United States and Canada discussion.

(Hernandez 1989, Norrbom 1990). The larvae breed in fruit; except for one proba¬ bly erroneous record, all reported host plants are species of Solanaceae. Both of the species discussed herein normally infest species of Solanum, but since 1921, electa has been found in pepper. Berlocher and Bush (1982) suggested that some of the Solanaceae-breeding species of Rhagoletis may be more closely related to Zonose¬ mata.

Key to U.S. and Canadian Species of Zonosemata A pair of black presutural spots present on scutum (fig. 532, a) in addition to other dark marks; black spot or stripe present on katepisternum (fig. 534, a); wing pattern as in fig. 530 .vittigera (Coquillett) No distinct black presutural markings on scutum (fig. 533, a) or katepister¬ num (fig. 535, a); wing pattern as in fig. 531 .electa (Say)

Zonosemata electa (Say) Pepper maggot, mouche du piment (Figs. 21, 531, 533, 535; Map 107) Trypeta electa Say 1830: 185 (syntypes, probably lost; Ind.). - Osten Sacken 1858: 79 (in catalog). - LeConte 1859: 369 (repeats orig. descr.). - Loew 1862c: 71 (review). - Loew 1873: 243 (review). - Gahan 1915: 74 (parasite). - Bush 1965: 310, 314 (type data). Tephritis flavonotata Macquart 1855a: 145 (female holotype, UMO; Md.). Macquart 1855b: 125 (repeats Macquart 1855a). - Doane 1899: 179 (taxonomy). - Stone 1951: 45 (synonymy, type data). - Bush 1965: 314 (type data). Spilographa electa: Loew 1873: 244, 329 (taxonomy). - Williston 1886: 307 (host). - W. A. Snow 1894: 161 (distribution, taxonomy). - Johnson 1895b: 337 (Fla.). Doane 1899: 178 (distribution). - F. H. Snow 1903: 219 (Kans.). - Aldrich 1905: 604 (in catalog). - Johnson 1913b: 83 (Fla.). - Britton 1920: 203 (Conn.). Peterson 1923: 1 (review). - Hall et al. 1926: 18 (N.J.). - Burdette 1930: 260 (biology). Trypeta (Spilographa) electa: Osten Sacken 1878: 190 (in catalog). Trypeta sp.: Ashmead 1889: 655 (parasite).

Zonosemata

Figures 530, 531. Right wings, Zonosemata spp. 530, vittigera (Coq.); 531, electa (Say). Figures 532, 533. Dorsal view, scutum, Zonosemata spp. 532, vittigera (Coq.); 533, electa (Say). Figures 534, 535. Side view of thorax, Zonosemata spp. 534, vittigera (Coq.); 535, electa (Say).

Zonosema electa: Coquillett 1899c: 261 (taxonomy). - V. T. Phillips (part) 1923: 128 (review). — Johnson 1925b: 260 (N. Engl.). — Leonard 1928: 851 (N.Y.). — Petch and Maltaise 1932: 56 (Que.). - Curran 1934: 290 (figure of head). Brimley 1938: 383 (N. Car.). - Flennig 1952: 219 (in catalog immature stages). Bray and Triplehorn 1953: 22 (plant associations). - Peterson 1953: 331, 333, 335, 337 (figures of larva). Zonosema flavonotata: Sturtevant 1925: 215 (seminal receptacles). Zonosemata electa: F. H. Benjamin 1934: 19 (review, Fla.). — Burdette 1935: 1 (hosts, biology). - V. T. Phillips 1946: 96, 129 (description, biology larva; hosts). Stone 1951: 47 (taxonomy, synonymy). - Plant Pest Control Div., USDA 1959a: 721 (review). - Foote 1960d: 148 (taxonomy). - Foote 1960h: 114 (review). -

Systematic Treatment of the Genera

Foott 1963: 75 (review, biology). - Bush 1965: 314 (revision). - Foote 1965a: 675 (in catalog). - Foott 1968a: 16 (host). - Foott 1968b: 18 (rearing). - Foott and Timmons 1970: 105 (review). - Bierne 1971: 68 (economic importance, biology). - Goeden 1971: 46 (host). - Wasbauer 1972: 142 (host). - Steyskal 1975b: 232 (in key to larvae). - Wharton and Marsh 1978: 149 (parasite). - Hernandez 1989:207 (in key). - Norrbom 1990: 55 (host). Z. electa is larger and less heavily marked with black than vit¬ tigera. It lacks a pair of dark spots on the scutum anterior to the transverse suture, recognition.

and the katepisternum, subscutellum, and mediotergite are unmarked. The wing patterns of the two species are similar, but the discal and subapical bands of electa are wider and tend to be connected at the posterior wing margin (fig. 531, b) in contrast to those of vittigera, in which these bands are completely separated (fig. 530, b). distribution.

Map 107.

HOSTS. Six species of Solarium, including eggplant, have been reported as hosts, as well as pepper, tomato, and a Physalis species. A species of Rosa also has been reported, but this record is probably based on a misidentification of Rhagoletis basiola, which is superficially similar to electa and which normally infests species of Rosa. Tomato apparently is rarely attacked, but Z. electa is economically important in areas where it attacks pepper and eggplant. Norrbom (1990) reported that males and females of electa were reared from Solarium melanocerasum in Mississippi, a record not found in the older literature. Detailed descriptions of adults and larvae were presented by Peter¬ son (1923), Benjamin (1934), Phillips (1946), and Bush (1965). discussion.

Zonosemata vittigera (Coquillett) (Figs. 28, 34, 530, 532, 534; Map 107) Trypeta electa [misidentification]: Cockerell 1898: 155 (N. Mex.). Zonosema vittigera Coquillett 1899c: 261 (male lectotype, NMNH; Eagle Pass, Tex.). - Bush 1965: 315 (lectotype designation). Spilographa vittigera: Aldrich 1905: 604 (in catalog). Zonosema electa var.: V. T. Phillips 1923: 128 (figure of wing). Zonosemata vittigera: F. H. Benjamin 1934: 19 (taxonomy). - V. T. Phillips 1946: 129 (host). - Aczel 1955b: 153 (description, distribution). - Foote 1960d: 148 (taxonomy). - Foote 1960h: 114 (review). - Cazier 1962: 181 (biology). - Bush 1965: 315 (revision). - Foote 1965a: 675 (in catalog). - Goeden 1971: 47, 48 (host). - Goeden and Ricker 1971: 417 (biology). - Wasbauer 1972: 142 (hosts). Steyskal 1975b: 232 (in key to larvae). - Wharton and Marsh 1978: 149 (para¬ site). - Whitman et al. 1988: 532 (spider mimicry). - Hernandez 1989: 207 (in key). — Norrbom 1990: 54 (taxonomy).

497

Zonosemata Zonosemata sp.: V. T. Phillips 1946: 100 (description of larva). - Hennig 1952: 220 (in catalog immature stages). Zonosemata variegata Aczel 1954b: 162 (nomen nudum). - Bush 1965: 317 (syn¬ onymy). Z. vittigera is easily distinguished from electa by the presence of a pair of sublateral dark spots on the scutum anterior to the transverse suture (fig. 532, a), the black subscutellum and mediotergite in most specimens, the black spot on the katepisternum (fig. 534, a), and the lack of a connection on the posterior wing recognition.

border between the transverse discal and subapical bands (compare figs. 530, b and 531, b). distribution.

Map 107.

Solanum eleagnifolium Cav. This host has now been recorded well east of the range of vittigera shown in Map 106. host.

A rather detailed account of the biology of vittigera was presented by Cazier (1962). The species is of no known economic importance. It has been reported as far south as central Mexico. Z. macgregori Hernandez, from Baja California, appears to be the most closely related species. discussion.

REFERENCES CITED

Aczel, M.L. 1939. Bohrfliegen-Studien. Zool. Anz. 125: 124—131. -. 1940. Neue Beitrage zur Systematik und Okologie der palaearktischen Trypetiden. Zool. Anz. 130: 234-242. -. 1949. Catalogo de la familia “Trypetidae” de la Region Neotropical. Acta Zool. Lilloana 7: 177-328. -. 1950. A revision of the genus “Xantbaciura” Hendel based on Argentine species. Acta Zool. Lilloana 8 (1949): 111 — 146. -. 1951. Generos y especies de la Tribu “Trypetinae.” II. Dos generos y tres especies nuevos de la Argentina. Acta Zool. Lilloana 9 (1950): 307—323. -. 1952a. Supplemento al “Catalogo de la familia Trypetidae de la Region neotropical.” Acta Zool. Lilloana 12 (1951): 117-133. -. 1952b. Revision parcial del genero Americano Paracantha Coquillett. Acta Zool. Lilloana 10: 199—243. -. 1952c. Further revision of the genus Xanthaciura Hendel. Acta Zool. Lilloana 10: 245280. -. 1952d. El genero Euaresta Loew (= Camaromyia Hendel) en la Region Neotropical. Rev. Chil. Entomol. 2: 147—172. -. 1953. La familia Tephritidae en la Region Neotropical. 1. Acta Zool. Lilloana 13: 97-

200. _. 1954a. Generos y especies de la tribus “Trypetini.” 3. Sobre los generos Rbagoletis, “Phorelliay Tomoplagioides. Dusenia 5: 71-94. _. 1954b. Generos y especies de la tribus Trypetini. 4. El genero Rhagoletotrypeta y nuevas especies de Tomoplagia y de Xonosemata. Dusenia 5: 137—164. _. 1955a. Fruit flies of the genus Tomoplagia Coquillett. Proc. U.S. Natl. Mus. 104(3343): 321-411. _. 1955b. The comparative morphology of the Tomoplagia Coquillett species. Dusenia 6: 139-170. _# 1955c. The genus Parastenopa Hendel (Diptera: Tephritidae). Wasmann J. Biol. 13: 167-187. . . ^ n _ 1958. Revision of the American species of the genus Acinia Robineau-Desvoidy. Rev. Bras. Entomol. 8: 75—106.

499

References Cited

Adams, C.C. 1915. An ecological study of prairie and forest invertebrates. Bull. Ill. State Lab Nat. Hist. 11(2): 33-280. Adams, C.F. 1904. Notes on and descriptions of North American Diptera. Kans Univ Sci Bull. 2(14): 433-455. Agee, H.R., and D.L. Chambers. 1980. Fruit fly quality monitoring, pp. 7-15. In T. Iwata [ed.], Proceedings of a symposium on fruit fly problems. Kyoto and Naha, Aug. 9-12, 1980. Natl. Inst. Agric. Sci., Yataba, Ibaraki 305, Japan. Agee, H.R., and M.L. Park. 1975. Use of the electroretinogram to measure the quality of the vision of the fruit fly. Environ. Lett. 10(2): 171-176. Agee, H.R., W.A. Phyllis, and D.L. Chambers. 1977. The compound eye of the Caribbean fruit fly and the apple maggot fly. Ann. Entomol. Soc. Am. 70(3): 359-364. Aldrich, J.M. 1905. A catalog of North American Diptera (or two-winged flies). Smithson. Misc. Collect. 46(1444): 1-680. -. 1907. Additions to my catalog of North American Diptera. J. N.Y. Entomol. Soc. 15:29. -. 1909. The fruit-infesting forms of the dipterous genus Rhagoletis, with one new species. Can. Entomol. 9: 69-72, pi. 4. -. 1910. A decennial confusion. Can. Entomol. 42: 99-100. . 1913. Collecting notes from the Great Basin and adjoining territory. Entomol News 24* 214-221. -. 1925. New Diptera or two-winged flies in the United States National Museum. Proc. U.S. Natl. Mus. 66(18): 1-36. --. 1929. A revision of two-winged flies of the genus Procecidochares in North America with an allied new genus. Proc. U.S. Natl. Mus. 76(2): 1-13. AhNiazee, M.T. 1973a. Cherry fruit fly in Oregon: a nomenclature problem. Bull. Entomol. Soc. Am. 19: 103-104. . 1973b. Biology and control of the western cherry fruit fly in Oregon. Proc. Oreg. Hortic. Soc. 64: 75-78. -• 1974a. The western cherry fruit fly, Rhagoletis indifferens (Diptera: Tephritidae). 1. Distribution of the diapausing pupae in the soil. Can. Entomol. 106: 909-912. -. 1974b. The western cherry fruit fly, Rhagoletis indifferens (Diptera: Tephritidae). 2. Aggressive behavior. Can. Entomol. 106: 1201-1204. . 1975. Susceptibility of diapausing pupae of Rhagoletis indifferens and a parasite to subfreezing temperature. Environ. Entomol. 4: 1011-1013. .1976. Thermal unit requirements for determining adult emergence of the western cherry fruit fly (Diptera: Tephritidae) in the Willamette Valley of Oregon. Environ Entomol 5397-402. -. 1978. The western cherry fruit fly (Diptera: Tephritidae). 3. Developing a management program by utilizing attractant traps as monitoring devices. Can. Entomol. 110: 11331139. —. 1979. A computerized phenology model for predicting biological events of Rhagoletis indifferens (Diptera: Tephritidae). Can. Entomol. Ill: 1101-1109. AhNiazee, M.T and R.D. Brown. 1974. A bibliography of North American cherry fruit flies (Diptera: Tephritidae). Bull. Entomol. Soc. Am. 20: 93-100. }977' Laboratory rearing of the western cherry fruit fly, Rhagoletis indifferens (Diptera: tephritidae): oviposition and larval diets. Can. Entomol. 109: 1227-1234. AhNiazee M.T., and R.L. Penrose. 1981. Apple maggot in Oregon: a possible new threat to the apple industry. Bull. Entomol. Soc. Am. 27: 245-246. Allen, T.C. 1931. Bacteria producing rot of apple in association with the apple maggot Rhagoletis pomonella. Phytopathology 21: 338. Allen, T.C., and C.L. Fluke, Jr. 1933. Notes on the life history of the apple maggot in Wisconsin. J. Econ. Entomol. 26: 1108-1112.

501

References Cited

Alien, T.C., and A.J. Ricker. 1932. A rot of apple caused by Pbytomonas melophthora, n. sp., following invasion by the apple maggot. Phytopathology 22: 557-571. Allen, T.C., J.A. Pinckard, and A.J. Ricker. 1934. Frequent association of Pbytomonas melopbtbora with various stages in the life cycle of the apple maggot, Rbagoletis pomonella. Phytopathology 24: 8-238. Ametangelo, J.R. 1974. Infestation of seeds of Ambrosia trifida, giant ragweed, by larval insects. Bios 45(1): 15-18. Am. Soc. Hortic. Sci. 1983. Symposium. The medfly in California: the threat, defense strat¬ egies, and public policy. Hortic. Sci. 18(1): 38-52 (special insert). Angeles, N. de J. 1966. Damage of Anastrepba mombinpraeoptans Sein in mangoes. Agron. Trop. (Ven.) 16(4): 299-302. Anonymous. 1915. Review of “Papaya fruit fly,” by Knab and Yothers. Exp. Stn. Rec. 32(1): 60—61. APHIS* 1983. Medfly and the aftermath. Symposium presented at the Entomological Society of America’s annual meeting, 1982. U.S. Dep. Agnc., APHIS 81-60. 58 pp. Arnaud, P.H., Jr. 1979. A catalog of the types of Diptera in the collection of the California Academy of Sciences. Myia 1: 1-505. Arnaud, P.H., and T.C. Owen. 1981. Charles Howard Curran (1894-1972) 1-393.

Myia 2-

Arrand, J.C., and W.S. Peters. 1968. A record of Rbagoletis indifferens Curran from Creston, B.C. J. Entomol. Soc. B.C. 65: 40. Ashley, T.R., and D.L. Chambers. 1979. Effects of parasite density and host availability on progeny production by Biosteres longicaudatus (Hymenoptera: Braconidae), a parasite of Anastrepba suspensa (Diptera: Tephritidae). Entomophaga 24: 363-369. Ashley, T.R., P.D. Greany, and D.L. Chambers. 1976. Adult emergence in Biosteres (Opius) longicaudatus and Anastrepba suspensa in relation to the temperature and moisture con¬ centration of the pupation medium. Fla. Entomol. 59: 391-396. Ashmead, W.H. 1889. Descriptions of new Braconidae in the collection of the U.S. National Museum. Proc. U.S. Natl. Mus. 11(1888): 611-671. -. 1890. On the Hymenoptera of Colorado: descriptions of new species, notes, and a list of the species found in the state. Colo. Biol. Assoc. Bull. 1: 405—449. -. 1892. Discovery of the genus Crataepus Forster in America, and the description of a new species. Can. Entomol. 24: 309—310. -. 1894. Descriptions of new parasitic Hymenoptera. Trans. Am. Entomol. Soc. 21: 318— 344. Auclair, J.L., A.A. Beaulieu, J.A. Doyle, R. Martineau, F.O. Morrison, R.O. Paradis, L. Raynault, and Frere Adrien Robert. 1964. French names of insects of Canada. Min. Agric. Col. Que., Publ. 295. 102 pp. Averill, A.L., and R.J. Prokopy. 1980. Release of oviposition-deterring pheromone by apple maggot flies, Rbagoletis pomonella (Walsh) (Diptera: Tephritidae). J. N.Y. Entomol. Soc. 88: 34. -. 1981. Oviposition-deterring fruit-marking pheromone in Rbagoletis pomonella. Fla. Entomol. 64: 222—226. -. 1982. Oviposition-deterring fruit-marking pheromone in Rbagoletis zepbyria. J. Ga. Entomol. Soc. 17: 315—319. -. 1989. Host-marking pheromones, pp. 207—219. In A.S. Robinson and G. Hooper [eds.], Fruit flies. Their biology, natural enemies and control. World Crop Pests, vol. 3A. Elsevier, Amsterdam. Ayers, E.L. 1957. The two medfly eradication programs in Florida. Proc. Fla. State Hortic. Soc. 70: 67-69. Ayers, E.L., and G.G. Rohwer. 1957. The Mediterranean fruit fly eradication campaign in Florida. Proc. Fla. State Hortic. Soc. 69 (1956): 12—15.

References Cited Babb, G.F. 1902. Note on Rbagoletis suavis Lw. with a description of the larva and puparium. Entomol. News 13: 242, pi. XIV. Back, E.A. 1914. The Mediterranean fruit fly in Bermuda. U.S. Dep. Agric. Bull. 161. 8 pp. Back, E.A., and C.E. Pemberton. 1914. Life history of the melon fly. J. Agric. Res. Ill: 269274. Baerwald, R.J., and G.M. Boush. 1967. Selection of a non-diapausing race of apple maggot. J. Econ. Entomol. 60: 682—684. Baker, A.C., W.E. Stone, C.C. Plummer, and M. McPhail. 1944. A review of studies on the Mexican fruit fly and related Mexican species. U.S. Dep. Agric. Misc. Publ. 531. 155 pp. Baker, C.F. 1895. Biological notes on some Colorado Diptera. Entomol. News 6: 173—174. -. 1904. Reports on Californian and Nevadan Diptera, 1. Invertebr. Pac. 1: 17—39. Balduf, W.V. 1957. The weight of puparia of Rbagoletis basiola (O.S.). Entomol. News 68: 163-165. -. 1958. The puparium of Rbagoletis basiola (O.S.)—variation in length and form. Ohio J. Sci. 58(1): 7-14. -. 1959. Obligatory and facultative insects in rose hips—their recognition and bionomics. Ill. Biol. Monogr. 26: 1-194. Ballou, L.M. 1925. Some dipterous galls of southern California. J. Entomol. Zook, Pomona Coll. 17: 1-8. -. 1926. Some dipterous galls of California. J. Entomol. Zook, Pomona Coll. 18: 25—32. Banham, F.L. 1971. Native hosts of western cherry fruit fly (Diptera: Tephritidae) in the Okanagan Valley of British Columbia. J. Entomol. Soc. B.C. 68: 29—32. -. 1972. Cherry fruit flies in the Okanagan Valley. Can. Agric. 17: 25-27. -. 1973. An evaluation of traps for the western cherry fruit fly (Diptera: Tephritidae). J. Entomol. Soc. B.C. 70: 13-16. Banks, N. 1912. The structure of certain dipterous larvae with particular reference to those in human foods. U.S. Dep. Agric. Tech. Ser. 22. 44 pp. -. 1926. Descriptions of a few new American Diptera. Psyche 33: 42—44. Baranowski, R.M. 1969. Ally found against Caribbean fruit fly, pp. 10-11. In Sunshine State Agric. Res. Rep., Nov. 1969, Gainesville, Fla. -. 1974. Release of Opius longicaudatus against Anastrepba suspensa in Florida. Fruit Fly News 3: 17. -. 1975. Bionomics and mass rearing of Anastrepba suspensa (Loew), p. 279. In Annu. Res. Rep., Inst. Food Agric. Sci., Univ. Fla., Gainesville. Baranowski, R.M., and R.W. Swanson. 1970. Introduction of Paracbasma ( = Opius) cereus (Flymenoptera: Braconidae) into Florida as a parasite of Anastrepba suspensa (Diptera: Tephritidae' Fla. Entomol. 53: 161-162. -. 1972. The utility of Paracbasma cereus (Hymenoptera: Braconidae) as a means of suppressing Anastrepba suspensa (Diptera: Tephritidae) populations. Proc. Tall Timbers Conf. 3: 249-252. Barnes, M.M. 1959. Radiotracer labeling of a natural tephritid population and of flight range of the walnut husk fly. Ann. Entomol. Soc. Am. 52: 90-92. Barnes, M.M., and J.C. Ortega. 1958. A glycine-sodium hydroxide solution as an attractant for walnut husk fly. J. Econ. Entomol. 51: 532-534. -. 1959. Experiments with protein hydrolysate bait sprays for control of the walnut husk fly. J. Econ. Entomol. 52: 279-285. Barnes, M.M., and FLT. Osborn. 1958. Attractants for the walnut husk fly. J. Econ. Entomol. 51: 686-689. Barrett, R.E. 1932. An annotated list of the insects and arachnids affecting the various species of walnuts or members of the genus Juglans. Univ. Calif. Publ. Entomol. 5(15): 275-309. Bateman, M.A. 1972. The ecology of fruit flies. Annu. Rev. Entomol. 17: 493-518. Bates, M. 1933a. Notes on American Trypetidae (Diptera). II. Psyche 40: 48-56. -. 1933b. Notes on West Indian Trypetidae. Bull. Brooklyn Entomol. Soc. 28: 160-171.

503

References Cited

-. 1934a. [Note], p. 295. In C.H. Curran, Families and genera of North American Diptera (privately published). -. 1934b. The peristigmal gland cells of trypetid larvae. Ann. Entomol. Soc. Am. 27: 1-4. -. 1934c. Notes on American Trypetidae. IV. Acrotaenia and similar genera. Rev. Entomol. 4: 1-17. -. 1935. Notes on American Trypetidae (Diptera). III. The genus Tephrella. Pan-Pac. Ento¬ mol. 11: 103—114. Batra, S.W.T. 1979. Reproductive behavior of Euaresta bella and E. festiva (Diptera: Tephritidae), potential agents for the biological control of adventive North American rag¬ weeds (Ambrosia spp.) in Eurasia. J. N.Y. Entomol. Soc. 87: 118-125. Batra, S.W.T., J.R. Coulson, P.H. Dunn, and P.E. Boldt. 1981. Insects and fungi associated with Carduus thistles (Compositae). U.S. Dep. Agric. Tech. Bull. 1616. 100 pp. Baulne, J.L. 1953. Amis et ennemis des cultures. Que. Soc. Prot. Plants, 32nd and 33rd reports (for 1951 and 1951, resp.). 232 pp. Beach, A.M. 1895. Some bred parasitic Hymenoptera in the Iowa Agricultural College collec¬ tion. Proc. Iowa Acad. Sci. (1894): 92-94. Beck, D.E. 1932. Life history notes and a study of the effects of humidity on adult emergence of Rbagoletis suavis Cress., from pupae at a constant temperature. J. N.Y. Entomol. Soc. 40: 497_499. Beckham, C.M., and H.H. Tippins. 1972. Observations on sunflower insects. J. Econ. Ento¬ mol. 65: 865—866. Beckwith, C.S. 1945. The insects of the cultivated blueberry, pp. 43-51. In The cultivated blueberry industry in New Jersey. N.J. Dep. Agric. Circ. 356. 54 pp. Beirne, B.P. 1971. Pest insects of annual crop plants in Canada, II. Diptera. Mem. Entomol. Soc. Can. 78: 48—70. Benbow, S.M., and D.E. Foster. 1982. Biology of Eutreta diana Osten Sacken on sand sage¬ brush, Artemisia filifolia Torr. (Diptera: Tephritidae). Pan-Pac. Entomol. 58: 19—24. Benjamin, D.M., and A.C. Flodson. 1949. A new bait trap collection record for Rbagoletis completa. J. Econ. Entomol. 42: 707—708. Benjamin, F.H. 1934. Descriptions of some native trypetid flies with notes on their habits. U.S. Dep. Agric. Tech. Bull. 401. 95 pp. Bennett, F.D. 1982. Report of work carried out April 1981-March 1982. Commonw. Inst. Biol. Control, London. 80 pp. Bennett, F.D., and R.M. Baranowski. 1981. Correction of the host record for Zeteticontus insularis (Howard) (Hymenoptera: Encyrtidae) in Florida. Fla. Entomol. 64: 197-198. Berg, G.H. 1979. Pictorial key to fruit fly larvae of the family Tephritidae. Org. Int. Reg. San. Agropec. (OIRSA), San Salvador. 36 pp. Berlocher, S.H. 1980a. An electrophoretic key for distinguishing species of the genus Rbagoletis (Diptera: Tephritidae) as larvae, pupae, or adults. Ann. Entomol. Soc. Am. 73: 131-137. -. 1980b. Genetics of the allozymes of the apple maggot. J. Hered. 71: 63-67. _. 1981. A comparison of molecular and morphological data, and phenetic and cladistic methods, in the estimation of the phylogeny in Rbagoletis (Diptera: Tephritidae), pp. 1-31. In M.W. Stock [ed.], Applications of genetics and cytology in insect systematics and evolu¬ tion. For. Wildl. Range Exp. Stn., Univ. Idaho, Moscow. 152 pp. _. 1984a. A new North American species of Rbagoletis (Diptera: Tephritidae), with records of host plants of Corwws-infesting Rbagoletis. J. Kans. Entomol. Soc. 57. 237—242. _# 1984b. Electrophoretic evidence for the validity of Euaresta stelligera Coquillett (Dip¬ tera: Tephritidae). J. Kans. Entomol. Soc. 57: 354—357. _ 1984c. Segregation and linkage of allozymes of the walnut husk fly. J. Hered. 75. 392— 396. . _ 1984d. Genetic changes coinciding with the colonization of California by the walnut husk fly, Rbagoletis completa. Evolution 38: 906-918.

504

References Cited

Berlocher, S.H., and G.L. Bush. 1982. An electrophoretic analysis of Rhagoletis (DipteraTephritidae) phylogeny. Syst. Zool. 31: 136-155. Berlocher, S.H., and D.C. Smith. 1983. Segregation and mapping of allozymes of the apple maggot fly. J. Hered. 74: 337-340. Berry, N.O. 1955. Mexican fruit fly and citrus black fly control in Mexico. J. Econ Entomol 48: 414-416. Berube, D.E. 1978a. Larval descriptions and biology of Tephritis dilacerata (Diptera: Tephritidae), a candidate for the biological control of Sonchus arvensis in Canada. Entomophaga 23: 69-82. . 1978b. The basis for host plant specificity in Tephritis dilacerata and T. formosa (Dip¬ tera: Tephritidae). Entomophaga 23: 331-337. Berube, D.E., and J.H. Myers. 1983. Reproductive isolation between Urophora affinis and U. quadrifasciata (Diptera: Tephritidae) in British Columbia. Can. J. Zool. 61: 787-791. Berube, D.E., and R.V. Zacharuk. 1983a. The abdominal musculature associated with oviposition in gall forming fruit flies in the genus Urophora. Can J. Zool. 61: 1805-1814. . 1983b. The ultrastructure of a novel muscle attachment in the tephritid fruit flv Urophora affinis. Can. J. Zool. 61: 1815-1823. Beutenmuller, W. 1892. Catalog of gall-producing insects found within fifty miles of New York City, with descriptions of their galls, and of some new species. Bull. Am Mus Nat Hist. 4(1): 245-278, pis. IX-XVI. Bezzi, M. 1909. Le specie dei genen Ceratitis, Anastrepha e Dacus. Portici, R. Scuola Sup Agnc., Lab. Zool. Gen. Agric. Bol. 3: 273-313. -. 1910. Restaurazione del genere Carpomyia (Rond.) A. Costa. Portici, R. Scuola Sup Agnc., Lab. Zool. Gen. Agric. Bol. 5: 3-33. ——. 1913 Indian trypaneids (fruit flies) in the collection of the Indian Museum, Calcutta Mem. Indian Mus. 3(3): 53-168. ——.1916. On the fruit flies of the genus Dacus (s.l.) occurring in India, Burma and Ceylon Bull. Entomol. Res. 7: 99-121. y . 1920. Species duae novae generis Oedaspis s.l. Broteria 18: 5-13. -. 1923. The genus Urophora in America. Trans. Am. Entomol. Soc. 64: 1-6, pi. 1. —. 1924a South African trypaneid Diptera in the collection of the South African Museum Ann. South Afr. Mus. 29: 449-577. • 1924b. Further notes on the Ethiopian fruit flies, with keys to all the known genera and species. Bull. Entomol. Res. 15: 73-118. Bezzi, M., and J.S. Tavares. 1916. Alguns muscideos cecidogenicos do Brasil. Broteria (Ser Zool.) 14: 155-170. Biggs, J.D. 1972. Aggressive behavior in the adult apple maggot (Diptera: Tephritidae). Can. Entomol. 104: 349-353. Bigot J.M.F. 1857. Dipteros, pp. 328-349. In R. de la Sagra, Historia fisica, politica y natural de las Isla de Cuba. Segunde parte. Historia natural. Paris (1856) 7. xxxii + 371 pp. . 1858. Insectes dipteres pour servir a la faune du Gabon. Arch. Entomol. 2: 131 -. 1859. Dipterorum aliquot nova genera. Rev. Mag. Zool. 7: 305-315. —. 1884 Description d’un nouveau genre et d’une nouvelle espece de Dipteres de la famille des Ortalidae. Bull. Soc. Entomol. Fr. 1884: xxix. Bird, R.D. 1927. Notes on insects bred from native and cultivated fruit trees and shrubs of southern Manitoba. Can. Entomol. 59: 124-128. Bissell, T.L. 1936. Dahlia insect pests. Gard. Chron. Am. 40: 233, 248. Blanc, F.L. 1959. A new species of Chaetostoma from California. Pan-Pac. Entomol. 35: 201— . 1970. Status of oriental fruit fly in California. Pan-Pac. Entomol. 46: 72. Blanc, F.L., and R.H. Foote. 1961. A new genus and five new species of California Tephritidae Pan-Pac. Entomol. 37: 73-83.

505

References Cited

.1987. Taxonomic observations on United States Tephritidae (Diptera), with descriptions of new species. Proc. Entomol. Soc. Wash. 89: 425-439. Blanc, F.L., and H.H. Keifer. 1955. The cherry fruit fly in North America. Calif. Dep. Aerie Mo. Bull. 44: 77-88. ‘ Blanchard, E.E. 1929. Descriptions of Argentine Diptera. Physis (B. Aires) 9: 458-465. Blanton, F.S. 1952. Eutreta sparsa found on chrysanthemum (Tephritidae). Entomol. News 63: 75. Boiler, E.F., and R.J. Prokopy. 1976. Bionomics and management of Rhagoletis. Annu. Rev. Entomol. 21: 23—246. Borror, D.J., D.M. DeLong, and C.A. Triplehorn. 1976. An introduction to the study of insects, 4th ed. Holt, Rinehart & Winston, New York. Bot, J. 1965. Census of a dacine population (Diptera: Tripetidae). J. Entomol. Soc. S. Afr. 28166-178. Boulanger, L.W., G.E. Stanton, and A.L. Padula. 1969. Continuous rearing of the apple maggot. J. Econ. Entomol. 62: 748-749. Bourne, A.I., W.H. Thies, and F.R. Shaw. 1934. Some observations on long-distance dispersal of apple maggot flies. J. Econ. Entomol. 27: 352-355. Boush, G.M., and R.J. Baerwald. 1967. Courtship behavior and evidence for a sex pheromone in the apple maggot parasite Opius alloeus (Hymenoptera: Braconidae). Ann. Entomol. Soc. Am. 60: 865—866. Boush, G.M., and F. Matsumura. 1967. Insecticidal degredation by Pseudomonas melopbthora, the bacterial symbiote of the apple maggot. J. Econ. Entomol. 60: 918-920. Boush, G.M., R.J. Baerwald, and S. Miyazaki. 1969. Development of a chemically defined diet for adults of the apple maggot based on amino acid analysis of honeydew. Ann. Entomol. Soc. Am. 62: 19—21. Boush, G.M., S.M. Saleh, and R.M. Baranowski. 1972. Bacteria associated with the Carib¬ bean fruit fly. Environ. Entomol. 1: 30—33. Boyce, A.M. 1929a. A new pest of English walnuts in California. J. Econ. Entomol. 22: 269270. -. 1929b. The walnut husk fly (Rhagoletis juglandis Cresson). J. Econ. Entomol. 22: 861—

866. -. 1930. The walnut husk fly. Calif. Dep. Agric. Mo. Bull. 19: 249—253. -. 1931. The diapause phenomenon in insects, with special reference to Rhagoletis completa Cress. J. Econ. Entomol. 24: 1018—1024. -. 1932. Mortality of Rhagoletis completa Cress. (Diptera: Trypetidae) through ingestion of solid materials. J. Econ. Entomol. 25: 1053—1059. -. 1933. Influence of host resistance and temperature during dormancy upon the seasonal history of the walnut husk fly, Rhagoletis completa Cress. J. Econ. Entomol. 26: 813-819. -. 1934. Bionomics of the walnut husk fly, Rhagoletis completa. Hilgardia 8: 363-579. Boyce, A.M., and B.R. Bartlett. 1941. Lures for the walnut husk fly. J. Econ. Entomol. 34: 318. Brandhorst, C.T. 1943. A study of the relationship existing between certain insects and some native western Kansas forbs and weedy plants. Trans. Kans. Acad. Sci. 46: 165-175. Bray, D.F., and C.A. Triplehorn. 1953. Survey of the insect fauna of red and pin oaks in Delaware. Del. Agric. Exp. Stn. Bull. 297. 28 pp. Breakey, E.P. 1945. Cherry fruit flies. Wash. Agric. Exp. Stn. Bull. 470: 119-120. Breland, O.P. 1938. Notes on sunflower Callimomidae with description of a new Zaglyptonotus. Ann. Entomol. Soc. Am. 31: 202—206. -. 1939. Additional notes on sunflower insects. Ann. Entomol. Soc. Am. 32: 719-726. Bridwell, J.C. 1923. Pupae of the walnut hull maggot living two years (Rhagoletis suavis Loew). J. Wash. Acad. Sci. 13: 262—263. Brimley, C.S. 1938. The insects of North Carolina. N.C. Dep. Agric., Div. Entomol. 560 pp.

References Cited

Brink, J.E. 1923. The sunflower maggot (Straussia longipennis Wied.). Entomol. Soc. Ont., Annu. Rep. for 1922: 72—74. Brittain, W.H. 1918. Miscellaneous notes on the apple maggot (1917) (Rhagoletis pomonella Walsh). Proc. Entomol. Soc. Nova Scotia (1917): 37—41. -. 1919. Further notes on the apple maggot (1918) (Rhagoletis pomonella Walsh). Proc. Entomol. Soc. Nova Scotia (1918): 15—23. Brittain, W.H., and C.A. Good. 1917. The apple maggot in Nova Scotia. Prov. Nova Scotia, Dep. Agric. Bull. 9: 1—70. Britton, W.E. 1906. Apple maggot infesting huckleberries. Conn. Agric. Exp. Stn., 29th Annu. Rep. (1905): 260. -. 1920. Check-list of the insects of Connecticut. Conn. Geol. Nat. Hist. Surv. Bull. No. 31. 397 pp. Brodie, W. 1892. Canadian galls and their occupants - Eurosta solidaginis, Fitch. Can. Ento¬ mol. 24: 137-139. Brogdon, J.E. 1956. The Mediterranean fruit fly and its importance in relation to mangoes, pp. 23-26. In Proc. 16th Annu. Mtg., Fla. Mango Forum, Gainesville. Brogdon, J.E., and D.O. Wolfenbarger. 1968. The papaya fruit fly. Fla. Agric. Ext. Circ. 136-B. 7 pp. Brooks, F.E. 1921. Walnut husk maggot. U.S. Dep. Agric. Bull. 992. 8 pp. Brown, A.C. 1937. Report of the Grove Inspection Department, July 1, 1934 to June 1, 1936, pp. 15—27. In State Plant Board Fla., 11th Bien. Rep. Gainesville. Brown, R.D., and M.T. AliNiazee. 1977. Synchronization of adult emergence of the western cherry fruit fly in the laboratory. Ann. Entomol. Soc. Am. 70: 678-680. Brues, C.T. 1924. The specificity of food plants in the evolution of phytophagous insects. Am. Nat. 58: 127-144. Burdette, R.C. 1930. The effect of talc on the oviposition of a trypetid. I. Econ. Entomol. 23: 260-265. -. 1935. The biology and control of the pepper maggot, Zonosemata electa Say. N.J. Agric. Exp. Stn. Bull. 585. 24 pp. Burditt, A.K., Jr. 1982. Anastrepha suspensa (Loew) (Diptera: Tephritidae) McPhail traps for survey and detection. Fla. Entomol. 65: 367-373. Burditt, A.K., Jr., and R.A. Baranowski. 1982. Management of arthropod pests for subtropi¬ cal fruits, pp. 85-90. In Tree fruit and nut pest management in the southeastern United States (D.T. Johnson, organizer). Misc. Publ., Entomol. Soc. Am. 12(2). Burditt, A.K., Jr., and T.P. McGovern. 1979. Chemicals tested as attractants for the Caribbean fruit fly, 1972-1978. U.S. Dep. Agric., ARM S-6. 47 pp. Burditt, A.K., Jr., D.L. Von Windeguth, and R.J. Knight, Jr. 1974. Induced infestation of fruit by the Caribbean fruit fly, Anastrepha suspensa (Loew). Proc. Fla. Hortic. Soc. 87: 387390. Buriff, C.R., and D.G. Davis. 1974. Tachinid flies captured in traps for the apple maggot. Environ. Entomol. 3: 572. Burk, T. 1981. Signaling and sex in acalyptrate flies. Fla. Entomol. 64: 30-43. -. 1983. Behavioral ecology of mating in the Caribbean fruit fly, Anastrepha suspensa (Loew) (Diptera: Tephritidae). Fla. Entomol. 76: 678-682. Burk, T.D., and J.C. Webb. 1983. Effect of male size on calling propensity, song parameters, and mating success in Caribbean fruit flies (Anastrepha suspensa (Loew)). Ann. Entomol. Soc. Am. 76: 678-682. Burks, B.D. 1943. The North American parasitic wasps of the genus Tetrastichus - a contribu¬ tion to biological control of insect pests. Proc. U.S. Natl. Mus. 93(3170): 505-608. -. 1955. A new species of the genus Halticoptera. Proc. Entomol. Soc. Wash. 57: 81-82. Bush, G.L. 1962. The cytotaxonomy of the larvae of some Mexican fruit flies in the genus Anastrepha (Tephritidae, Diptera). Psyche 69: 87-101.

507

References Cited

• 1965. The genus Zonosemata, with notes on the cytology of two species (Diptera Tephritidae). Psyche 72: 307-323. . 1966a. The taxonomy, cytology, and evolution of the genus Rhagoletis in North Amer¬ ica (Diptera: Tephritidae). Bull. Mus. Comp. Zool. 134: 431-562. -. 1966b. Female heterogamety in the family Tephritidae (Acalyptratae, Diptera). Am. Nat. 100: 119-126. -. 1969a. Sympatric host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae). Evolution 23: 237-251. -. 1969b. Mating behavior, host specificity, and the ecological significance of sibling species in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae). Am. Nat. 103 (934): 669-672. -. 1974. The mechanism of sympatric host race formation in the true fruit flies (Tephritidae), pp. 3—23. In M.J.B. White [ed.], Genetic mechanisms of speciation in insects. Australian and New Zealand Book Co., Sydney. Bush, G.L., and M.D. Huettel. 1970. Cytogenetics and description of a new North American species of the Neotropical genus Cecidocharella (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 63: 88—91. Bush, G.L., and G.B. Kitto. 1977. Application of genetics to insect systematics and analysis of species differences, pp. 89—118. In J.A. Romberger et al. [eds.], Biosystematics in agricul¬ ture. Beltsville Symposia in Agricultural Research. Allanheld, Osmun, Montclair, N.J. xii + 340 pp. Bush, G.L., and S.C. Taylor. 1969. The cytogenetics of Procecidochares. I. The mitotic and polytene chromosomes of the Pamakani fly, P. utilis Stone (Tephritidae, Diptera). Caryologia 22: 311-322. Butcher, F.G. 1952. The occurrence of papaya fruit fly in mango. Proc. Fla. State Horde. Soc. 65: 196. Butt, F.H. 1937. The posterior stigmatic apparatus of trypetid larvae. Ann. Entomol. Soc. Am. 30: 487-490. Byers, G.W., F. Blank, W.J. Hansen, D.F. Beneway, and R.W. Frederickson. 1962. Catalog of the types in the Snow Entomological Museum, pt. Ill (Diptera). Kans. Univ. Sci. Bull. 43: 131-181. Caesar, L. 1915. Cherry fruit flies. Entomol. Soc. Ont., 45th Annu. Rep. (1914): 107-112. -. 1917. Insects attacking fruit trees. Ont. Dep. Agric. Bull. 250. 55 pp. -. 1929. The apple maggot outbreak of 1926 to 1928. Entomol. Soc. Ont., 59th Annu. Rep. (1928): 93-95. Caesar, L., and W.A. Ross. 1919. The apple maggot. Ont. Dep. Agric., Fruit Br., Bull. 271. 32 pp. Cameron, E. 1941. The biology and post-embryonic development of Opius illicis n. sp., a parasite of the holly leaf miner (Phytomyza illicis Curt.). Parasitology 33: 8-39. Cameron, P.J., and F.O. Morrison. 1974a. Sampling methods for estimating the abundance and distribution of all life stages of the apple maggot, Rhagoletis pomonella (Diptera: Tephritidae). Can. Entomol. 106: 1025-1034. _. 1974b. Psilus sp. (Hymenoptera: Diaprididae), a parasite of the pupal stage of the apple maggot, Rhagoletis pomonella (Diptera: Tephritidae) in southwestern Quebec. Phy¬ toprotection 55: 13—16. _. 1977. Analysis of mortality in the apple maggot, Rhagoletis pomonella (Diptera: Tephritidae), in Quebec. Can. Entomol. 109: 769-787. Cane, J.H., and F.F. Kurczewski. 1976. Mortality factors affecting Eurosta solidaginis (Dip¬ tera: Tephritidae). J. N.Y. Entomol. Soc. 84: 976—982. Capek, M. 1971. The possibility of control of imported weeds of the genus Solidago L. in Europe. Acta Inst. For. Zvol. 2: 429-441. Carroll, L.E., and R.A. Wharton. 1989. Morphology of the immature stages of Anastrepha ludens (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 82: 201-214.

508

References Cited

Carstenson, G., and H.E. Jaques. 1942. A preliminary study of the insect galls of Iowa. Proc. Iowa Acad. Sci. 49: 525-527. Carter, W. 1952. Recent developments in Oriental fruit fly research. J. Econ. Entomol 45274-279. -. 1953. The Oriental fruit fly, pp. 551-559. In U.S. Dep. Agric. Yearb. for 1952. U.S. Government Printing Office, Washington, D.C. -. 1973. Insects in relation to plant disease. Wiley-Interscience, New York. 759 pp. Castrejon Ayala, F. 1987. Aspectos de la biologia y habitos de Toxotrypana curvicauda Gerst. (Diptera: Tephritidae) en condiciones de laboratono y su distribucion en una plantacion de Carica papaya L. en Yautepec, Mor. Tesis de Biologo, Escuela Nacional de Ciencias Biologicas, Mexico, D.F. 49 pp. Caudell, A.N. 1902. Notes on Colorado insects. U.S. Dep. Agric., Div. Entomol. Bull. 58 (n. ser.): 35-38. Cavalloro, R. 1983. Fruit flies of economic importance. Proceedings of the CEC/IOBC Inter¬ national Symposium, Athens, Greece, 16-19 November 1982. Balkema, Rotterdam. 642 pp. Cavender, G.L., and R.D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Ann. Entomol. Soc Am 75* 400406. -. 1983. On distinguishing Trupanea bisetosa (Coquillett) from T. nigricornis (Coquillett) (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 85: 275-281. -. 1984. The life history of Paracantha cultaris (Coquillett) on wild sunflower, Heliantbus annuus ssp. lenticularis (Douglas) Cockerell, in southern California (Diptera: Tephritidae). Pan-Pac. Entomol. 60: 213-218. Cazier, M.A. 1962. Notes on the bionomics of Zonosemata vittigera (Coquillett), a fruit fly on Solanum. Pan-Pac. Entomol. 38: 181-186. ChagHon, G. 1901. Preliminary list, No. 1, of Canadian Diptera. Entomol. Student 2: 5-8, Chambers, D.L. 1972. Treating tephritids with attractants to enhance their effectiveness in sterile release programs. J. Econ. Entomol. 65: 279-282. -. 1975. Quality in mass-produced insects: definition and evaluation, pp. 19-32. In Con¬ trolling fruit flies by the sterile insect technique. IAEA, Vienna. 175 pp. .1976. Situacion actual de las mosca de la fruta del Caribe, Anastrepba suspensa Loew. Folia Entomol. Mex. 34: 25-35. Champlain, A.B., and J.N. Knull. 1923. Notes on Pennsylvania Diptera. Entomol News 34211-215. Chandler, F.B. 1943. Low-bush blueberries. Maine Agric. Exp. Stn. Bull. 423: 105-130. Chapman, P.J. 1931. Apple maggot studies in 1930. J. Econ. Entomol. 24: 686—691. M933. Viability of eggs and larvae of the apple maggot (Rbagoletis pomonella (Walsh)) at 32°F. N.Y. (Geneva) State Agric. Exp. Stn. Tech. Bull. 206. 19 pp. Chapman, P.J., and O.H. Hammer. 1934. A study of apple maggot control measures. N.Y. State Agric. Exp. Stn. Bull. 644. 40 pp. Chapman, P.J., and A.D. Hess. 1941. Mortality of the apple maggot in fruit held in cold storage. U.S. Dep. Agric. Circ. 600. 9 pp. Chittenden, F.H. 1904. Rbagoletis cingulata, natural history. U.S. Dep. Agric., Div. Entomol Bull. 44: 70-75. -. 1909. Some insects injurious to truck crops. The parsnip leaf miner. The parsley stalk weevil. The celery caterpillar. U.S. Dep. Agric., Bur. Entomol. Bull. 82 (2): 9-24. Ch™,M.L., and J.L. Nation. 1975. The effect of age upon fatty acid composition of adult Caribbean fruit flies, Anastrepba suspensa. Entomol. Exp. Appl. 18: 515-517. Christenson, L.D. 1958. Recent progress in the development of procedures for eradicating and controlling tropical fruit flies. Proc. 10th Int. Congr. Entomol. 3 (1956): 11-16. . 1963. Tropical fruit fly menace. Smithson. Inst. Annu. Rep. for 1962: 441-448.

509

References Cited Christenson, L.D., and R.H. Foote. 1960. Biology of fruit flies. Annu. Rev. Entomol. 5- 171192. Christenson, L.D., L.F. Steiner, and J.W. Balock. 1956. Status of Mediterranean fruit fly research. Mediterranean Fruit Fly Quarantine Fiearing, Washington, D.C., May 9, 1956 (mimeo.). 11 pp. Cirio, U. 1972. Osservazioni sul comportamento di ovideposizione della Rhagoletis completa Cresson (Diptera: Trypetidae) in laboratorio. Proc. Congr. 9th Ital. Entomol. (Siena), pp. 99-117. Clausen, C.P. 1956a. Biological control of fruit flies. J. Econ. Entomol. 49: 176-178. -. 1956b. Biological control of insect pests in the continental United States. U.S. Dep. Agric. Tech. Bull. 1139. 150 pp. Cleveland, M.L., and D. W. Plamilton. 1958. The insect fauna of apple trees in southern Indiana, 1956 and 1957. Ind. Acad. Sci. 68: 205-217. Cockerell, T.D.A. 1889a. Thistle insects. In Colo. Biol. Assoc., 6th Rep. Custer Co. (Colo.) Courant, Jan. 16, 1889. -. 1889b. Contributions towards a list of the fauna and flora of Wet Mountain Valley, Colorado. I. West. Am. Sci. 6(47): 103-106. -. 1889c. Entomological notes from Colorado. Entomol. Mo. Mag. 25: 324. -. 1889d. Notes from Colorado. Entomol. Mo. Mag. 25: 363. -. 1890a. The evolution of insect galls. Entomologist 23: 74-76 (March). -. 1890b. Trypeta bigeloviae, n. sp. Entomol. Mo. Mag. 26: 324 (Dec.). -. 1893a. Trypeta, Clisiocampa and Ammalo. Can. Entomol. 25: 112. -. 1893b. The entomology of the mid-alpine zone of Custer Co., Colorado. Trans Am. Entomol. Soc. 20: 305—370. -. 1898. Contributions to the entomology of New Mexico. II. Some records of Diptera. Davenport Acad. Nat. Sci. 7: 149-156. -. 1899. List of the dipterous galls described from New Mexico. Southwest 1899: 91. -. 1900a. Note on Trypeta notata. J. N.Y. Entomol. Soc. 8: 198. -. 1900b. Scriptotricha or Paracantba? Ann. Mag. Nat. Hist (Ser.7) 5: 400. -. 1902. Some insects of the Hudsonian Zone in New Mexico. VIII. Psyche 4: 346—348. -. 1914. The anatomy of Heliantbus. Entomologist 47: 191-196. -. 1917. The fauna of Boulder Co., Colorado, III. Univ. Colo. Studies 12: 5-20. Cogan, B.H., and H.K. Munro. 1983. Family Tephritidae, pp. 518-554. In R.W. Crosskey et al. [eds.], Catalogue of the Diptera of the Afrotropical Region. Br. Mus. (Nat. Hist.), London. 1,437 pp. Cole, F.R. 1912. Some Diptera of Laguna Beach, pp. 150-162. In First Annu. Rep., Laguna Mar. Lab., Laguna Beach, Fla. -. 1921. Diptera from the Pribiloff Islands, Alaska. Proc. Calif. Acad. Sci., 4th ser. 11(14): 169-177. -. 1923. Diptera from the islands and adjacent shores of the Gulf of California. II. General report. Proc. Calif. Acad. Sci., 4th ser. 12: 457—481. -. 1927. A study of the terminal abdominal structures of male Diptera (two-winged flies). Proc. Calif. Acad. Sci. 16(14): 397—499. Cole, F.R., and A.L. Lovett. 1919. New Oregon Diptera. Proc. Calif. Acad. Sci. 9(7): 221255, pis. 14-19. _. 1921. An annotated list of the Diptera (flies) of Oregon. Proc. Calif. Acad. Sci., 4th ser. 11(15): 197-344. Cole, F.R., and E.I. Schlinger. 1969. The flies of western North America. Univ. Calif. Press, Berkeley. 693 pp. Collin, J.E. 1947. The British genera of Trypetidae, with notes on a few species. Entomol. Rec. J. Var. 59 (suppl.): 1-8. Commonwealth Institute of Entomology. 1951-1960. Distribution maps of insect pests, Series A, London.

510

References Cited

-. -. -. -. -. -. -.

1951. Map No. 1, Ceratitis capitata (Wied.). 1954. Map No. 48, Rhagoletis pomonella (Walsh). 1956. Map No. 64, Dacus (Strumeta) cucurbitae Coq. 1958a. Map No. 88, Anastrepha fraterculus (Wied.). 1958b. Map No. 89, Anastrepha ludens (Lw.). 1958c. Map No. 90, Anastrepha mombinpraeoptans Sein. 1960. Map No. 109, Dacus dorsalis Hend.

Comstock, J.H. 1882. Report on miscellaneous insects. The apple maggot. Rep. Comm. Agric [U.S. Dep. Agric.], 1881-1882, pp. 195-198. Conley, H.J. 1975. Caribfly disrupts exports. Citrus Ind. 56: 12-13. Cook, A.J. 1889. The apple maggot —Trypeta pomonella Walsh. Mich. State Board Agric. 28th Annu. Rep., Sect. 28: 152. Cook, M.T. 1903. Galls and insects producing them. Ohio Nat. 3: 419-436, pis. 13-18. -. 1905. Insect galls of Indiana, pp. 801-867. In Ind. Dep. Geol. Natl. Res. Annu. Rep. 29. 888 pp. -. 1907. The insect galls of Indiana. Proc. Ind. Acad. Sci. 1907: 1-11 (reprint). -.1910. The insect galls of Michigan. Mich. Geol. Biol. Surv. Publ. 1 (Biol. Ser.): 23-33. Coquillett, D.W. 1894. New North American Trypetidae. Can. Entomol. 26: 71-75. -. 1895. Two dipterous leaf miners on garden vegetables. Insect Life 7: 381-384. -. 1899a. Order Diptera, pp. 341-346. In D.S. Jordan, The fur seals and Fur Seal Islands on the North Pacific Ocean, vol. 4 (1898). 384 pp. -. 1899b. A new trypetid from Hawaii. Entomol. News 10: 129-130. -. 1899c. Notes and descriptions of Trypetidae. J. N.Y. Entomol. Soc. 7: 259-268. . 1900. Papers from the Harriman Alaska expedition. IX. Entomological results (3): Diptera. Proc. Wash. Acad. Sci. 2: 389-464. -. 1902a. New Diptera from North America. Proc. U.S. Natl. Mus. 35: 83-126. -. 1902b. New Acalyptrate Diptera from North America. J. N.Y. Entomol. Soc 10- 177191. —. 1904. Diptera from southern Texas with descriptions of new species. J. N.Y Entomol Soc. 12: 31-35. —. 1908. New genera and species of Diptera. Proc. Entomol. Soc. Wash. 9 (1907)* 144148. -. 1910. The type-species of the North American genera of Diptera. Proc. U.S. Natl Mus 37(1719): 499-647. Cordley, A.B. 1889. The apple maggot. Orchard and Garden 11: 192. Cosens, A. 1912. A contribution to the morphology and biology of insect galls. Toronto Univ. Studies, Biol. Ser. 13: 297-387. Couper, W. 1864. List of Coleoptera and Diptera taken at Quebec and other parts of L. Canada. Trans. Lit. Hist. Soc. Que. 1864: 3-21. Cowperthwaite, W.G. 1962. The medfly strikes the third time. Proc. Fla. State Hortic Soc 7519-20. Cox, J.A. 1952. The cherry fruit fly in Erie County. Pa. Agr. Exp. Stn. Bull. 548. 17 pp. Crandell, H.A. 1939. The biology of Pachycrepoideus duhius Ashmead, a pteromalid parasite of Piophila caseyi Linne. Ann. Entomol. Soc. Am. 39: 632-654. Crawford, D.L. 1910. The Mexican orange maggot. Pomona Coll. J. Entomol. 2: 321-332. . 1913. Control of the orange maggot (Trypeta ludens). Mex. Gulf Coast Citrus Assoc. Circ. 1.16 pp. . 1927. Investigation of Mexican fruit fly (Anastrepha ludens Loew) in Mexico. Calif. Dep. Agric. Mo. Bull. 16: 421-446. Cresson, E.T., Jr. 1907. Some North American Diptera from the southwest (Paper II). Trans. Am. Entomol. Soc. 33: 99-198, pi. 1. . 1914a. Some nomenclatorial notes on the dipterous family Trypetidae. Entomol News 25: 275-279.

References Cited

-. 1914b. More nomenclatorial notes on Trypetidae (Diptera). Entomol. News 25: 323. . 1920. Descriptions of new North American acalyptrate Diptera. II Sapromyzidae). Entomol. News 31: 65-67.

(Trypetidae,

-. 1929. A revision of the North American fruit flies of the genus Rhagoletis. Trans. Am. Entomol. Soc. 55(935): 401-414, pi. XVI. . 1931. Notes on the abstersa-group of the genus Tephritis, and a description of a new species from California. Entomol. News 42: 3-5. Crevecoeur, F.F. 1906. Additions to the list of Kansas Diptera. Trans. Kans. Acad. Sci. 20(1): 90-96. Crnjar, R.M., and R.J. Prokopy. 1982. Morphological and electrophysiological mapping of tarsal chemoreceptors of oviposition-deterring pheromone in Rhagoletis pomonella flies. J. Insect Physiol. 28: 393-400. Crnjar, R.M., R.J. Prokopy, and V.G. Dethier. 1979. Electrophysiological identification of oviposition-deterring pheromone receptors in Rhagoletis pomonella (Diptera: Tephritidae). J. N.Y. Entomol. Soc. 86 (1978): 283-284. Cunningham, R.I. 1966. Sex identification of pupae of three species of fruit flies (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 59: 864-865. Curran, C.H. 1923. Two varieties of Eurosta solidaginis Fitch. Entomol. News 34: 302. -. 1924a. Rhagoletis symphoricarpi, a new trypaneid from British Columbia. Can. Ento¬ mol. 56: 62—63. -. 1924b. Rhagoletis pomonella and two allied species. Entomol. Soc. Ont., Annu. Rep. 33: 56-57. -. 1925. Note on the rearing of Eurosta solidaginis and variety fascipennis (Trypaneidae, Dipt.). Can. Entomol. 57: 128—129. -. 1928a. Diptera or two-winged flies. In Insects of Puerto Rico and the Virgin Islands. Sci. Surv. Porto Rico and Virgin Is. 11(1): 1—118. -. 1928b. Diptera. In The Entomological Record, 1927. Entomol. Soc. Ont., Annu. Rep. 58: 99-101. -. 1930. Report on the Diptera collected at the station for the study of insects, Harriman Interstate Park, N.Y. Bull. Am. Mus. Nat. Hist. 61(1931): 21—115. -. 1932a. New North American Diptera, with notes on others. Am. Mus. Novit. 526: 1— 13. -. 1932b. New species of Trypaneidae, with key to the North American genera. Am. Mus. Novit. 556: 1-19. -. 1934. The families and genera of North American Diptera. C.H. Curran (privately printed). Currie, G.A. 1932. Oviposition stimuli of the burr-seed fly, Euaresta aequalis Loew (Diptera: Trypetidae). Bull. Entomol. Res. 23: 191 — 193, pi. VIII. Daecke, E. 1906. Minutes, Feldman Collecting Social. Tephritis platyptera. Entomol. News 17: 312. -. 1910. Trypetid galls and Eurosta elsa n. sp. Entomol. News 21: 341-343, pi. X. d’Araujo e Silva, A., C.N. Goncalves, A.N. Galvao, A.J.L. Goncalves, J. Gomes, M. do Nascimento Silva, and L. de Simoni. 1968. Quarto catalogo des insectes que vivem nas plantas do Brasil. Parte II, 1° Tomo. Ministerio da Agricultura, Laboratorio Cantral de Patalogia Vegetal, Rio de Janeiro. Darby, H.H., and E.M. Kapp. 1934. Studies on the Mexican fruit fly, Anastrepha ludens (Loew). U.S. Dep. Agric. Tech. Bull. 444. 20 pp. Darlington, E.P. 1943. “Bred” or “reared” and note on the blueberry fruit fly. Entomol. News 54: 217-219. Davis, G.C. 1889. A new departure by the apple maggot —Trypeta pomonella. Ohio Farmer 76:291. Davis, J.C., H.R. Agee, and E.A. Ellis. 1983. Comparative ultrastructure of the compound eye

512

References Cited

of the wild, laboratory reared, and irradiated Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 76: 322-332. Davis, W.T. 1931. The elliptical goldenrod gall, its maker, and destroyer. Bull. Brooklyn Entomol. Soc. 26: 120-123. Dean, R.W. 1932. The alimentary canal of the apple maggot, Rhagoletis pomonella Walsh. Ann. Entomol. Soc. Am. 25: 210-223. -. 1933. Morphology of the digestive tract of the apple maggot fly, Rhagoletis pomonella Walsh. N.Y. Agric. Exp. Stn. Tech. Bull. 215. 17 pp. -. 1935a. Anatomy and postpupal development of the female reproductive system of the apple maggot fly, Rhagoletis pomonella Walsh. N.Y. Agric. Exp. Stn. Tech. Bull. 229. 31 pp. -. 1935b. Preoviposition period of the apple maggot fly, Rhagoletis pomonella Walsh, in eastern New York. J. Econ. Entomol. 28: 504. -. 1938. Experiments on rearing apple maggot adults. J. Econ. Entomol. 31: 241-244. -. 1941. Attraction of Rhagoletis pomonella adults to protein baits. J. Econ. Entomol 34123. -. 1942a. Development of the female reproductive system in Rhagoletis pomonella Walsh. Ann. Entomol. Soc. Am. 35: 397-410. -. 1942b. Emergence, migration and control of the apple maggot fly, pp.291-295. In 87th Annu. Mtg., N.Y. Hortic. Soc. -. 1969. Infestation of peaches by Rhagoletis suavis. J. Econ. Entomol. 62: 940-941. Dean, R.W., and P.J. Chapman. 1973. Bionomics of the apple maggot in eastern New York. Search (Cornell Univ.) 3(10): 1-64. Denmark, H.A. 1956. The Mediterranean fruit fly infests Florida again. Fla Entomol 39* 85-87. Detmers, F. 1927. Canada thistle, Cirsium arvense Tourn. Field thistle, creeping thistle. Ohio Agric. Exp. Stn. Bull. 414. 56 pp. Dickens, J.C., E. Solis, and W.G. Hart. 1982. Sexual development and mating behavior of the Mexican fruit fly, Anastrepha ludens (Loew). Southwest. Entomol. 7: 9-15. Diehl, S.R., and G.L. Bush. 1983. The use of trace element analysis for determining the larval host plant of adult Rhagoletis, pp. 276-284. In R. Cavalloro [ed.], Fruit flies of economic importance. Proceedings of the CEC/IOBC International Symposium, Athens, Greece, 16— 19 November 1982. Balkema, Rotterdam. Dirks, C.O. 1935. Larval production and adult emergence of the apple fruit fly in relation to apple varieties. J. Econ. Entomol. 28: 198-203. Doane, R.W. 1898. A new trypetid of economic importance. Entomol. News 9: 69-72. -. 1899. Notes on Trypetidae with descriptions of new species. J. N.Y. Entomol Soc 7177-193, pis. Ill, IV. -. 1900. Additional notes on Trypetidae. J. N.Y. Entomol. Soc. 8: 47-48. Dodson, G. 1978. Morphology of the reproductive system in Anastrepha suspensa (Loew) and notes on related species. Fla. Entomol. 61: 231-239. -. 1982. Mating and territoriality in wild Anastrepha suspensa (Loew) Tephritidae) in field cages. J. Ga. Entomol. Soc. 17: 189-200.

(Diptera:

. 1986. Lek mating system and large male aggressive advantage in a gall-forming tephritid (Diptera: Tephritidae). Ethology 72: 99-108. . 1987a. Host-plant records and life history notes on New Mexico Tephritidae (Diptera). Proc. Entomol. Soc. Wash. 89: 607-615. -. 1987b. Biological observations on Aciurina trixa and Valentibulla dodsoni (Diptera: Tephritidae) in New Mexico. Ann. Entomol. Soc. Am. 80: 494-500. Dodson, G., and S.B. George. 1986. Examination of two morphs of gall-forming Aciurina (Diptera: Tephritidae). Ecological and genetic evidence for species. Biol. I. Linn Soc 2963-79. ... Downes, W. 1919. Popular and practical entomology. The apple maggot in British Columbia. Can. Entomol. 51: 2-4.

513

References Cited

Drew, R.A.I. 1972. The generic and subgeneric classification of Dacini (Diptera: Tephritidae) from the South Pacific area. J. Aust. Entomol. Soc. 11: 1—22. -. 1974. Revised descriptions of species of Dacini (Diptera: Tephritidae) from the South Pacific area. II. The Strumeta group of subgenera of genus Dacus. Queensl. Dep. Prim. Ind., Div. Plant Ind. Bull. 653. 101 pp. -. 1989a. The taxonomy and distribution of tropical and subtropical Dacinae, pp. 9—14. In A.S. Robinson and G. Hooper [eds.], Fruit flies. Their biology, natural enemies and control, vol. 3A. Elsevier, New York. 372 pp. -. 1989b. The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Queensland Mus. Mem. 26. 521 pp. Drew, R.A.I., G.S.H. Hooper, and M.A. Bateman. 1978. Economic fruit flies of the South Pacific region. Watson, Ferguson, Brisbane. 137 pp. Eaton, E.L., and I.V. Hall. 1961. The blueberry in the Atlantic provinces. Can. Dep. Agric. Publ. 754. 35 pp. Ebeling, W. 1959. Subtropical fruit pests. Univ. Calif., Div. Agric. Sci. 463 pp. Eden, W.G. 1970. Research on Caribbean fruit fly at University of Florida. Int. Atomic Energy Agency, Proc. Ser., Symp., pp. 150—151. Eichmann, R.D. 1936. The cherry aphis and cherry maggot of the Flathead Valley. Mont. Agric. Exp. Stn. Bull. 313. 11 pp. Eide, P.M. 1946. The cherry fruit fly problem in eastern Washington. Proc. Wash. State Hortic. Assoc. (1945): 55—61. -. 1947. Cherry fruit fly control in the lower Yakima Valley. Proc. Wash. State Hortic. Assoc. (1946): 119-128. Eide, P.M., F.T. Lynd, and H.S. Telford. 1949. The cherry fruit fly in eastern Washington. Wash. Agric. Exp. Stn. Circ. 72. 8 pp. Ellertson, F.E. 1961. New host records for cherry fruit fly in Oregon. Pan-Pac. Entomol. 37: 116. Elzinga, R.A., and A.B. Broce. 1986. Labellar modifications of Muscomorpha flies. Ann. Entomol. Soc. Am. 79: 150—209. Emmart, E.W. 1933. The eggs of four species of fruit flies of the genus Anastrepba. Proc. Entomol. Soc. Wash. 35: 184-191, pis. 7, 8. -. 1935. Studies of the chromosomes of Anastrepba. 1. The chromosomes of the fruit fly, Anastrepba ludens Loew. Proc. Entomol. Soc. Wash. 37: 119—136, pis. 11 — 16. Essig, E.O. 1938. Insects of western North America, 1st ed. Macmillan, New York. 1,035 pp. -. 1958. Insects and mites of western North America, 2nd ed. Macmillan, New York. 1,050 pp. Evenhuis, N.L. [ed.]. 1989a. Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Museum Special Publication 86. Bishop Museum Press, Honolulu, Hawaii and E.J. Brill, Leiden. 1,155 pp. -. 1989b. Tephritidae, pp. 2-4. In N.L. Evenhuis [ed.], Pacific Basin Diptera News, issue 1. Bishop Museum Press, Honolulu, Hawaii. Fabricius, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species adiectis synonymis, locis, descriptiombus, observationibus. Flensburg & Leipzig. 832 pp. -. 1794. Entomologia systemica emendata et aucta. 4. Copenhagen. 472 pp. -. 1805. Systema antliatorum secundum ordines, genera, species adjectis synonymis, locus, observationibus, descriptionibus. Brunswick. 403 pp. Fallen, C.F. 1814. Beskrifning ofver de i Sverige funne tistel-flugor, horande till Dipter-slogtet Tepbritis. K. Svensk. Vetenskakad. Handl. 35: 156—177. -. 1826. Supplementum 2. Dipterorum Sveciae. Lund, pp. 9-16. Farleman, M.G. 1932. Note on cherry fruit fly oviposition. J. Econ. Entomol. 5: 1048. -. 1933. Observations on the habits of flies belonging to the genus Rbagoletis. J. Econ. Entomol. 26: 825—828.

514

References Cited Felt, E.P. 1898. Trypeta canadensis Loew, gooseberry fruit fly. N.Y. State Mus. Bull. 5: 160163. -. 1918. Key to American insect galls. N.Y. State Mus. Bull. 200. 310 pp. ——. 1940. Plant galls and gall makers. Comstock, Ithaca, N.Y. viii + 364 pp. Field, A. 1904. Trypeta solidaginis. Entomol. News 15: 309. Finney, G.L. 1953. Oriental fruit fly studies. Calif. Agric. 7(6): 11-12. Fischer, M. 1964. Opiinae obszaru nearktycznego (Hymenoptera: Bracomdae). Czecs I. Pol. Pismo Entomol. 34: 197-530. Fitch, A. 1855. Report on the noxious, beneficial and other insects of the state of New York I N.Y. Agric. Soc. Trans. (1854) 14: 705-880. . 1856. First and second report on noxious, beneficial and other insects of the state of New York. Albany. 176 pp.. Fleschner, C.A. 1963. Releases of recently reported insect parasites and predators in Califor¬ nia, 1960-1961. Pan-Pac. Entomol. 39: 114-116. Fleury, A.C. 1932. Mexican fruit fly infestation in the Lower Rio Grande Valley Texas. Calif Dep. Agric. Mo. Bull. 21: 316-321. Flint, C.L. [ed.]. 1890. A treatise on some of the insects injurious to vegetation, by Thaddeus William Harris, M.D. Orange Judd Co., N.Y., 640 pp, 8 pis. Fhtters, N.E., and P.S. Messenger. 1958. Effect of temperature and humidity on development and distribution of Hawaiian and Mexican fruit flies. J. Rio Grande Valley Hortic. Soc. 12: Flitters, N.E., P.S. Messenger, and C.N. Husman. 1956. Bioclimatic cabinets used in studies on the Mexican fruit fly and pink bollworm. U.S. Dep. Agric. ARS-33-33. 12 pp. Fluke, C.L., Jr., and T.C. Allen. 1931. The role of yeast in life history studies of the apple maggot, Rhagoletis pomonella. J. Econ. Entomol. 24: 77-80. Foote, B.A. 1965. Biology and immature stages of eastern ragweed flies (Tephritidae) Proc No. Centr. Br., Entomol. Soc. Am. 20: 105-106. ——.1967. Biology and immature stages of fruit flies: The genus Icterica (DipteraTephritidae). Ann. Entomol. Soc. Am. 60: 1295-1305. ——. 1984. Host plant records for North American ragweed flies (Diptera: Tephritidae) Entomol. News 95(2): 51-54. Foote, R.H. 1958. The genus Euarestoides in the United States and Mexico. Ann Entomol Soc. Am. 51: 288-293. . 1959a. A new North American species of Tephritis, with some observations on its generic position. Bull. Brooklyn Entomol. Soc. 54: 13-17. . 1959b. Notes on the genus Euleia Walker in North America. J. Kans. Entomol. Soc. 32: 14 5 15 0 • . 1959c. A new synonymy in the Tephritidae. Proc. Entomol. Soc. Wash. 61: 59. —. 1960a. Paraterellia, a new genus of Tephritidae from the western United States Ann Entomol. Soc. Am. 53: 121-125. 253—260.

The §enus TryPeta in America north of Mexico. Ann. Entomol. Soc. Am. 53:

—. 1960C A new North American fruit fly genus, lephritidae). Ann. Entomol. Soc. Am. 53: 671-675.

Procecidocbaroides (Diptera:

—. 1960d. A new tephritid genus, Rhagoletoides, with notes on its distribution and systematic position. Entomol. News 71: 145-149. —. 1960e. The genus Tephritis Latreille in the Nearctic Region north of Mexico: Descrip¬ tions ol tour new species and notes on others. J. Kans. Entomol. Soc. 33: 71-85. —. 1960f. The species of the genus Neotephritis Hendel in America north of Mexico 1 N Y Entomol. Soc. 68: 145-151. ' T,,19D60.^ A revision of the genus Trupanea in America north of Mexico. U.S. Dep. Agric lech. Bull. 1214. 29 pp. b

515

References Cited

-. 1960h. Notes on some North American Tephritidae, with descriptions of two new genera and two new species. Proc. Biol. Soc. Wash. 73: 107—117. -. 1961. Eurostina Curran, a new synonym. Proc. Entomol. Soc. Wash. 63: 28. -. 1962. The types of North American Tephritidae in the Snow Museum, the University of Kansas. J. Kans. Entomol. Soc. 35: 170-179. -. 1964a. A new synonym in the genus Eurosta (Tephritidae). Proc. Entomol. Soc. Wash. 66: 61. -. 1964b. Acinia picturata (Snow), a resurrected synonym. Proc. Entomol. Soc. Wash. 66: 84. -. 1964c. A review of the genus Xanthomyia Phillips. Proc. Entomol. Soc. Wash. 66: 140. -. 1964d. Notes on the Walker types of New World Tephritidae (Diptera). J. Can. Entomol. Soc. 37: 316-326. -. 1965a. Family Tephritidae, pp. 658-678. In A. Stone et al. [eds.], A catalog of the Diptera of America North of Mexico. U.S. Dep. Agric. Agric. Handbook 276. 1,696 pp. -. 1965b. A study of the types of Tephritidae described by F.M. van der Wulp in “Biologia Centrali-Americana” (Diptera). J. Kans. Entomol. Soc. 38: 236-247. -. 1966a. The genus Rhagoletotrypeta, with a new Nearctic species. Ann. Entomol. Soc. Am. 59: 803-807. -. 1966b. Notes on the types of Tephritidae described by R.W. Doane (Diptera). Proc. Entomol. Soc. Wash. 68: 120—126. -. 1967a. Some Texas records of the genus Hexacbaeta Loew (Diptera: Tephritidae) Ann. Entomol. Soc. Am. 60: 1327—1328. -. 1967b. Family Tephritidae, No. 57, pp. 57.1-57.91. In M. Vanzolini [ed.], A catalog of the Diptera of the Americas south of the United States. Dep. Zool., Seer. Agric., Sao Paulo, Brazil. -. 1980. Fruit fly genera south of the United States. U.S. Dep. Agric. Tech. Bull. 1600. 77 pp. -. 1981. The genus Rhagoletis Loew south of the United States (Diptera: Tephritidae). U.S. Dep. Agric. Tech. Bull. 1607. 75 pp. -. 1984. Family Tephritidae, pp. 66—188. In A. Soos and L. Papp [eds.], Catalog of Palaearctic Diptera, vol. 9 (Micropezidae-Agromyzidae). Akademiai Kaido, Budapest, Hun¬ gary. 460 pp. Foote, R.H., and F.L. Blanc. 1959. A new genus of North American fruit flies. Pan-Pac. Entomol. 35: 149—156. -. 1963. The fruit flies or Tephritidae of California. Calif. Insect Surv. Bull. 7: 1-117. -. 1979. New species of Tephritidae from the United States, Mexico, and Guatemala, with revisionary notes. Pan-Pac. Entomol. 55: 161—179. Foote, R.H., and A. Freidberg. 1980. The taxonomy and nomenclature of some Palaearctic Tephritidae. J. Wash. Acad. Sci. 70: 29—34. Foote, R.H., and G.C. Steyskal. 1987. Tephritidae, pp. 817-831. In J.F. McAlpine et al. [coords.], Manual of Nearctic Diptera, vol. 2. Res. Br., Agric. Can. Mon. 27. 1,332 pp. Foott, W.H. 1963. The biology and control of the pepper maggot, Zonosemata electa (Say), in southwestern Ontario. Proc. Entomol. Soc. Ont. 93 (1962): 75—81. -. 1968a. The importance of Solanum carolinense L. as a host of the pepper maggot, Zonosemata electa (Say) in southwestern Ontario. Proc. Entomol. Soc. Ont. (1967) 98: 16— 19. -. 1968b. Laboratory rearing of the pepper maggot, Zonosemata electa (Say). Proc. Ento¬ mol. Soc. Ont. (1967) 98: 18—21. Foott, W.H., and P.R. Timmons. 1970. Pepper insects in southwestern Ontario. Proc. Ento¬ mol. Soc. Ont. (1969) 100: 105-110. Forsell, M.J. 1947. The Tephritidae of the Puget Sound region. Northwest Sci. 21: 167.

516

References Cited

Foster, D.E., and S.K. Benbow. 1978. A gall-forming fly attacks sagebrush. Range Wildl. Mgmt. 8:48. Foxlee, H.R. 1957. Diptera taken at Robson, B.C. Proc. Entomol. Soc. B.C. 53 (1956): 3439. Frank, A. 1924. The cherry fruit fly. Proc. Wash. State Hortic. Soc. (1923): 42-45. Frauenfeld, G.R. von. 1857. Beitrage zur Naturgeschichte der Trypeten nebst Beschreibung einiger neue Arten. Sitzungber. K.-Akad. Wiss. Math-Naturw. Classe, Wien 22: 523-557. Freidberg, A. 1981. Mating behavior of Tephritidae). Israel J. Entomol. 15: 89-95.

Schistopterum

moebiusi

Becker

(Diptera:

-. 1984. Gall Tephritidae (Diptera), pp. 129-167. In T.N. Ananthakrishnan [ed.l, Biologv of gall insects. Oxford & IBH, New Delhi. -. 1985. The genus Craspedoxantha Bezzi (Diptera: Tephritidae: Terelliinae). Ann. Natal Mus. 27: 183-206. -. 1987. Ortbocanthoides aristae, a remarkable new genus and species of Tephritidae (Diptera) from Mt. Kenya. Ann. Natal Mus. 28: 551-559. Freidberg, A., and W.N. Mathis. 1986. Studies of Terelliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithson. Contr. Zool. 439. 75 pp. Frey, R. 1945. Tiergeographischen Studien uber die Dipterenfauna der Azoren. Soc. Sci. Fenn Comm. Biol. 8(10): 1-114. Frias, D.A. 1981. Biologia evolutiva de dipteros Otitidae y Tephritidae (Diptera Acalvotratae) Ph.D. thesis, Universidad de Chile. 243 pp. -. 1985. Quatro nuevas especies chilenas delo genero Tephritidae). Rev. Bras. Zool. 2: 363-381.

Trupanea Schrank (Diptera

-. 1988. Tephritis marisolae, nueva especie chilena del genero Tepbritis Latreille (DipteraTephritidae). Rev. Chil. Entomol. 16: 77-81. Frick, K.E. 1949. The present status of the cherry fruit fly in the Yakima Valley. Proc. Wash State Fiortic. Assoc. 45: 99-110. -. 1952. Determining emergence of the cherry fruit fly with ammonium carbonate bait traps. J. Econ. Entomol. 45: 262-263. . 1953. Inspection methods to detect cherry fruit fly larvae in cherries. J. Econ Entomol 46: 893-894. . 1957. Further biological studies on the western cherry fruit fly. J. Econ. Entomol. 50: . 1964. Some endemic insects that feed upon introduced tansy ragwort (Senecio jacobaea) in western United States. Ann. Entomol. Soc. Am. 57: 707-710. -. 1971. The biology of Trypeta angustigena Foote in central and coastal California - host plants and notes (Diptera: Tephritidae). J. Wash. Acad. Sci. 61: 20-24. . 1972. Third list of insects that feed upon tansy ragwort, Senecio jacobaea, in the western United States. Ann. Entomol. Soc. Am. 65: 629-631. Frick, K.E., and R.B. Hawkes. 1970. Additional insects that feed upon tansy ragwort, Senecio

jacobeae, an introduced weedy plant, in the western United States. Ann. Entomol. Soc. Am 63: 1085—1089. Frick, K.E., Fl.G. Simkover, and H.S. Telford. 1954. Bionomics of the cherry fruit fly in eastern Washington. Wash. Agric. Exp. Stn. Tech. Bull. 3. 66 pp. Fnngs, H., and M. Frings. 1955. The location of the contact chemoreceptors of the cherry fruit 435^^^°^?/S C*ngUlata’ an 31 23 2,« -. 1967a. Wooden apples lure costly pest. Front. Plant Sci. 19: 4-5. . 1967b. Factors influencing effectiveness of artificial oviposition devices for apple mag¬ got. J. Econ. Entomol. 60: 950-955. . 1967c. Artificial diet for apple maggot larvae. J. Econ. Entomol. 60: 1161-1162. . 1968a. Visual responses of apple maggot flies, Rhagoletis pomonella (Diptera: Tephritidae): orchard studies. Entomol. Exp. Appl. 11: 403-422. . 1968b. Sticky spheres for estimating apple maggot adult abundance. J. Econ. Entomol. 61: 1082—1085. . 1968c. Influence of photoperiod, temperature, and food on initiation of diapause in the apple maggot. Can. Entomol. 100: 318-329. . 1972a. Evidence for a marking pheromone deterring repeated oviposition in adult apple maggot flies. Environ. Entomol. 1: 326-332. ——. 1972b. Response of apple maggot flies to rectangles of different colors and shades. Environ. Entomol. 1: 720-726. ——. 1973. Dark enamel spheres capture as many apple maggot flies as fluorescent spheres Environ. Entomol. 2: 953-954. . 1975a. Apple maggot control by sticky red spheres. J. Econ. Entomol. 68: 197-198. ——. 1975b. Mating behavior in Rhagoletis pomonella (Diptera: Tephritidae). V Virgin female attraction to male odor. Can. Entomol. 107: 905-908. . 1975c. Oviposition-deterring fruit marking pheromone in Rhagoletis fausta. Environ Entomol. 4: 298-300. ——. 1975d. Selective new trap for Rhagoletis cingulata and R. pomonella flies. Environ Entomol. 4: 420-424. —1976a. Feeding, mating, and oviposition activities of Rhagoletis fausta flies in nature Ann. Entomol. Soc. Am. 69: 899-904. ~j 1976b. Sticky spheres for trapping apple maggot flies. Mass. Fruit Notes 41(6): —. 1977a. Attraction of Rhagoletis flies (Diptera: Tephritidae) to red spheres of different sizes. Can. Entomol. 109: 593-596. -—. 1977b. Stimuli influencing trophic relations in Tephritidae, No. 265, pp. 305-336. In omportement des insectes et milieu trophique (Colloques Internationaux de Centre Na¬ tional de la Recherche Scientifique). —. 1980. Mating behavior of frugivorous Tephritidae, pp. 37-46. In T. Iwata [ed.l, Proceedmgs ol a Symposium on Fruit Fly Problems, Kyoto and Naha, Aug. 9-12 1980. Natl. Inst Agric. Sci., Yatabe, Ibaraki 305, Japan. 1^la; OvlPositl°n-deterring pheromone system of apple maggot flies, pp. 477-494 In ’iooiUTL wd Mana§ement of msect pests with semiochemicals. Plenum, New York. —. 1981b. The Mediterranean fruit fly in Massachusetts: can it happen? Fruit Notes, Coop. Ext. Serv., Umv. Mass. 46(4): 5-8. —. 1981c. Epideictic pheromones that influence spacing patterns of phytophagous insects,

539

References Cited

pp. 181-213. In D.A. Nortduld [ed.], Semiochemicals: their role in pest control. Wiley, New York. -. 1982. Getting to know a fruit fly. Ga. Entomol. 17(4)(2nd Suppl.): 30-38. Prokopy, R.J., and S.H. Berlocher. 1980. Establishment of Rhagoletis pomonella (Diptera: Tephritidae) on rose hips in southern New England. Can. Entomol. 12: 1319-1320. Prokopy, R.J., and G.L. Bush. 1972a. Mating behavior in Rhagoletis pomonella (Diptera: Tephritidae). III. Male aggregation in response to an arrestant. Can. Entomol. 104: 275283. -. 1972b. Apple maggot infestation of pear. J. Econ. Entomol. 65: 597. -. 1973a. Mating behavior of Rhagoletis pomonella (Diptera: Tephritidae). IV. Courtship. Can. Entomol. 105: 873—891. -. 1973b. Ovipositional responses to different sizes of artificial fruit by flies of the Rhagoletis pomonella species group. Ann. Entomol. Soc. Am. 66: 927—929. -. 1973c. Oviposition by grouped and isolated apple maggot flies. Ann. Entomol. Soc. Am. 66: 1197-1200. Prokopy, R.J., and W.M. Coli. 1978. Selective traps for monitoring Rhagoletis mendax flies. Protect. Ecol. 1: 45—53. Prokopy, R.J., and K.I. Hauschild. 1979. Comparative effectiveness of sticky red spheres and Pherocon AM standard traps for monitoring apple maggot flies in commercial orchards. Environ. Entomol. 8: 696—700. Prokopy, R.J., and J. Hendrichs. 1979. Mating behavior of Ceratitis capitata on a field-caged host tree. Ann. Entomol. Soc. Am. 72: 642—648. Prokopy, R.J., and E.D. Owens. 1978. Visual generalist visual specialist phytophagous insects: host selection behavior and application to management. Entomol. Exp. Appl. 24: 409—420. Prokopy, R.J., and P.J. Spatcher. 1977. Location of receptors for oviposition-deterring phe¬ romone in Rhagoletis pomonella flies. Ann. Entomol. Soc. Am. 70: 960—962. Prokopy, R.J., and R.P. Webster. 1978. Oviposition-deterring pheromone of Rhagoletis pomonella. A kairomone for its parasitoid Opius lectus. J. Chem. Ecol. 4: 481—494. Prokopy, R.J., E.W. Bennett, and G.L. Bush. 1971. Mating behavior in Rhagoletis pomonella (Diptera: Tephritidae). 1. Site of assembly. Can. Entomol. 103: 1405—1409. -. 1972. Mating behavior in Rhagoletis pomonella (Diptera: Tephritidae). II. Temporal organization. Can. Entomol. 104: 97—104. Prokopy, R.J., V. Moericke, and G.L. Bush. 1973. Attraction of apple maggot flies to odor of apples. Environ. Entomol. 2: 743—749. Prokopy, R.J., W.H. Reissig, and V. Moericke. 1976. Marking pheromones deterring repeated oviposition in Rhagoletis flies. Entomol. Exp. Appl. 20: 170—178. Prokopy, R.J., P.D. Greany, and D.L. Chambers. 1977. Oviposition-deterring pheromone in Anastrepha suspensa. Environ. Entomol. 6: 463—465. Prokopy, R.J., A.L. Averill, C.M. Bardinelli, E.S. Bowdan, S.S. Cooley, R.M. Crnjar, E.A. Dundulis, C.A. Roitberg, P.J. Spatcher, J.H. Tumlinson, and B.L. Weeks. 1982a. Site of production of an oviposition-deterring pheromone component in Rhagoletis pomonella flies. J. Insect Physiol. 28: 1 — 10. Prokopy, R.J., A.L. Averill, S.S. Cooley, and C.A. Roitberg. 1982b. Associative learning in egglaying site selection by apple maggot flies. Science 218: 76—77. Proverbs, M.D. 1953. Note on the occurrence of Rhagoletis fausta (O.S.) (Diptera: Tephritidae) in the Okanagan Valley. Proc. Entomol. Soc. B.C. 49(1952): 18. Quaintance, A.L. 1908. The apple maggot or “railroad worm.” U.S. Dep. Agric. Bur. Ento¬ mol. Circ. 101. 12 pp. Quayle, H.J. 1931. Possible future distribution and abundance of the Mediterranean fruit fly [Ceratitis capitata Wied.) in the United States. J. Econ. Entomol. 24: 1064-1066. -. 1938. Insects of citrus and other subtropical fruits. Comstock, Ithaca, N.Y. 583 pp. Quisenberry, B.F. 1949a. The genus Oxyna in the Nearctic Region north of Mexico. Pan-Pac. Entomol. 25: 71—76.

References Cited

-. 1949b. A new genus of Tephritidae near Xanthomyia. Bull. Brooklyn Entomol Soc 4449-52. * -. 1949c. Notes and descriptions of North American Tephritidae. I. Kans. Entomol Soc 22: 81-88. -. 1950. The genus Euaresta in the United States (Diptera: Tephritidae). J. N.Y Entomol Soc. 58: 9-38. -. 1951. A study of the genus Tephritis Latreille in the Nearctic Region north of Mexico (Diptera: Tephritidae). J. Kans. Entomol. Soc. 24: 56-72. Raine, J., and H. Andison. 1958. Life histories and control of cherry fruit flies on Vancouver Island, British Columbia. J. Econ. Entomol. 51: 592-595. Ratner, S.S., and J.G. Stoffolano, Jr. 1982. Development of the oesophageal bulb of the apple

Rbagoletis pomonella (Diptera: Tephritidae): morphological, histological, and histochemical study. Ann. Entomol. Soc. Am. 75: 555-562. -. 1984. Ultrastructural changes of the oesophageal bulb of the adult female apple maggot, Rbagoletis pomonella (Walsh) (Diptera: Tephritidae). Int. J. Insect Morphol. Embrvol 13* 191-208. 7 ‘ ' Rau, P. 1922. Ecological and behavior notes on Missouri insects. Trans. St. Louis Acad Sci 24(7): 1-71. * * Redfern, M. 1968. The natural history of spear thistle heads. Field Studies 2: 669-717. Reissig, W.H. 1974a. Field tests of traps and lures for the apple maggot. ]. Econ. Entomol 67484-486. * * -. 1974b. Field tests of the response of Rbagoletis pomonella to apples. Environ. Entomol. 3: 733—736. . 1975a. Evaluation of traps for apple maggot in unsprayed and commercial apple or¬ chards. J. Econ. Entomol. 68: 445-448. . 1975b. Performance of apple maggot traps in various apple tree canopy positions. I. Econ. Entomol. 68: 534-538. . 1976. Comparison of traps and lures for Rbagoletis fausta and Rbagoletis cingulata. I. Econ. Entomol. 69: 634-643. . 1977. Capture of released apple maggot flies, Rbagoletis pomonella Tephritidae) on sticky panels. Can. Entomol. 109: 631-634.

(Diptera:

Reissig, W.H., and D.C. Smith. 1978. Bionomics of Rbagoletis pomonella in Crataegus. Ann. Entomol. Soc. Am. 71: 155-159. Rhode, R.H. 1976. The history and current status of the Mediterranean fruit fly in North America. J. Rio Grande Valley Hortic. Soc. 30: 11-17. Rhode, R.H., and M. Sanchez R. 1982. Field evaluation of McPhail and gallon plastic tub traps against the Mexican fruit fly. Southwest. Entomol. 7: 98-100. Ride, W.D.L. [ed.]. 1985. International code of zoological nomenclature, 3rd ed. International Trust for Zoological Nomenclature, London. 338 pp. Riedl, H., R.G. Hyslop, W.W. Barnett, W.W. Coates, L.B. Fitch, J.L. Joos, W.H. Olson, J.C. Profita, and W.R. Schreader. 1981. New monitoring methods for the walnut husk fly Calif Agric. 35(9/10): 21-22. Ries, D.T. 1927. The apple maggot in Michigan. J. Econ. Entomol. 20: 144-146. . 1935. Biological study of the walnut husk fly (Rbagoletis suavis Loew). Pap Mich Acad. Sci. Arts Lett. 20(1934): 717-723. Riley, C.V. 1889. The Morelos orange fruit worm. Insect Life 1: 45-47. ——. 1891. A new peach pest. U.S. Dep. Agric., Rep. Entomol. for 1890, pp. 255-257. Riley, C.V., and L.O. Howard. 1890. Some of the bred parasitic Hymenoptera in the National Collection. Insect Life 2: 348-353. -. 1891a. A peach pest in Bermuda. Insect Life 3: 5-8. — . 1891b. Professor Harvey’s bulletin on the apple maggot. Insect Life 3: 253—254. Rivard, I. 1967. Opius lectus and O. alloeus, larval parasites of the apple maggot, Rbagoletis pomonella, in Quebec. Can. Entomol. 99: 895-896.

541

References Cited

-. 1968. Synopsis et bibliographic annotee sur la mouche de la pomme, Rbagoletis pomonella (Walsh) (Dipteres: Tephritidae). Mem. Soc. Entomol. Que. 2: 1-158. -. 1972. Piegeage de la mouche de la pomme a l’aide de pomme artificielles engluees. Phytoprotection 53: 62—70. -. 1973. Inventaire historique sur les fluctuations de population de la mouche de la pomme, Rbagoletis pomonella (Walsh), dans le sudouest du Quebec. Ann. Entomol. Soc. Que. 18: 126-137. Robertson, H.A. 1924. The rose curculio in Manitoba, with notes on other insects affecting roses. Entomol. Soc. Ont., Annu. Rep. 54: 12—16. Robineau-Desvoidy, J.B. 1830. Essai sur les Myodaires. Memoires presentes par divers savans a l’Academie Royale des Sciences de l’Institut de France. Sciences Mathematiques et Physi¬ ques, vol. 2. 813 pp. Robinson, A.S., and G. Hooper [eds.]. 1989. Fruit flies. Their biology, natural enemies and control, vol. 3A. Elsevier, New York. 372 pp. Rock, G.C., and D.R. Yeargan. 1974. Flight activity and population estimates of four apple insect species as determined by pheromone traps. Environ. Entomol. 3: 508—510. Rohdendorf, B.B. 1961. Palaearctic fruit-flies of the genus Rbagoletis Loew and closely related genera. Entomol. Rev. 40: 89-102 (English translation). Rohwer, G.G. 1958. The Mediterranean fruit fly in Florida—past, present, and future. Fla. Entomol. 41: 23—25. Rohwer, G.G., and D.L. Williamson. 1983. Pest risk evaluation in regulatory entomology. Bull. Entomol. Soc. Am. 29(4): 41—46. Roitberg, B.D., and R.J. Prokopy. 1981. Experience required for pheromone recognition by the apple maggot fly. Nature 292(5823): 540—541. -. 1982. Influence of intertree distance on foraging behavior of Rbagoletis pomonella in the field. Ecol. Entomol. 7: 437-442. Roitberg, B.D., J.C. van Lenteren, J.J.M. van Alphen, F. Galis, and R.J. Prokopy. 1982. Foraging behavior of Rbagoletis pomonella, a parasite of hawthorn (Crataegus viridis) in nature. J. Anim. Ecol. 51: 307—325. Rondani, C. 1856. Dipterologiae Italicae prodromus. Parmae, vol. 1. 288 pp. -. 1870a. Dipterologiae Italicae prodromus. vol. 7, pars. 4 (sect. 1): Genera italica ordinis dipterorum ordinatum disposita et distincta et in familia et stirpes aggregata. 59 pp. Parma. -. 1870b. Ortalidinae Italicae collectae, distinctae et in ordinum depositae. Dipterologiae Italicae prodromi, pars VII, Fasc. 4, Sect. 1. Bol. Soc. Entomol. Ital. 2: 5—34, 105—133. -. 1871a. Dipterologiae Italicae prodromus. vol. 7, Pars 4 (sect. 2). Genera Italica ordinis dipterorum ordinatum disposita et distincta et in familias et stirpes aggregata. 49 pp, Parma. -. 1871b. Ortalidinae Italicae collectae, distinctae, et in ordinum dispositae. Diptero¬ logiae Italicae Prodromus, pars VII, Fasc. 4 (sect. 2). Bol. Soc. Entomol. Ital. 3: 3—24, 161-188. Rosewall, O.W. 1922. Insects of the yellow thistle. Entomol. News 33: 176—180. Ross, W.A. 1913. Recent work on the apple maggot in Ontario. Entomol. Soc. Ont., Annu. Rep. 43: 67—72. Rossiter, M.C., D.J. Howard, and G.L. Bush. 1983. Symbiotic bacteria of fruit flies, pp. 7784. In R. Cavalloro [ed.], Fruit flies of economic importance. Proceedings of the CEC/IOBC International Symposium, Athens, Greece, 16—19 November 1982. Balkema, Rotterdam. Roth, H., and H.H. Richardson. 1970. Ethylene dibromide, methyl bromide, and ethylene chlorobromide as fumigants for control of apple and blueberry maggots in fruit. J. Econ. Entomol. 63: 496-499. Rubio, R.E., and M.W. McFadden. 1966. Isolation and identification of bacteria in the diges¬ tive tract of the Mexican fruit fly, Anastrepba ludens (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 59: 1015—1016. Rust, E.W. 1918. Anastrepba fratercuius Wied.—a severe menace to the southern United States. J. Econ. Entomol. 11: 457—467.

542

References Cited

Ryckman, R.E. 1951. Injurious effects of Paracantha culta on Canadian thistle in Wisconsin. J. Econ. Entomol. 44: 127. Sabrosky, C.W. 1967. Corrections to A Catalog of the Diptera of America North of Mexico. Bull. Entomol. Soc. Am. 13: 115-125. -. 1971. Additional corrections to A Catalog of the Diptera of America North of Mexico. Bull. Entomol. Soc. Am. 17: 83-88. Sabrosky, C.W., and E.C. Zimmerman. 1966. Trypetesinae and Trypetesini (Lacordaire): proposed emendation of family-group names under the plenary powers (Insecta: Coleoptera) Z.N.(S.) 1733. Bull. Zool. Nomencl. 23: 46—47. Sanders, R.R. 1964. Contributions from the Los Angeles Museum-Channel Islands biological survey 38. Diptera from San Nicolas Island and Point Mugu, California. Bull. South. Calif. Acad. Sci. 63: 21-25. Sarma, R., G.B. Kitto, S. Berlocher, and G.L. Bush. 1987. Biochemical and immunological studies on an alpha-glycerophosphate dehydrogenase from the tephritid fly, Anastrepha suspensa. Arch. Inst. Biochem. Physiol.4: 271-286. Saunders, W.W. 1841. Descriptions of four new dipterous insects from central and northern India. Trans. Entomol. Soc. Lond. 3: 59-61. Say, T. 1830. Descriptions of North American dipterous insects. J. Acad. Nat Sci Phila 6183-188. Schiner, I.R. 1868. 1. Diptera, 388 pp., 4 pis. In B. von Wiillerstorff-Urbair [in charge], Reise der osterreichen Fregatte Novara. Zool., vol. 2, Abt. 1, [Sect.] B. Vienna. Schmidt, C.T. 1967. A fruit-fly attacking coffee cherries in Sao Tome. Garcia de Orta (Lisboa) 15: 329-331. ' Schrank, F. 1795. Naturhistorische und okonomische Briefe iiber das Donaumoor. 211 pp pi. Mannheim.

1

Schulz, J.T., and W.V. Lipp. 1969. The status of the sunflower insect complex in the Red River Valley of North Dakota. Proc. N. Centr. Br., Entomol. Soc. Am. 24- 99-100 Schwitzgebel, R.B., and D.A. Wilbur. 1943. Diptera associated with ironweed, Vernonia interior, in Kansas. J. Kans. Entomol. Soc. 16: 4-13. Sein, F., Jr. 1933. Anastrepha (Trypetidae, Diptera) fruit flies in Puerto Rico. I. Dep. Aerie Puerto Rico 17: 183-196. 6 Severin, H.H.P. 1913a. Precautions taken and the danger of introducing the Mediterranean fruit fly (Ceratitis capitata Wied.) into the United States. J. Econ. Entomol. 6: 68-73. -. 1913b. The life history of the Mediterranean fruit fly (Ceratitis capitata Wied.) with a list of fruits attacked in the United States. J. Econ. Entomol. 6: 399-403. —. 1917a. Life history, habits, natural enemies and methods of control of the currant fruit fly (Epochra canadensis Loew). Maine Agric. Exp. Stn. Bull. 264: 177-246. . 1917b. Dark currant fruit fly in California. Rhagoltis [sic] ribicola Doane. Calif State Comm. Hortic. Mo. Bull. 6(6): 258. . 1918a. Fruit flies of economic importance in California. Currant fruit fly (Epochra canadensis (Loew)). Calif. Dep. Agric. Mo. Bull. 7: 201-206. ——. 1918b. A native food plant of Rhagoletis fausta O.S. J. Econ. Entomol. 11: 325-327 Sharp, J L. 1976. Comparison of flight ability of wild-type and laboratory-reared Caribbean fruit flies on a flight mill. J. Ga. Entomol. Soc. 11: 255-258. . 1978. Tethered flight of apple maggot flies. Fla. Entomol. 61: 199-200. ——. 1979. A method to determine wing dimensions of insects. Fla. Entomol. 62: 69-71. Sharp, JX. and D.L. Chambers. 1983. Aggregation response of Anastrepha suspensa (Dip¬ tera: Tephritidae) to proteins and amino acids. Environ. Entomol. 12: 923-928. -1984. Consumption of carbohydrates, proteins, and amino acids by Anastrepha suspen¬ sa (Loew) (Diptera: Tephritidae) in the laboratory. Environ. Entomol. 13: 768-773. Sharp, J.L., and P.J. Landolt. 1984. Gustatory and olfactory behavior of the papaya fruit fly

Toxotrypana curvicauda Gerstaecker (Diptera: Tephritidae), in the laboratory, with notes on longevity. J. Ga. Entomol. Soc. 19: 176-182.

543

References Cited Sharp, J.L., and H.F. Little. 1982. Effect of mechanical disturbance on median dorsal flight muscle development of Anastrepha suspensa. Ann. Entomol. Soc. Am. 75: 146—149. Sharp, J.L., and J.C. Webb. 1977. Flight performance and signaling sound of irradiated or unirradiated Anastrepha suspensa. Proc. Hawaii. Entomol. Soc. 22: 525—532. Sharp, J.L., T.R. Ashley, D.R. Bennett, and B.J. Smittle. 1975. Emergence, longevity, fecundity, and sterility of Anastrepha suspensa (Diptera: Tephritidae) irradiated in nitrogen. J. Ga. Entomol. Soc. 19: 241—250. Sharp, J.L., T.C. Carlysle, and H.F. Little. 1980. Missing indirect flight muscles in Anastrepha suspensa with droopy wing syndrome. Ann. Entomol. Soc. Am. 73: 239-243. Sharp, J.L., E.F. Boiler, and D.L. Chambers. 1983. Selection for flight propensity of laboratory and wild strains of Anastrepha suspensa and Ceratitis capitata (Diptera: Tephritidae). J. Econ. Entomol. 76: 302—305. Shaw, J.G. 1947. Hosts and distribution of Anastrepha serpentina in northeastern Mexico. J. Econ. Entomol. 40: 34—40. -. 1962. Species of the spatulata group of Anastrepha. J. Kans. Entomol. Soc. 35: 408— 414. Shepherd, D.R. 1957. Eradication of Mediterranean fruit fly in Florida. F.A.O. Plant Prot. Bull. 5: 101-103. Shervis, L.J., G.M. Bousch, and C.F. Koval. 1970. Infestation of sour cherries by the apple maggot: confirmation of a previously uncertain host status. J. Econ. Entomol. 63: 294-295. Shewell, G.E. 1961. Notes on three European Diptera recently discovered in Canada. Can. Entomol. 93: 1044—1047. Shiraki, T. 1933. A systematic study of the Trypetidae in the Japanese Empire. Mem. Fac. Sci. Agric., Taihoku Imp. Univ. 8 (Entomol. No. 2). 509 pp. Shorthouse, J.D. 1980. Modification of the flower heads of Sonchus arvensis (Family Compositae) by the gall former Tephritis dilacerata (Order Diptera, Family Tephritidae). Can. J. Bot. 58: 1534-1540. Silverman, J., and R.D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp., on the ragweed, Ambrosia dumosa (Gray) Paine, in southern California (Diptera: Tephritidae). Pan-Pac. Entomol. 56: 283-288. Simanton, W.A. 1958. Studies of Mediterranean fruit fly lures in Florida. J. Econ. Entomol. 51: 679-682. Simkover, H.G. 1953. Rhagoletis cingulata on wild and cultivated cherries in eastern Wash¬ ington. J. Econ. Entomol. 46: 896-897. Simon, J.P. 1969. Comparative serology of a complex species group of food plant specialists: the Rhagoletis pomonella complex. Syst. Zool. 18: 169—184. Sivinski, J. 1984. Effect of sexual experience on male mating success in a lek-forming tephritid, Anastrepha suspensa (Loew). Fla. Entomol. 67: 126—130. Slingerland, M.V. 1899. The cherry fruit fly, a new cherry pest. N.Y. Agric. Exp. Stn. Bull. 172: 23-41. -. 1902. The scientific name of the cherry fruit fly. Can. Entomol. 34: 28. Slosson, A.T. 1895. Additional list of insects taken in the alpine region of Mt. Washington. Entomol. News 6: 4—7. Smith, D.C., and R.J. Prokopy. 1980. Mating behavior of Rhagoletis pomonella (Diptera: Tephritidae). VI. Site of early-season encounters. Can. Entomol. 112: 585-590. -. 1981. Seasonal and diurnal activity of Rhagoletis mendax flies in nature. Ann. Entomol. Soc. Am. 74: 462—466. -. 1982. Mating behavior in Rhagoletis mendax (Diptera: Tephritidae) flies in nature. Ann. Entomol. Soc. Am. 75: 388—392. Snodgrass, R.E. 1923. The anatomy and metamorphosis of the apple maggot (Rhagoletis pomonella Walsh). J. Wash. Acad. Sci. 13: 260—261. -. 1924. Anatomy and metamorphosis of the apple maggot, Rhagoletis pomonella Walsh. J. Agric. Res. 28: 1—36.

544

References Cited

Snow, F.H. 1903. A preliminary list of the Diptera of Kansas. Kans. Univ. Sci. Bull 2- 211—

221.

-. 1904. Lists of Coleoptera, Lepidoptera, Diptera, and Hemiptera collected in Arizona by the entomological expeditions of the University of Kansas in 1902 and 1903 Kans Univ Sci. Bull. 2: 323-350. Snow, W.A. 1894. Descriptions of North American Trypetidae, with notes. Kans Univ Sci Quart. 2: 159-174. -. 1895. On Toxotrypana of Gerstaecker. Kans. Univ. Sci. Quart. 4: 117-119. Snyder, A.J. 1898. Trypeta solidaginis. Can. Entomol. 30: 99-100. Solferini, V.N., and J.S. Morgante. 1987. Karyotype study of eight species of Anastrepha (Diptera: Tephritidae). Caryologia 43: 229—241. Spishakoff, L.M. 1966. Laboratory rearing of Anastrepha serpentina. I. Econ. Entomol 59-

1010-1011.

’

’

Spider, G.H., J.W. Armstrong, and H.M. Coney. 1984. Mediterranean fruit fly (Diptera: Tephritidae). Host status of commercial lemon. J. Econ. Entomol. 77: 1441-1444. Stanton, G.E. 1971. Reactions of caged adult apple maggots, Rhagoletis pomonella (Diptera: Tephritidae) to the properties of reflected light and color. J. Ga. Entomol. Soc. 6: 61-68. -.

1972. Continuous laboratory rearing of multivoltine apple maggots, Rhagoletis

pomonella. J. Ga. Entomol. Soc. 7: 254-259. Stebbins, F.A. 1910. Insect galls of Springfield, Massachusetts, and vicinity. Springfield Mus. Nat. Hist. Bull. 2. 138 pp. Steck, G.J. 1981. North American Terelliinae (Diptera: Tephritidae): Biochemical systematics and evolution of larval feeding niches and adult life histories. Ph.D. dissertation, University of Texas, Austin. . 1984. Chaetostomella undosa (Diptera: Tephritidae): Biology, ecology, and larval des¬ cription. Ann. Entomol. Soc. Am. 77: 669-678. -. 1991. Biochemical systematics of Anastrepha fraterculus and related species (DipteraTephritidae). Ann. Entomol. Soc. Am. 84(1): 10-28. Steck, G.J., and R.A. Wharton. 1985. Descriptions of immature stages of Eutreta (DipteraTephritidae). J. Kans. Entomol. Soc. 59: 296-302. -. 1989. Description of immature stages of Anastrepha interrupta, A. limae, and A. grandis (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 81: 994-1003. Steck, G.J, L.E. Carroll, H.E. Celedonio-Hurtado, and J. Guillen-Aguilar. 1990. Methods for identification of Anastrepha larvae (Diptera: Tephritidae), and key to 13 species. Proc Entomd. Soc. Wash. 92: 333-346. Stegmaier, C.E., Jr. 1967a. Pluchea odorata, a new host record for Acinia picturata (DipteraTephritidae). Fla. Entomol. 50: 53-55. -. 1967b. Notes on a seed-feeding Tephritidae, Paracantha forficula (Diptera), in Florida Fla. Entomol. 50: 157-160. —.1968a. Host plant records of Dyseuaresta mexicana (Diptera: Tephritidae) with notes on its life history in Florida. Fla. Entomol. 51: 19-21. -. 1968b. Engeron, a host plant genus of tephntids (Diptera). Fla. Entomol. 51: 45-50. —. 1968c. Notes on the biology of Trupanea actinobola. Fla. Entomol. 51: 95-99. Steiner, L.F. 1957. Low-cost plastic fruit fly trap. J. Econ. Entomol. 50: 508-509. . 1965. A rapid method for identifying dye-marked fruit flies. J. Econ. Entomol. 58: 3743 73 • Ste;ner• L.F., D.H. Miyashita, and L.D. Christenson. 1957. Angelica oils as Mediterranean fruit fly lures. J. Econ. Entomol. 50: 505. Steiner, L.F., G.G. Rohwer, E.L. Ayers, and L.D. Christenson. 1961. The role of attractants in the^recent Mediterranean fruit fly eradication program in Florida. J. Econ. Entomol. 54: Stephenson, B.C., and B.B. McClung. 1966. Mediterranean fruit fly eradication in the lower Rio Grande Valley. Bull. Entomol. Soc. Am. 12: 374.

545

References Cited Stevenson, A.B. 1963. Laboratory emergence of adults from overwintering pupae of the apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae). Can. Entomol. 95: 1154— 1159. Steyskal, G.C. 1945. Behavior of Strauzia longipennis var. vittigera Loew. Bull. Brooklyn Entomol. Soc. 39 (1944): 156. -. 1957. Notes on color and pattern of eye in Diptera. II. Bull. Brooklyn Entomol. Soc. 52: 89-94. -. 1972a. The dipterous fauna of the heads of Bidens pilosa re-examined. Fla. Entomol. 55: 87-88. -. 1972b. A new species of Myoleja with a key to North American species (Diptera: Tephritidae). Fla. Entomol. 55: 207—212. -. 1972c. A preliminary key to the species of Neotepbritis Hendel (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 74: 414—416. -. 1972d. A correction to Sturtevant’s paper on the seminal receptacles, etc., of the Diptera, 1926. Proc. Entomol. Soc. Wash. 74: 130. -. 1973. Distinguishing characters of larvae of the walnut husk maggots of the genus Rhagoletis (Diptera: Tephritidae). Coop. Econ. Insect Rep. 23: 522. -. 1974. Recent advances in the primary classification of the Diptera. Ann. Entomol. Soc. Am. 67: 513-517. -. 1975a. Anastrepha obliqua (Macquart) the prior name for Anastrepha mombinpraeoptans Sein (Fruit flies, Tephritidae, Diptera). Coop. Econ. Insect Rep. 25: 357-358. -. 1975b. Recognition characters for larvae of the genus Zonosemata (Diptera: Tephritidae). Coop. Econ. Insect Rep. 25: 231—232. -. 1977a. Two new Neotropical fruit flies of the genus Anastrepha, with notes on generic synonymy (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 77: 75—81. -. 1977b. Pictorial key to species of the genus Anastrepha (Diptera: Tephritidae). Special Publ., Entomol. Soc. Wash. 35 pp. -. 1977c. History and use of the McPhail trap. Fla. Entomol. 60: 11 — 16. -. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Special Publ., Entomol. Soc. Wash. 61 pp. -. 1981. A new species of Rhagoletotrypeta (Diptera: Tephritidae) from Texas, with a key to the known species. Proc. Entomol. Soc. Wash. 83: 707—712. -. 1982. A second species of Ceratitis (Diptera: Tephritidae) adventive in the New World. Proc. Entomol. Soc. Wash. 84: 165—166. -. 1984. A synoptic revision of the genus Aciurina Curran, 1932 (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 86: 582—598. -. 1986. Taxonomy of the adults of the genus Strauzia Robineau-Desvoidy (Diptera: Tephritidae). Insecta Mundi 1: 101 — 117. Steyskal, G.C., and R.H. Foote. 1963. Notes on the genus Icterica Loew in North America. Proc. Entomol. Soc. Wash. 64(1962): 166. -. 1977. Revisionary notes on North American Tephritidae (Diptera), with keys and des¬ criptions of new species. Proc. Entomol. Soc. Wash. 77: 146—155. Steyskal, G.C., W.L. Murphy, and E.M. Hoover [eds.]. 1986. Insects and mites: techniques for collection and preservation. U.S. Dep. Agric. Misc. Publ. 1443. 103 pp. Still, G.W. 1960. An improved trap for deciduous tree fruit flies. J. Econ. Entomol. 53: 967. Stinner, B.R., and W.G. Abrahamson. 1979. Energetics of the Solidago canadensis stem-gall insect-parasitoid guild interaction. Ecology 69: 918—926. Stirret, G.M. 1936. The walnut husk fly, Rhagoletis suavis Loew, in Ontario. Annu. Rep. Entomol. Soc. Ont. 66: 69. Stoetzel, M.B. [ed.]. 1989. Common names of insects & related organisms, 1989. Ento¬ mological Society of America, Lanham, Md. Stoffolano, J.G., and L.R.S. Yin. 1987. Structure and function of the ovipositor and associated

References Cited

sensilla of the apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae). Int. J. Insect Morphol. Embryol. 16: 41—69. Stoltzfus, W.B. 1974. Biology and larval description of Procecidocharoides penelope (Diptera: Tephritidae). J. Wash. Acad. Sci. 64: 12-14. -. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae). Iowa State J. Res 51(4): 369-438. -. 1988. The taxonomy and biology of Strauzia (Diptera: Tephritidae). J. Iowa Acad. Sci 95: 117-126. -. 1989. A non-gall forming Eurosta solidaginis (Diptera: Tephritidae). J. Iowa Acad. Sci 96: 50-51. Stoltzfus, W.B., and B.A. Foote. 1965. The use of froth masses in courtship of Eutreta (Dip¬ tera: Tephritidae). Proc. Entomol. Soc. Wash. 67: 263-264. Stone, A. 1939a. A revision of the genus Pseudodacus. Rev. Entomol. 10: 282-289. -. 1939b. A new genus of Trypetidae near Anastrepha (Diptera). J. Wash. Acad. Sci. 29: 340-350. -. 1941. The generic names of Meigen 1800 and their proper application (Diptera). Ann. Entomol. Soc. Am. 34: 404-418. -. 1942a. The fruitflies of the genus Anastrepha. U.S. Dep. Agric. Misc. Publ. 439: 1-112. -. 1942b. New species of Anastrepha and notes on others. J. Wash. Acad. Sci. 32: 298304. -. 1951. The Rhagoletis of roses. Proc. Entomol. Soc. Wash. 53: 45-48. Stone, A., C.W. Sabrosky, W.W. Wirth, R.H. Foote, and J.R. Coulson [eds.]. 1965. A catalog of the Diptera of America north of Mexico. U.S. Dep. Agric. Government Printing Office, Washington, D.C. Story, J.M. 1977. Effect of Urophora affinis on spotted knapweed seed production in Mon¬ tana, p. 208. In Proc. Knapweed Symposium, Kamloops, B.C. -. 1985. First report of the dispersal into Montana of Urophora quadrifasciata (Diptera: Tephritidae), a fly released in Canada for the biological control of diffuse and spotted knapweed. Can. Entomol. 118: 1061-1062. Story, J.M., and N.L. Anderson. 1978. Release and establishment of Urophora affinis (Dip¬ tera: Tephritidae) on spotted knapweed in western Montana. Environ. Entomol 7* 445448. Story, J.M., and R.M. Nowiersky. 1984. Increase and dispersal of Urophora affinis (Diptera: Tephritidae) on spotted knapweed in western Montana. Environ. Entomol. 13: 1151-1156. Story, J.M., R.M. Nowiersky, and K.M. Boggs. 1987. Distribution of Urophora affinis and U. quadrifasciata, two flies introduced for biological control of spotted knapweed (Centaurea maculosa) in Montana. Weed Sci. 35: 145-148. Strickland, E.H. 1938. An annotated list of the Diptera (flies) of Alberta. Can. I. Res (Sec D) 16: 175-219. -. 1946. An annotated list of the Diptera (flies) of Alberta, additions and corrections. Can J. Res. (Sec. D) 24: 157-173. -. 1953. The ptilinal armature of flies. Can. J. Zool. 31: 263-299. Strong, L.A. 1930. The Mediterranean fruit fly eradication campaign. J. Econ. Entomol 23509-512. Sturtevant, A.H. 1918. Acalypterae (Diptera) collected in Mobile Co., Alabama. J. N.Y. Entomol. Soc. 26: 34-40. . 1925—1926. The seminal receptacles and accessory glands of the Diptera, with special reference to the Acalypterae. J. N.Y. Entomol. Soc. 33: 195-215; 34: 1-21. Sutherland, D.W.S. 1978. Common names of insects and related organisms (1978 revision). Entomol. Soc. Am. Spec. Publ. 78-1. 132 pp. Swanson, R.W., and R.M. Baranowski. 1972. Host range and infestation by the Caribbean fruit fly, Anastrepha suspensa (Diptera: Tephritidae), in south Florida. Proc. Fla. State Hortic. Soc. 85: 271-274.

547

References Cited Sweet, H.E. 1930. An ecological study of the animal life associated with Artemisia californica Less at Claremont, California. J. Entomol. Zool., Pomona College 22: 75-151. Szentesi, A., P.D. Greany, and D.L. Chambers. 1979. Oviposition behavior of laboratoryreared and wild Caribbean fruit flies (Anastrepha suspensa) (Diptera: Tephritidae): I. Selected chemical influences. Entomol. Exp. Appl. 26: 227—238. Taschenbert, E.F., F. Lopez-D., and L.F. Steiner. 1974. Responses of maturing larvae of Ana¬ strepha suspensa to light and to immersion in water. J. Econ. Entomol. 67: 731—734. Tauber, M.J., and C.A. Tauber. 1967. Reproductive behavior and biology of the gall-former Aciurina ferruginea (Doane) (Diptera: Tephritidae). Can. J. Zool. 45: 907-913. -. 1968. Biology of the gall former Procecidochares stonei on a composite. Ann. Entomol. Soc. Am. 61: 553—554. Tauber, M.J., and C.A. Toschi. 1965a. Bionomics of Euleia fratria (Loew) (Diptera: Tephritidae). I. Life history and mating behavior. Can. J. Zool. 43: 369-379. -. 1965b. Life history and mating behavior of Tephritis stigmatica (Coquillett) (Diptera: Tephritidae). Pan-Pac. Entomol. 41: 73—79. Teskey, H.J. 1981. Key to families—larvae, pp. 125-147. In J.F. McAlpine et al. [coords.], Manual of Nearctic Diptera, vol. 1. Res. Br., Agric. Can. Monogr. 27, Ottawa. Thomas, F.L. 1914. Three new species of Trypetidae from Colorado. Can. Entomol. 46: 425— 428. Thompson, F.C. 1989. Biosystematic information systems. Systematic Entomology Labora¬ tory, USDA, Washington, D.C. 7 pp. Thompson, F.C., and L.E. Carroll. 1991. Fruit fly expert system and biosystematic data base. Systematic Entomology Laboratory, USDA, Washington, D.C. 8 pp. Thompson, M.T. 1907. Three galls made by cyclorrhaphous flies. Psyche 14: 71-74. Thomson, C.G. 1869. Diptera, species novas descripsit. In Konglige Svenska Fregatten Eugenies Resa Omkring Jorden, Pt. 2, Zool. 1. Insekter, pp. 443-614. Tilden, J.W. 1951. The insect associates of Baccharis pilularis de Candolle. Microentomology 16: 149-188. Tomlinson, W.E. 1951. Influence of temperature on emergence of the blueberry maggot. J. Econ. Entomol. 44: 266. Tonks, N.V. 1953. Annotated list of insects and mites collected on brambles in the lower Fraser Valley, British Columbia, 1951. Proc. Entomol. Soc. B.C. 49: 27-29. Townsend, C.H.T. 1893a. A general summary of the known larval food habits of acalyptrate Muscidae. Can. Entomol. 25: 10—16. -. 1893b. A trypetid bred from galls on Bigelovia. Can. Entomol. 25: 48—52. Treherne, R.C. 1917. Popular and practical entomology. The apple maggot in British Colum¬ bia. Can. Entomol. 49: 329—330. Trottier, R., I. Rivard, and W.T.A. Neilson. 1975. Bait traps for monitoring apple maggot activity and their use for timing control sprays. J. Econ. Entomol. 68: 211—213. Tsiropolous, G.T. 1976. Bacteria associated with the walnut husk fly, Rhagoletis completa. Environ. Entomol. 5: 83—86. Tucker, E.S. 1907. Some results of desultory collecting of insects in Kansas and Colorado. Kans. Univ. Sci. Bull. 4: 51—108. -. 1908. Additional results of collecting insects in Kansas and Colorado. Trans. Kans. Acad. Sci. 22: 276—304. Uhler, L.D. 1951. Biology and ecology of the goldenrod fly, Eurosta solidaginis (Fitch). Cornell Univ. Agric. Exp. Stn. Mem. 300. 51 pp. -. 1961. Mortality of the goldenrod gall fly, Eurosta solidaginis, in the vicinity of Ithaca, New York. Ecology 42: 215-216. Van Duzee, M.C. 1911. A list of Diptera taken at Kearney, Ontario, in July, 1909. Can. Entomol. 43: 237—244. Varley, G.C. 1937. The life history of some trypetid flies, with descriptions of the early stages. Proc. Roy. Entomol. Soc. Lond. (Ser. A) 12: 109-122.

References Cited

-. 1947. The natural control of population balance in the knapwweed gall fly (Urophora jaceana). J. Anim. Ecol. 16: 139-187. Vasey, C.E., and E. Ritter. 1987. Antennal sensilla and setal patterns of the goldenrod gall fly, Eurosta solidaginis (Fitch) (Diptera: Tephritidae). J. N.Y. Entomol. Soc. 95: 452-455. Viereck, H.L. 1913. Descriptions of ten new genera and twenty three new species of ich¬ neumon flies. Proc. U.S. Natl. Mus. 44(1968): 555-568. -. 1917. Hymenoptera of Connecticut. Conn. State Geol. Nat. Hist. Surv. Bull. 22. 824 pp. von Windeguth, D.L., W.H. Pierce, and L.F. Steiner. 1972. Infestations of Anastrepha suspensa in fruit on Key West, Florida, and adjacent islands. Fla. Entomol. 56: 127-131. von Windeguth, D.L., A. Arner, J.B. Owens, and A.K. Burditt, Jr. 1976. Comparison of natural field infestation versus laboratory infestability of Marsh white grapefruit by the Caribbean fruit fly, Anastrepha suspensa. Proc. Fla. State Hortic. Soc. 89: 248-249. Walker, F. 1835. Descriptions of the British tephritites. Entomol. Mag. (Lond.) 3: 57-85. -. 1849. List of the specimens of dipterous insects in the collection of the British Museum. Part IV (Fam. Muscidae), pp. 689-1172. London. -. 1856. Catalog of the dipterous insects collected in Singapore and Malacca by Mr. A.R. Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond., Zool. 1: 4-39. -. 1857. Characters of undescribed Diptera in the collection of W.W. Saunders. Trans. Entomol. Soc. Lond. (n. ser.) 4: 119-158. Walker, F.H., Jr. 1936. Observations on sunflower insects in Kansas. J. Kans. Entomol Soc 916-25. Walker, M.H. 1929. Government wages winning fight on fruit fly. Fla. Grower 37(7): 10. Walsh, B.D. 1866. Answers to correspondents. Practical Entomol. 1: 114. -. 1867. The apple-worm and the apple-maggot. Carpocapsa pomonella (Linnaeus); Trypeta pomonella (new species). Am. J. Hortic. Florist’s Companion 2: 338-343. -. 1868. The apple maggot fly (Trypeta pomonella, Walsh). Trans. Ill. State Hortic. Soc. 1867 (appendix), pp. 29-33, 36-37. Walsh, B.D., and C.V. Riley. 1869. Answers to correspondents. Am. Entomol. 1: 59. Walsh, J. 1981. Medfly continues to bug California. Science 214(4526): 1221-1222, 1224. Walton, R. 1988. Characteristics of galls of Eurosta solidaginis (Diptera: Tephritidae) and their relation to mortality of gall inhabitants. Environ. Entomol. 17: 654-659. Wangberg, J.K. 1977. A new Tetrastichus parasitizing tephritid gallformers on Chrysotbamnus in Idaho. Pan-Pac. Entomol. 53: 237-240. . 1978a. Biology of gall formers of the genus Valentibulla on rabbitbrush in Idaho. L Kans. Entomol. Soc. 51: 472-483. 1978b. Biology of the tephritid gall formers (Diptera: Tephritidae) on rabbitbrush, Chrysothamnus spp., in Idaho (abstract). Folia Entomol. Mex., Nos. 39-40, June. . 1980. Comparative biology of gall formers in the genus Procecidochares (Diptera: Tephritidae) on rabbitbrush in Idaho. J. Kans. Entomol. Soc. 53: 401-420. -. 1981. Gall-forming habits of Aciurina species (Diptera: Tephritidae) on rabbitbrush (Compositae: Chrysothamnus spp.) in Idaho. J. Kans. Entomol. Soc. 54: 711-732. Wasbauer, M.S. 1963. Taxonomic discrimination of Rhagoletis pomonella from the eastern United States and Rhagoletis zephyria from California. Calif. Dep. Agric. T-9 (mimeo)

6 pp. . 1964. Recognition of fruit flies likely to be encountered in Mexican fruit fly traps in southern California. Calif. Dep. Agric. T-10 (mimeo). 8 pp. -. 1970. The type material in the collection of the California Department of Agriculture. Pan-Pac. Entomol. 46: 47-50. ' annotated host catalog of the fruit flies of America north of Mexico (Diptera: Tephnndae) Occas. Pap. 19, Bur. Entomol., Calif. Dep. Agric., Sacramento. 172 pp. Washburn, F.L. 1905. The Diptera of Minnesota. Minn. Agric. Exp. Stn. Bull. 93: 19168. . 1906. Additional Minnesota Diptera. Minn. Agric. Exp. Stn. Bull. 100: 79-82.

549

References Cited

Waterston, J.M. 1941. A list of food-plants of some Bermuda insects. Bermuda Dep. Agric., Hamilton. 63 pp. -. 1944. Fruit growing in Bermuda. Report of the plant pathologist on a visit to the United States of America and the Bahamas to obtain information on the growing of citrus and other subtropical fruits, Aug. 15-Dec. 24, 1943. Hamilton, Bermuda. 125 pp. Watson, A.K. 1979. Biological weed control program in Quebec. Phytoprotection 60: 136144. Webb, J.C., J.L. Sharp, D.L. Chambers, and J.D. Benner. 1976a. Acoustical properties of the flight activities of the Caribbean fruit fly. J. Exp. Biol. 64: 761-772. Webb, J.C., J.L. Sharp, D.L. Chambers, J.J. McDow, and J.C. Benner. 1976b. Analysis and identification of sounds produced by the male Caribbean fruit fly, Anastrepha suspensa. Ann. Entomol. Soc. Am. 69: 415—420. Webb, J.C., H.R. Agee, N.C. Leppla, and C.O. Calkins. 1981. Monitoring insect quality. Trans. Am. Soc. Agric. Eng. 24: 476—479. Webb, J.C., T. Burk, and J. Sivinski. 1983. Attraction of female Caribbean fruit flies (Anastrepba suspensa (Loew)) to males and male-produced stimuli in field cages. Ann. Entomol. Soc. Am. 76: 996-998. Webster, F.M. 1894. Biological notes on reared parasitic Hymenoptera of Ohio and Indiana, with descriptions of 13 new species by W.H. Ashmead. J. Cincinnati Soc. Nat. Hist. 17: 34— 55. Webster, R.P., and J.G. Stoffolano, Jr. 1978. The influence of diet on the maturation of the reproductive system of the apple maggot, Rhagoletis pomonella. Ann. Entomol. Soc. Am. 71: 844-849. Webster, R.P., J.G. Stoffolano, Jr., and R.J. Prokopy. 1979. Long-term intake of protein and sucrose in relation to reproductive behavior of wild and laboratory-cultured Rhagoletis

pomonella. Ann. Entomol. Soc. Am. 72: 41—46. Weed, C.M. 1896. The apple maggot or railroad worm. N.H. Agric. Exp. Stn. Bull. 35: 3135. Weems, H.V., Jr. 1963. Mexican fruit fly (Anastrepha ludens (Loew) (Diptera: Tephritidae). Fla. Dep. Agric., Bur. Plant Ind., Entomol. Circ. 16. 2 pp. -. 1964a. Oriental fruit fly (Dacus dorsalis Hendel). Fla. Dep. Agric., Bur. Plant Ind., Entomol. Circ. 21. 2 pp. -. 1964b. Melon fly (Dacus cucurbitae Coquillett). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 29. 2pp. -. 1965a. Queensland fruit fly (Dacus tryoni Froggatt)) (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 34. 2 pp. -. 1965b. Anastrepha suspensa (Loew). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 38. 4 pp. -. 1966. The Caribbean fruit fly in Florida. Fla. State Hortic. Soc. 79: 401—405. -. 1967a. Anastrepha interrupta Stone (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 61. 2 pp. -. 1967b. Anastrepha nigrifascia Stone (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 66. 1 p. -. 1968a. Anastrepha ocresia (Walker) (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 71. 2 pp. -. 1968b. Anastrepha edentata Stone (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 76. 2 pp. -. 1969a. Papaya fruit fly (Toxotrypana curvicauda Gerst.), 1860 (Diptera: Tephritidae). Fla. Dep. Agric., Div. Plant Ind., Entomol. Circ. 86. 2 pp. -. 1969b. Anastrepha serpentina (Wiedemann) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 91. 2 pp. -. 1970. West Indian fruit fly, Anastrepha mombinpraeoptans Sein (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 101. 2 pp.

References Cited

-. 1972a. Cherry fruit fly, cherry maggot, Rhagoletis cingulata (Loew) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 116. 2 pp. -. 1972b. Apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv, Div. Plant Ind., Entomol. Circ. 126. 2 pp. -. 1980. Anastrepha fraterculus (Wiedemann) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 217. 4 pp. -. 1981a. Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 230. 12 pp. -. 1981b. Major fruit flies of the world. Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Leaflet 3 (rev.). 8 pp. --. 1982. Anastrepha striata Schiner (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 245. 2 pp. -. 1987. Guava fruit fly (Dacus (Strumeta) correctus (Bezzi)) (Diptera: Tephritidae). Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Entomol. Circ. 291. 4 pp. Weigel, R., and E. Dilks. 1950. The ball gall of goldenrod as laboratory material. Turtox News 28: 134-137. Wellhouse, W.H. 1920. Wild hawthorn as hosts of apple, pear, and quince pests. T. Econ. Entomol. 13: 388-391. . 1922. The insect fauna of the genus Crataegus. N.Y. Agric. Exp. Stn. Mem. 56- 10451136. Westcott, R.L. 1982. Differentiating adults of apple maggot, Rhagoletis pomonella (Walsh), from snowberry maggot, Rhagoletis zephyria Snow (Diptera: Tephritidae), in Oregon. PanPac. Entomol. 58: 25-30. Westdahl, P.FL, and C.F. Barrett. 1960. Life-history and habits of the sunflower maggot, Strauzia longipennis (Wied.) in Manitoba. Can. Entomol. 92: 481-488. -. 1962. Injury by the sunflower maggot, Strauzia longipennis (Wied.). Can. I Plant Sci 42: 11-14. Westwood, J.O. 1840. Order XIII. Diptera Aristotle (Antliata Fabricius, Halteriptera Clairv.), pp. 125—158. In J.O. Westwood, An introduction to the modern classification of insects. Synopsis of the genera of British Insects. London. 158 pp. Wharton, R.A., and P.M. Marsh. 1978. New World Opiinae (Hymenoptera: Braconidae) parasitic on Tephritidae (Diptera). J. Wash. Acad. Sci. 68: 147-167. White, I.M. 1986. A new species of Paroxyna Hendel and notes on the nomenclature of other British Tephritidae (Diptera). Entomol. Mo. Mag. 122: 145-163. -. 1987. The Linnaean species of the family Tephritidae (Insecta: Diptera). Zool 1 Linn Soc. 90: 99-107. . 1988. Tephritid flies - Diptera: Tephritidae, pp. 1-134. In P.C. Barnard and R.R. Askew feds.], Handbooks for the identification of British insects, vol. 10, pt. 5a. Royal Entomologi¬ cal Society of London. White, I.M., and S.L. Clement. 1987. Systematic notes on Urophora (Diptera: Tephritidae) species associated with Centaurea solstitialis (Asteraceae, Cardueae) and other palaearctic weeds adventive in North America. Proc. Entomol. Soc. Wash. 89: 571-580. White, I.M., and V.A. Korneyev. 1989. A revision of the western Palaearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae). Syst. Entomol. 14: 327-374. White, I.M., and K. Marquardt. 1989. A revision of the genus Chaetorellia Hendel (Diptera: Tephritidae) including a new species associated with spotted knapweed, Centaurea mac¬ ulosa Lam. (Asteraceae). Bull. Entomol. Res. 79: 453-487. White, I.M., K. Groppe, and R. Sobhian. 1990. Tephritids of knapweeds, starthistles and safflower: results of a host choice experiment and the taxonomy of Terellia luteola (Wiedemann) (Diptera: Tephritidae). Bull. Entomol. Res. 80: 107-111. Whitman, D.W., L. Orsak, and E. Greene. 1988. Spider mimicry in fruit flies (DipteraTephritidae): further experiments on the deterrence of jumping spiders (Araneae: Salticidae) by Zonosemata vittigera (Coquillett). Ann. Entomol. Soc. Am. 81: 532-536.

551

References Cited Whitney, L.A. 1916. The yellow currant and gooseberry fruit fly, Epochra canadensis (Loew). Calif. Dep. Agric. Mo. Bull. 5: 152—156. Wiedemann, C.R.W. 1824. Munus rectoris in Academia Christiana Albertina aditurus Ana¬ lecta entomologica ex Museo Regio Havniensi. Kiel. 60 pp. -. 1830. Ausser europaische zweifliigelige Insekten, vol. 2. pp. xii + 684, pis. 7—10. Wilde, W.H.A., and H.W. Goble. 1966. A note on Rhagoletis pomonella (Walsh) trapping (Diptera: Tephritidae). Can. Entomol. 96: 145—146. Williston, S.W. 1886. Dipterological notes and descriptions. Trans. Am. Entomol. Soc. 13: 287-307. -. 1896. Manual of the families and genera of North American Diptera, 2nd ed. Hathaway, New Haven, Conn, iiv + 167 pp. -. 1908. Manual of North American Diptera, 3rd ed. Hathaway, New Haven, Conn. 405 pp. Wilson, H.F., and A.L. Lovett. 1913. Miscellaneous insect pests of orchard and garden. Oreg. Agric. Exp. Stn. Bienn. Crop Pest Hortic. Rep., 1911-1912: 147—165. Winn, A.F., and G. Beaulieu. 1915. A preliminary list of the insects of the Province of Quebec. Part II - Diptera. Que. Soc. Prot. Plants, Seventh Rep. (Suppl.): 153—154. Winterringer, G.S. 1961. Some plant galls of Illinois. Story of Illinois Series 12: 1—57. Ill. State Mus., Urbana. Witherell, P.C. 1982. Efficacy of two types of survey traps for Caribbean fruit fly, Anastrepha suspensa (Loew). Fla. Entomol. 65: 580—581. Wolfenbarger, D.O. 1957. Notes and comments on the second Mediterranean fruit fly infesta¬ tion. Fla. Entomol. 40: 29—31. Wolfenbarger, D.O., and S.D. Walker. 1974. Two major pest problems of papaya. Proc. Fla. State Hortic. Soc. 87: 384—385. Wolfenbarger, D.O., H.H. Samol, and D.H. Habeck. 1976. Dispersal distances of the Carib¬ bean fruit fly, corn planthopper, and Cuban May beetle. Res. Pop. Ecol. 18: 118—122. Wood, M. 1989. Friendly flies may stop starthistle. Agric. Res. 37(3): 9. Woods, W.C. 1914. A note on Rhagoletis pomonella on blueberries. J. Econ. Entomol. 7: 398-399. -. 1915. Biosteres rhagoletis Richmond, sp. n., a parasite of Rhagoletis pomonella Walsh. Can. Entomol. 47: 293—295. -. 1916. Blueberry insects in Maine. Maine Agric. Exp. Stn. Bull. 244 (1915): 249-288. Woodworth, C.W. 1913. Guide to California Insects. The Law Press, Berkeley, Calif. 360 pp. Wright, R.H., and R.E. Burgess. 1975. Molecular coding of olfactory specificity. Can. J. Zool. 53: 1247-1253. Wulp, F.M. van der. 1867. Eenige Noord-Americaanscher Diptera. Tijdschr. Entomol. 10 (= ser.2,2): 125-164. -. 1883. Amerikaanischer Diptera. Tijdschr. Entomol. 26: 1—59. -. 1899-1900. Group Trypetinae. In F.D. Godman and O. Salvin, Biologia CentraliAmericana, Insecta, Diptera. London, vol. 2, pp. 401—428 (pp. 401—416 issued 1899, pp. 417—428 issued 1900). Zack, R.S. 1984. Catalog of types in the James Entomological Collection. Melanderia 42: 141. Zia, Y. 1937. Studies on the Trypetidae or fruit flies of China. Sinensia 8: 103-226. Zia, Y., and S.H. Chen. 1938. Trypetidae of North China. Sinensia 9: 1-180. Zucchi, R.A. 1979a. Novas especies de Anastrepha Schiner, 1868 (Diptera: Tephritidae). Rev. Bras. Entomol. 23(1): 35—41. -. 1979b. Sobre os tipos de Anastrepha parallela (Wiedemann, 1830), de A. striata Schiner, 1868, e de A. zernyi Lima, 1934 (Diptera: Tephritidae). Rev. Bras. Entomol. 23(4): 263266. Zwolfer, H. 1967. Observations on Urophora cardui L. (Trypetidae). Commonw. Inst. Biol. Contr., Weed Projects for Canada Prog. Rep. 19: 1-11.

552

References Cited

-. 1968. Untersuchungen zur biologischen Bekampfung von Centaurea solstitialis L.— Strukturmarkmalen der Wirtspflanze als Ausloser des Eiablageverhaltens bei Urophora siruna-seva (Hg.) (Dipt., Trypetidae). Z. Angew. Entomol. 61: 119-130. -. 1969. JJrophora siruna-seva (Hg.) (Dipt., Trypetidae), a potential insect for the biolog¬ ical control of Centaurea solstitialis in California. Commonw. Inst. Biol. Contr. Tech Bull 11: 105-155. -. 1970. Investigations on the host specificity of Urophora affinis Frfld. Commonw. Inst. Biol. Contr., Weed Projects for Canada Progr. Rep. 25: 1-28 (unpublished). -. 1972. Investigations on Urophora stylata Fabr., a possible agent for the biological control of Cirsium vulgare in Canada. Commonw. Inst. Biol. Contr., Weed Projects for Canada Progr. Rep. 29: 1—20. -. 1983. Life systems and strategies of resource exploitation in tephritids, pp. 16-30. In R. Cavallaro [ed.], Fruit flies of economic importance. Balkema, Rotterdam. 642 pp. -. 1988. Evolutionary and ecological relationships of the insect fauna of thistles. Annu. Rev. Entomol. 33: 103-122. Zwolfer, H., and P. Harris. 1971. Host specificity determination of insects for biological control of weeds. Annu. Rev. Entomol. 16: 159-178. Zwolfer, H., W. Englert, and W. Patullo. 1970. Investigations on the biology, population ecology, and the distribution of Urophora cardui L. Commonw. Inst. Biol. Contr., Weed Projects for Canada Progr. Rep. 27: 1-17.

INDEX OF INSECT NAMES

Names in italics are junior synonyms, errors, or emendations. Numbers in italics indicate pages where genera and species occur in keys; numbers in boldface indicate pages where the main mention occurs.

abstersus (Loew), Euarestoides 32, 154, 155, 156, 156, 160, 161 Acalyptratae 17, 18 Acanthiophilus Becker 32 Acanthonevrini 24 acbillae [error], Neaspilota 229 achilleae Johnson, Neaspilota 224, 225, 226, 228, 241 Acidia Robineau-Desvoidy 23, 449 Acidigona [error] 65 Acidogona Loew 21, 31, 57, 65, 323, 490, 491 acidusa (Walker), Anastrepha 101 Acinia Robineau-Desvoidy 2, 21, 25, 33, 62, 67, 243 Aciura Robineau-Desvoidy 69 Aciurina Curran 21, 28, 29, 59, 69, 75, 76, 77, 78, 79, 476 Aciurini 24, 25, 33 Aciuropsis Hardy 23 acrolophi White and Marquardt, Chaetorellia 2 Acrotaenia Loew 9, 21, 26, 27, 46, 64, 81 Acrotaeniacantha Hering 26 Acrotaeniini 21, 25, 26, 27, 30, 33, 46 Acrotoxa Loew 83 actinibola [error], Trupanea 427 actinobola (Loew), Trupanea 418, 422, 423, 424, 426, 426, 427 acutangulus (Thomson), Euarestoides 32, 63, 155, 156, 157, 158, 159, 160, 161

acuticornis (Steyskal), Rhagoletis 223, 270, 324, 325, 326, 327, 328, 333, 334 adelphica Hendel, Euaresta 137 Adramini 22 adspersa Coquillett, Euaresta 150 aenigma Freidberg and Mathis, Neaspilota 224, 228, 229, 230, 240, 241, 242 aequalis (Loew), Euaresta 48, 142, 143, 144, 145, 146, 152, 154 affinis (Frauenfeld), Urophora 46, 460, 461, 462, 462, 463, 469, 473, 474 affinis Quisenberry, Stenopa 368, 370, 370, 371, 373 affinis (Snow), Neotephritis 246 ageratae Benjamin, Trupanea 418, 419, 425, 428, 429, 440, 445, 447 Aischrocrania Hendel 23 alani Steyskal, Procecidochares 304, 318 alba (Loew), Neaspilota 222, 224, 225, 226, 230, 231, 237 albiceps (Loew), Paroxyna 61, 277, 278, 280, 281, 286, 293 albidipennis (Loew), Neaspilota 226, 231, 232 albipennis [error], Neaspilota 231 albiscutellata Say, Rhagoletis 355 albiseta Freidberg and Mathis, Neaspilota 225, 227, 232 Alcidia [error] 163 aldricbii (Doane), Tephritis 394

553

554

Index of Insect Names

Aleomyia Phillips 368, 369, 471 alpha Phillips, Rhynencina 368, 369 alternata (Fallen), Rhagoletis 322, 336 alternata group, Rhagoletis 325, 328, 336, 352 aluea (Walker), Eurosta 168 amabilis, Hexachaeta 207 American currant fruit fly 360

americana Hering, Paroxyna 289, 290 Anastrepha Schiner 2, 5, 7, 9, 11, 15, 20, 22, 34, 35, 39, 46, 50, 84, 206, 324, 415 angusta Banks, Eutreta 183, 184, 186, 188, 194, 198

australis Aldrich, Procecidochares 305, 306, 307, 309, 310, 311, 313 australis Hering, Chaetorellia 2 baccharis (Coquillett), Tephritis 180, 388, 389, 390, 397, 401, 407 Bactrocera Macquart 2, 5, 6, 20, 22, 34, 42, 46, 47, 112, 132, 133, 415 Baryglossa Bezzi 24 Baryplegma Wulp 26, 46

angustigena Foote, Trypeta 23, 449, 450, 451, 452, 453, 454, 457

basiola (Osten Sacken), Rhagoletis 252, 324, 325, 327, 328, 333, 335, 350, 352, 493, 496 beameri Norrbom and Ming, Oedicarena 249, 250, 250, 251

angustipennis (Loew), Tephritis 390, 391, 393 annulata Aczel, Rhagoletotrypeta 365, 367

bella (Loew), Euaresta 143, 144, 145, 147, 148, 149, 150, 151, 153

anthracina (Doane), Procecidochares 305, 305, 306, 310, 312, 313

bellula Snow, Euaresta 143, 145, 148, 149, 151 153

apicata (Thomas), Mylogymnocarena 56, 213, 214, 215, 215, 216

berberis Curran, Rhagoletis 326, 331, 332, 336, 337, 343, 360, 361

aplopappi (Coquillett), Aciurina 71, 73, 74, 79, 81

bicolor (Foote), Mylogymnocarena 213, 215, 216 bicolor (Stone), Anastrepha 85, 86, 89, 90, 91, 92, 94, 103

appendiculata Freidberg and Mathis, Neaspilota 225, 228, 229, 232, 233, 239 apple fruit fly 351 apple maggot 355 apple maggot fly 351 apple worm 355 araneosa (Coquillett), Tephritis 58, 61, 391, 392, 394, 395, 402, 404, 408 araneosa complex, Tephritis 400, 402, 404, 407

arcualis (Walker), Euaresta 146 arculata (Loew), Strauzia 374, 375, 376, 377, 378 arizonaensis Quisenberry, Tephritis 390, 391, 396, 397 arizonensis Malloch, Trupanea 418, 419, 421, 425, 429, 430, 434, 439

armata Robineau-Desvoidy, Strauzia 380, 381

bigeloviae (Cockerell), Aciurina 60, 72, 73, 74, 77, 78, 79

bigeloviae Townsend, Eurosta 74 bipuncta (Aczel), Xanthaciura 46, 481, 482 bipunctata Aczel, Chrysaciura 33 bisetosa (Coquillett), Trupanea 60, 417, 418, 419, 424, 430, 431, 436, 441, 442 bivittatus (Bigot), Dacus 46, 115, 132, 133 black cherry fruit fly 347 black fruit fly 105 black-bodied cherry fruit fly 347 blantoni Hering, Procecidochares 305, 306, 310, 311, 314 Blepharoneura Loew 24 blueberry fruit fly 353 blueberry maggot 353

armatus Fabricius, Dacus 132

borrichia Bush and Huettel, Cecidocharella 28, 51, 121, 123, 268

arnaudi Foote, Euarestoides 155, 156, 158, 160, 161

boycei Cresson, Rhagoletis 326, 327, 328, 338, 350

arnicae (Linnaeus), Tephritis 25, 387 artemisiae (Fabricius), Trypeta 448 Aschiza 17

Brachycera 17

Asilomorpha 17 Aspilomyia Hendel 222 Aspilota Loew 222, 230 asteri [error], Eurosta 177

asteris Flarris, Eurosta 175, 176, 177 aterrima (Doane), Oxyna 259, 260, 261 atra (Loew), Procecidochares 30, 51, 120, 303, 304, 305, 307, 308, 314, 317 augustipennis [error], Tephritis 394 aurantiaca (Doane), Xenochaeta 491, 491, 492 493

aurifera (Thomson), Dioxyna 135, 136

brevinervis Malloch, Xanthaciura 485 brunneistigma [error], Neaspilota 233 bnmneistigmata [error], Neaspilota 233 brunneostigma [error], Neaspilota 233 brunneostigmata Doane, Neaspilota 226, 233, 234, 238 bullans (Wiedemann), Euaresta 49, 142, 143, 144, 150, 151, 152, 154 bumelia fruit fly 103 Caenoriata Foote 26

calcitrapae Robineau-Desvoidy, Urellia 416 calcitrapae White and Korneyev, Urophora affinis ssp. 464

555

Index of Insect Names

California [error], Chetostoma 129 californica (Coquillett), Valentibulla 476, 477, 478, 480 californica Doane, Tephritis 390, 391, 396, 398, 404, 405, 406 californica Malloch, Trupanea 418, 422, 423, 424, 426, 427, 432 californica Novak, Paroxyna 278, 279, 281, 282 californicum Blanc, Chetostoma 127, 229, 129, 130 caliginosus Foote, Procecidocharoides 32 9, 320, 321, 322 caliptera (Say), Eutreta 183, 186, 187, 188, 194, 197, 198 Callachna Aldrich 21, 28, 50, 120, 122, 303 callistigma Freidberg and Mathis, Neaspilota 225, 226, 235, 240 calyptera [error or emendation], Eutreta 187 Calyptratae 17 Camaromyia Hendel 142 Campiglossa Rondani 32, 202 canadensis (Loew), Epochra 54, 139, 140 candidipennis Foote, Tephritis 392, 392, 393, 395, 396, 400, 402, 404, 408 capitata (Wiedemann), Ceratitis 22, 49, 51, 121, 124, 125, 126, 324 cardui (Linnaeus), Urophora 457, 459, 462, 463, 464, 470 Carib fly 109 Caribbean fruit fly 109 Carphotricha Loew 31, 262 Carphotriche [error] 266 Carphotrichia [error] 267 Carpomyia Costa 324 Carpomyina Norrbom 20, 23, 271, 324, 366 Carpotricha [error] 262, 265 caurina (Doane), Urophora 48, 459, 461, 465, 466, 468 Cecidocharella Hendel 21, 25, 28, 29, 52, 120, 122, 303, 318, 370 Cecidochares 28, 30 Cecidocharina Hering 21, 25, 27, 28, 29, 30 Cecidocharini 27 celery fly 162 Celidosphenella Hendel 32 Cerajocera Rondani 253 cerasi Linnaeus, Rhagoletis 323 Ceratites [error] 123 Ceratitini Bezzi 20, 22, 25 Ceratitis Macleay 20, 22, 42, 52, 123 Ceratodacus Hendel 24 Chaetorellia Hendel 2, 26 Chaetostoma Loew 128 Chaetostomella Hendel 5, 21, 25, 52, 126, 222, 223, 253, 270 cherry fruit fly 340, 348 cherry fruit fly, black 347 cherry fruit fly, black-bodied 347

cherry fruit fly, eastern 340 cherry fruit fly, western 348 cherry fruit fly, whitebanded 340 cherry maggot 340 Chetostoma Rondani 20, 23, 24, 55, 128, 248, 449 chiclayae Greene, Anastrepha 87, 88, 89, 92, 93, 95, 98 chilensis (Macquart), Dioxyna 134 chionanthi Bush, Rhagoletis 326, 329, 330, 339, 341, 349, 354 Chrysaciura Aczel 33, 46, 481, 482 Chrysotrypanea Malloch 25 chrysura (Thomson), Xanthaciura 481, 483, 483, 484, 486 cingulata group, Rhagoletis 326, 330, 351, 354 cingulata (Loew), Rhagoletis 326, 329, 330, 331, 339, 340, 349, 350, 354 circinata (Loew), Icterica 208, 210, 210 cislimitensis Steyskal, Cryptotreta 63, 131 citriperda Macleay, Ceratitis 123 claripennis Foote, Urophora 26, 458, 459, 466, 467, 469, 475 clathrata (Loew), Paroxyna 134, 136, 137, 278, 279, 283, 287 coalita Blanc, Eutreta 181, 182, 189, 190, 199 coloradensis Quisenberry, Paroxyna 276, 277, 284, 285, 286, 298 comma (Wiedemann), Eurosta 46, 167, 168, 168, 172, 173 completa Cresson, Rhagoletis 326, 327, 328, 331, 341, 342, 345, 351 concolor (Wulp), Trypeta 23, 449 confluens (Loew), Strauzia 374, 382, 383 conformis (Walker), Bactrocera 112 confusa (Curran), Eurosta 174 conjuncta (Adams), Trupanea 295, 418, 419, 424, 433, 434 connexionis Benjamin, Xanthaciura 482, 483, 484, 485, 486 conspurcata Doane, Eutreta 46, 171, 172 corniculata (Zetterstedt), Acinia 67 cornifera (Walker), Strauzia 381 cormgera (Walker), Strauzia 380, 381 cornivora Bush, Rhagoletis 326, 329, 330, 343, 344, 353, 359, 364 corpulenta Cresson, Paroxyna 289, 291 correcta (Bezzi), Bactrocera 112, 113, 114, 114, 115 crepidus (Hering), Carphotricha 31 cressoni Aczel, Tomoplagia 60, 411, 411, 412, 413, 414 cribrata (Wulp), Eurosta 46, 56, 166, 167, 170, 171, 174 Cryptaciura Hendel 23 Cryptotreta Blanc and Foote 21, 31, 64, 130 cucurbitae (Coquillett), Bactrocera 112, 113, 115 cucurbitat [error], Bactrocera 116

556

Index of Insect Names

culta (Wiedemann), Paracantha 46, 60, 262, 263, 264, 264, 265, 268, 270

Dracontomyia Becker 28, 29, 370 dreisbachi Foote, Euarestoides 32, 155

cultaris (Coquillett), Paracantha 60, 263, 264, 265, 266, 269 currant fly 140 currant fly, dark 360 currant fly, yellow 140 currant fruit fly 140

dreisbachorum Novak, Paroxyna 279, 284, 286 291, 296 dupla (Cresson), Paroxyna 276, 277, 282, 285, 286, 298, 301 Durosta [error] 165

currant fruit fly, American 360

Dyseuaresta Hendel 21, 28, 32, 59, 137

dubia (Johnson), Trypeta 455

curvicauda Gerstaecker, Toxotrypana 23, 48, 413, 414, 415 eastern cherry fruit fly 340

curvinerve Rondani, Chetostoma 128 cutta [error], Paracantha 264

ebbettsi Bush, Rhagoletis 326, 331, 332, 344, 345, 351, 360

dacetoptera Phillips, Trupanea 418, 419, 421, 424, 429, 432, 434, 439, 446 Dacidae Munro 35 daciformis Bezzi, Anastrepha 83 Dacina 18 Dacinae Loew 11, 15, 18, 20, 22, 25, 46, 112, 415 Dacini 15, 20, 22, 25, 112 Dacus Fabricius 2, 5, 6, 20, 22, 35, 42, 46, 47, 112, 132, 415 daphne (Wiedemann), Trupanea 427, 430, 440 447 dark currant fly 360 decora Stoltzfus, Eutreta 179, 181, 182, 189 199, 200 dentata (Stone), Anastrepha 49, 86, 87, 89, 90, 91, 92, 94, 103, 105 dertona (Walker), Eurosta 168

eclipta Benjamin, Trupanea 418, 420, 422, 425, 435, 444 edentata Stone, Anastrepha 86, 87, 89, 92, 93 95 electa (Say), Zonosemata 49, 492, 493, 494 494 495, 497 electellum (Hulst) [error], Neotephritis 246 electromorpha Berlocher, Rhagoletis 326, 331, 332, 345, 346, 351, 354, 360, 363 elegans (Blanchard), Parastenopa 24 elegans Hendel, Cecidocharella 122 elizabethae Quisenberry, Chetostoma 130 elongata Malloch, Paracantha 269 elsa Daecke, Eurosta 46, 168, 170 Ensina Robineau-Desvoidy 33, 55 Epochra Loew 20, 22, 55, 129, 140, 162 Epochroa [error] 140 Euaresta Loew 2, 9, 21, 27, 30, 32, 33, 62, 137, 138, 141, 388, 476

despecta [misidentification], Paroxyna 284 diana (Osten Sacken), Eutreta 63, 179, 180, 181, 182, 184, 190, 191, 192, 197

Euarestia [error], Rhagoletis 147 euarestoides (Bates), Valentibulla 479

dichromata Snow, Xenochaeta 58, 490, 491, 492 Dictyotrypeta Hendel 29, 30, 31

Euarestoides Benjamin 21, 30, 32, 64, 154, 213 Euarestopsis Hering 26

difficilis americana Hering, Paroxyna 289 diffusa (Snow), Gymnocarena 56, 201, 203, 204 diffusia [error], Gymnocarena 204

Eucosmoptera Phillips 216, 481 Euleia Walker 20, 23, 24, 57, 161, 248, 373, 449

dilacerata (Loew), Tephntis 61, 390, 391, 393 398, 400, 407

Euphranta Loew 22

Dioxyna Frey 5, 21, 31, 32, 62, 133, 142, 201, 259, 275, 388 disrupta Cockerell, Aciurma 74, 75 distincta Greene, Anastrepha 84, 89, 91, 93, 94 distincta Quisenberry, Paroxyna 277, 278, 284 285 Dithryca Rondani 27, 28 Dithrycini Hendel 14, 21, 27, 29, 31 Ditrichini 27 divisa Stoltzfus, Eutreta 180, 181, 182, 191 193, 197

192

dodsoni Foote, Valentibulla 476, 477, 478 dolosa Benjamin, Neaspilota 225, 227, 235 236 237

Euphrantini Hering 20, 22, 24, 140 Euraesta [error] 147 Euribia Meigen 67, 457 Eurosta Loew 21, 28, 29, 46, 57, 131, 165, 212, 476, 487, 490 Eurostina Curran 28, 46, 165 Eurostina Foote, Blanc and Norrbom 21, 29 euryptera (Loew), Paroxyna 280 Eutreta Loew 2, 5, 15, 21, 24, 28, 29, 30, 31, 46, 64, 179, 185, 212, 214, 487, 490, 491 Eutretini Munro 14, 21, 25, 27, 28, 29, 30, 31, 212, 487 Eutretosoma Hendel 29, 30 excelsa Aczel, Xanthaciura 33, 482 eximia (Wiedemann), Hexachaeta 205, 206

donysa (Walker), Eurosta 173 dorsalis (Hendel), Bactrocera 48, 112, 113 114 115, 116, 118, 119

facialis Curran, Eutreta 196 fallax Lima, Hexachaeta 206, 207, 207, 208, 209

Index of Insect Names

farinata Novak, Paroxyna 134, 136, 137, 278, 279, 185, 287, 288 fasciata (Adams), Tephritis 397 fascipennis Curran, Eurosta 46, 167, 168, 177, 178 fausta (Osten Sacken), Rhagoletis 325, 326, 327, 328, 334, 345, 347, 362 femoralis (Thomson), Trupanea 402, 417, 418, 419, 424, 430, 434, 435, 441, 447 fenestra Stoltzfus, Eutreta 181, 182, 189, 192, 199, 200 fenestrata Foote, Metatephritis 63, 180, 213, 214 fenestrata Snow, Eurosta 166, 167, 169, 170, 171, 172, 173 ferruginea (Doane), Aciurina 72, 76 festiva (Loew), Euaresta 61, 141, 143, 144, 146, 147, 149, 152, 154 fimbrata [error], Paracantha 266 fimbriata (Macquart), Paracantha 264 finalis (Loew), Neotephritis 61, 243, 244, 244, 245, 247 flava Foote, Gymnocarena 201, 203, 204, 204, 205 flaveola Coquillett, Trypeta 450, 451, 452, 452, 454 flavescens Robineau-Desvoidy, Oxyna 259 flavicans Robineau-Desvoidy, Orellia 253 flavinotata [error], Rhagoletis 330 flavipennis (Loew), Oxyna 259 flavipes Aldrich, Procecidochares 53, 304, 305, 310, 312, 315 flavissimus Foote, Procecidocharoides 319, 320, 321, 322 flavonotata (Macquart), Zonosemata 333, 334, 335, 494, 495 flavus (Adams), Euarestoides 32, 156, 158, 160 florescentiae [misidentification], Orellia 257, 258 floridana Ibrahim, Neaspilota 224, 226, 230, 231, 237 floridensis Foote, Eurosta 167, 168, 169, 170, 172, 173 floriscentiae [error], Orellia 258 fly, apple maggot 355 fly, Carib 109 fly, celery 162 fly, melon 115 fly, peacock 1 fly, walnut husk 341 footei Freidberg and Mathis, Neaspilota 225, 227, 228, 237 footeorum Novak, Paroxyna 276, 277, 287, 289, 298 Forellia Robineau-Desvoidy 448 forficula Benjamin, Paracantha 263, 264, 267, 268, 269 formosa (Coquillett), Urophora 459, 461, 467, 471 fractura (Coquillett), Trypeta 450, 451, 454 franciscana Hering, Paroxyna 290

fratercula [error], Anastrepha 96 fraterculus (Wiedemann), Anastrepha 83, 84, 88, 89, 91, 93, 95, 96, 99, 102, 110, 111 frateria [error], Euleia 162 fratria (Loew), Euleia 23, 56, 162, 162, 163, 164, 165, 449 frontalis Curran, Eutreta 183, 185, 191, 194, 198 fruit fly, American currant 360 fruit fly, apple 355 * fruit fly, black 105 fruit fly, black cherry 347 fruit fly, black-bodied cherry 347 fruit fly, blueberry 353 fruit fly, Bumelia 103 fruit fly, Caribbean 109 fruit fly, cherry 340, 348 fruit fly, currant 140 fruit fly, dark currant 360 fruit fly, eastern cherry 340 fruit fly, guava 114 fruit fly, Malaysian 118 fruit fly, Mediterranean 124 fruit fly, Mexican 98 fruit fly, Oriental 116 fruit fly, papaya 415 fruit fly, peach 119 fruit fly, pepper 118 fruit fly, pumpkin 132 fruit fly, Queensland 118 fruit fly, solanaceous 118 fruit fly, South American 95 fruit fly, West Indian 101 fruit fly, western cherry 348 fruit fly, whitebanded cherry 340 fucata (Fabricius), Acinia 68, 69 furcifera (Bigot), Toxotrypana 414, 416 fuscata Foote, Gonioxyna 32, 61, 202, 215 gemella (Coquillett), Euaresta 145, 146 geminata (Loew), Jamesomyia 58, 211, 212 genalis Malloch, Paracantha 263, 264, 268, 269 genalis (Thomson), Paroxyna 61, 63, 279, 280, 288, 290, 296 gentilis Hering, Paracantha 15, 262, 263, 264, 266, 267, 268, 269 germanna (Walker), Acinia 68 Gerrhoceras Hering 29 gibba (Loew), Callachna 28, 49, 51, 120, 121,

122 gibbosa [error], Callachna 122 gigantei Steyskal, Strauzia 376, 377, 378 goldenrod gall fly 175 Gonioxyna Hendel 5, 21, 31, 32, 62, 142, 200, 259, 295, 388 grindeliae Aldrich, Procecidochares 305, 306, 307, 310, 312, 313, 396 grindeliae (Coquillett), Urophora 459, 460, 467, 468,472

558

Index of Insect Names

guava fruit fly 114 gusano de la naranja 98 guttularis (Meigen), Dithryca 27 Gymnocarena Hering 21, 33, 57, 202, 215, 373 hamata (Loew), Anastrepha 105 hamifera Loew, Trypeta 448 hebes Curran, Trupanea 443 Hendrella Munro 27 heraclei (Linnaeus), Euleia 161, 162 hespera Banks, Eutreta 180, 183, 184, 185, 188, 194, 197 Hetschkomyia Hendel 28 Hexachaeta Loew 7, 20, 24, 50, 205 Hexaptilona Hering 24 Homoeothrix Hering 33 Hoplandromyia Bezzi 23 humilis (Loew), Dioxyna 135, 136 Icaria Schiner 179 Icterica Loew 15, 21, 33, 57, 209, 490 idahoensis Steyskal, Aciurina 70, 71, 73, 77 immaculata Blanc, Paraterellia 49, 271, 272, 273, 274 immaculata (Macquart), Trypeta 448 imperfecta (Coquillett), Trupanea 418, 419, 421, 425, 436, 437 inaequalis (Coquillett), Trypeta 450, 451, 452, 454, 455, 456 indifferens Curran, Rhagoletis 54, 326, 329, 330, 339, 341, 348 inermis Robineau-Desvoidy, Strauzia 273, 280 inornata (Coquillett), Neotephritis 246 insecta (Loew), Xanthaciura 58, 482, 483, 484,

485 intermedia (Loew), Strauzia 374, 375, 378, 379 intermedia Stoltzfus, Eutreta 46, 183, 194, 195,

200 interrupta Stone, Anastrepha 87, 88, 89, 92, 97, 98, 107 intrudens Aldrich, Rhagoletis 347 isochela Freidberg and Mathis, Neaspilota 225, 227, 228, 233, 236, 238 jaceae Robineau-Desvoidy, Acinia 67 jaceana (Hering), Urophora 460, 462, 463, 469, 473 jaculata Rondani, Urophora 460, 461, 462, 469, 472, 473 jamesi Novak, Paroxyna 277, 278, 289, 291, 293, 298 Jamesomyia Quisenberry 21, 31, 57, 65, 211, 487, 490, 491 johnsoni Thomas, Aciurina 80 jonesi Curran, Euaresta 143, 145, 148, 150, 151,

153 jonesi Curran, Eutreta 190 jonesi Curran, Trupanea 418, 420, 422, 423, 425, 435, 437, 438, 442, 444

juglandis Cresson, Rhagoletis 326, 327, 328, 338, 350 juniperina Marcovitch, Rhagoletis 324, 326, 331, 332, 333, 350, 351, 360 labecula Foote, Tephritis 389, 390, 401, 408, 409, 436 Laksayetsa Foote 29, 30 Lamproxyna Hendel 33 Lamproxynella Hering 32, 138 lantana gall fly 180 lateralis (Wiedemann), Eurosta 168, 169, 173 lathana Stone, Anastrepha 46, 99, 100 latifrons (Hendel), Bactrocera 112, 113, 114, 118 latifrons (Loew), Eurosta 165, 166, 167, 171, 172, 174 latifrons (Wulp), Oedicarena 51, 248, 249, 250, 250, 251 leavittensis Blanc, Tephritis 391, 392, 395, 396, 397, 402, 404, 408 Lilloaciura Aczel 33 limae Stone, Anastrepha 87, 88, 89, 92, 98 limata (Coquillett), Myoleja 24, 56, 217, 218, 219, 219, 220 liogaster (Thomson), Euleia 162 longicornis Macquart, Bactrocera 112 longicornis Snow, Eutreta 181, 182, 191, 193, 196, 198 longimacula Greene, Anastrepha 110 longipennis (Wiedemann), Strauzia 54, 56, 373, 374, 375, 376, 377, 378, 380, 381, 383, 384, 386, 387 longirostris Johnson, Rhynencina 48, 368, 369 longitudinalis (Loew), Strauzia 374, 381, 383 lucida (Fallen), Myoleja 216 Lucumaphila Stone 83, 84 ludens (Loew), Anastrepha 4, 5, 6, 8, 46, 84, 85, 86, 88, 89, 90, 91, 94, 98, 106 lunifera (Hering), Epochra 141 lurida (Loew), Chaetostomella 126 lutea (Coquillett), Aciurina 71, 76, 77, 78 macgregori Hernandez, Zonosemata 497 maculata (Cole), Aciurina 60, 70, 71, 73, 75,

78 maculata (Curran), Peronyma 301, 302 maculifemorata Hering, Paroxyna 292 maculifermorta [error], Paroxyna 293 maculigera Foote, Trupanea 418, 419, 421, 425, 429, 430, 434, 439, 446 maculosa (Coquillett), Trypeta 449, 450, 451, 454, 455, 456, 457 maggot, apple 355 maggot, blueberry 353 maggot, pepper 494 maggot, snowberry 364 maggot, sunflower 380 maggot, walnut husk 362 magniceps Hendel, Gonioxyna 200

559

Index of Insect Names Malaysian fruit fly 118 malgassa Munro, Ceratitis 124 marcetiae (Bezzi and Tavares), Parastenopa 24 marginepunctata Macquart, Paroxyna 46, 264, 266 Matsumuracidia Ito 23 maverna [error], Trupanea 440 medfly 124, 126 Mediterranean fruit fly 124, 324 meigenii (Loew), Rhagoletis 46, 325, 327, 328, 336, 337, 350, 352 melaneura [error], Acidogona 67 melanura (Loew), Acidogona 56, 65, 66, 67 melon fly 115 mendax Curran, Rhagoletis 326, 329, 330, 343, 344, 353, 359, 364 Metatephritis Foote 21, 30, 31, 62, 180, 213 mevarna (Walker), Trupanea 418, 420, 421, 422, 425, 426, 428, 439, 440, 445, 447 Mexican fruit fly 98 mexicana (Aczel), Aciurina 72, 73, 74, 79, 81 mexicana Aczel, Gymnocarena 215 mexicana Malloch, Paracantha 269 mexicana (Wiedemann), Dyseuaresta 60, 138, 139 michiganensis Quisenberry, Tephritis 391, 392, 403, 406 microsetulosa Malloch, Trupanea 432 microstigma Curran, Trupanea 437 Mikimyia Bigot 414, 416 mimetica Malloch, Paracantha 269 minuta (Snow), Procecidochares 304, 306, 311, 314 modocorum Blanc, Eutreta 181, 182, 189, 196 mombimpreoptans [error], Anastrepha 101 mombinpraeoptans Sein, Anastrepha 101 mombinpraeoptera [error], Anastrepha 101 Monacrostichus Bezzi 112 montana (Snow), Procecidochares 304, 305, 306, 311, 315, 317 Morelos orange worm 98 mouche de l’airelle 353 mouche de la pomme 355 mouche des brous du noyer 341 mouche du groseillier 140 mouche du piment 494 mouche du tournesol 380 munda (Coquillett), Valentibulla 53, 476, 477, 478, 479, 480 mundula (Coquillett), Valentibulla 477 mundulata Foote, Valentibulla 476, 477, 478, 479, 480 murina (Doane), Paroxyna 277, 278, 280, 292 Muscomorpha 17 Mylogymnocarena Foote 21, 33, 57, 203, 214 Myoleja Rondani 5, 21, 23, 24, 57, 129, 216, 373 Myopites Blot 26 Myopitinae 8 Myopitini Bezzi 14, 21, 24, 25, 26, 30, 369, 458

navajorum Blanc, Eutreta 46, 195 Neaspilota Osten Sacken 5, 9, 13, 21, 25, 30, 55, 127, 222, 223, 253, 270 Nematocera 17 Neoaspilota [error] 231 Neorellia Freidberg and Mathis 223, 226 Neorhabdochaeta Malloch 29, 30 Neorhagoletis Hendel 28, 29 Neospilota [error] 222, 240 Neotaracia Foote 26 Neotephritis Hendel 21, 33, 62, 65, 155, 213, 243, 388 nicholsoni Benjamin, Eurosta 173, 174 nigricornis (Coquillett), Trupanea 417, 418, 419, 424, 431, 436, 441 nigricornis (Doane), Myoleja 217, 218, 219, 220, 221 nigrifascia Stone, Anastrepha 87, 88, 92, 100 noctipennis Stoltzfus, Strauzia 374, 378, 383, 384 Noeeta Robineau-Desvoidy 28, 29, 31, 488 nora (Doane), Xanthomyia 488, 488, 489, 490 northern walnut husk fly 362 notata (Coquillett), Aciurina 72, 73, 79 novaeboracensis (Fitch), Eutreta 183, 184, 186, 187, 194, 195, 197 nuphera (Say), Eurosta 175 obfuscata (Wulp), Oedicarena 250, 251 obliqua (Macquart), Anastrepha 84, 87, 88, 89, 96, 101, 102, 110, 111 obliqua (Say), Tomoplagia 410, 411, 412, 413 obscura Hendel, Hexachaeta 206, 207, 207, 208 obscuripennis (Loew), Paroxyna 298 obscuripennis (Loew), Tetreuaresta 27 obscuripennis (Snow), Paroxyna 298 occidentalis Adams, Trupanea 435 occidentalis Novak, Paroxyna 277, 278, 291, 293, 294, 296, 298 occidentalis (Snow), Orellia 53, 128, 251, 254, 254, 255, 256, 258 ocbilleae [error], Neaspilota 229 ocresia (Walker), Anastrepha 85, 86, 92, 103, 105 Oedaspidina 27, 28, 29 Oedaspidini 24, 27 Oedaspis Loew 27, 28, 303, 307 Oedaspis-solidago Patton 303, 307 Oediaspis [error] 308 Oedicarena Loew 9, 11, 20, 23, 50, 202, 247, 373, 494 ogloblini (Blanchard), Anomoia 24 onopordinis (Fabricius), Euleia 161 onotrophes (Loew), Chaetostomella 126, 127 opaca (Coquillett), Aciurina 72, 73 , 75, 76, 77, 80 opacipennis (Foote), Paroxyna 46, 202, 276, 284, 293 orange fruit worm 98

560

Index of Insect Names

orange worm 98 oregona Curran, Eutreta 180, 181, 184, 189, 191, 196, 198 Orella [error] 258 Orellia Robineau-Desvoidy 5, 21, 26, 30, 55, 127, 128, 222, 223, 253, 270 oriental fruit fly 116 Orotava Frey 33, 210 osmanthi Bush, Rhagoletis 326, 329, 330, 339, 341, 348, 354 Ostracocoelia Giglio-Tos 29 Otitidae 18 ovatipennis Foote, Tephritis 391, 392, 395, 396, 402, 404, 405, 408 Oxyna Robineau-Desvoidy 5, 21, 31, 59, 201,

259 pacifica Curran, Aciurina 78 pacifica Curran, Eutreta 186 pacifica Doane, Tephritis 394 pallens Coquillett, Anastrepha 85, 86, 90, 91, 93, 94, 103 pallida (Cole), Eurosta 130, 131

piccola [error], Dioxyna 136 picturara [error], Acinia 68 picturata (Snow), Acinia 61, 66, 68, 69 Plagiotoma Loew 410, 411 plantaginis (Haliday), Paroxyna 407 Platensina 28 Platensinini 27, 28 platyptera (Loew), Xanthomyia 58, 437, 488,

489 Platystomatidae 18 Plaumannimyia Hering 32 pleuralis Aldrich, Procecidochares 304, 305, 309, 313, 315 Polionota Wulp 26, 27 polita (Loew), Procecidochares 304, 305, 311, 313, 315, 316 pollinosa Curran, Eutreta 181, 182, 189, 190, 195, 199 Polymorphomyia Snow 31 pomonella group, Rhagoletis 326, 330, 346, 348, 354, 359, 361, 363 pomonella (Walsh), Rhagoletis 15, 56, 324, 326, 329, 330, 331, 332, 341, 343, 353, 354, 355, 356, 364, 365

pallidipennis (Cresson), Paroxyna 223, 276, 277, 284, 295

Prececidochares [error] 303

palmeri Jenkins, Tephritis 391, 396, 399, 404, 405

Procecidochares Hendel 2, 6, 21, 25, 28, 29, 52, 120, 122, 303, 322

palpalis (Coquillett), Oxyna 60, 259, 260, 261 palposa (Loew), Orellia 53, 254, 255, 256, 258 papaya fruit fly 415

Procecidocharoides Foote 21, 28, 52, 122, 223, 270, 303, 318 Protensina Hendel 33

Paracantha Coquillett 5, 21, 29, 30, 46, 59, 262, 490, 491

Pseudacrotaenia Hendel 26, 46 Pseudeuleia Hering 217

Paracanthella Hendel 31 Paranoeeta Shiraki 31, 488 Parastenopa Hendel 23

Pseudeutreta Hendel 30, 31 Pseudodacus Hendel 83, 84, 92, 104 Pseudoedaspis Hendel 32, 138 Pseudopolionota Lima 26

Paraterellia Foote 20, 23, 50, 223, 270 Paroxyma [error] 275

pseudovicina Hering, Trupanea 418, 422, 423, 425, 437, 442, 444

Paroxyna Hendel 5, 21, 28, 31, 32, 46, 62, 134, 136, 142, 201, 259, 274, 388 parsnip leaf miner 162 peach fruit fly 119 peacock fly 1

pubescens Freidberg and Mathis, Neaspilota 224, 225, 226, 232, 235, 238 pucciola [error], Dioxyna 136

penelope (Osten Sacken), Procecidocharoides 28, 53, 318, 319, 320, 321, 322 pepper fruit fly 118 pepper maggot 494

pumpkin fruit fly 132

perfecta (Loew), Strauzia 374, 375, 376, 384, 385 Peronyma Loew 7, 21, 28, 29, 46, 52, 301, 303, 411

pullatus Foote, Procecidocharoides 53, 319, 320, 321, 322 punctata (Schrank), Orellia 253 punctistigma Benjamin, Neaspilota 225, 227, 238, 239 pura Boheman, Tephritis 406 pura (Loew), Tephritis 391, 392, 403, 405, 406, 410

persimilis Bush, Rhagoletis 326, 329, 330, 345, 346, 351, 354, 360, 363

pygmaea Novak, Paroxyna 277, 278, 293, 294,

persuasa (Osten Sacken), Oedicarena 248, 249 250, 251, 252

Pyrgotidae 18 Pyrgotoides Curran 29

Phaeogramma Grimshaw 32 Phasmatocephala Hering 179 Phobema Aldrich 83, 84

quadrifasciata (Macquart), Peronyma 46, 51, 301, 302, 308

Phorellia Robineau-Desvoidy 448 Phytalmiini 24 picciola (Bigot), Dioxyna 135, 136, 137

295

quadrifasciata (Meigen), Urophora 46, 459, 462, 463, 464, 465, 470 Queensland fruit fly 118

561

Index of Insect Names

radiata Schrank, Trupanea 416 radifera (Coquillett), Trupanea 418, 419, 422, 425, 435, 438, 443 railroad worm 355 rava Foote, Neotephritis 244, 245, 246, 247 regalis Munro, Parafreutreta 27 reticulata Snow, Eutreta 46, 170, 172 Rhabdochaeta de Meijere 29 Rhachiptera Bigot 25, 30 Rhacochlaena Loew 22 Rhagoletis Loew 2, 15, 20, 22, 23, 46, 55, 65, 140, 162, 248, 322 331, 365, 366, 367, 370, 493, 494 Rhagoletoides Foote 248 Rhagoletotrypeta Aczel 9, 20, 23, 55, 365 Rhagolitis [error] 347 rhino Steyskal, Myoleja 217, 218, 219, 220, 221 rhinophora Hering, Eutreta 179 Rhithrum Hendel 33 Rhynencina Johnson 21, 26, 30, 47, 368, 369, 458 ribicola Doane, Rhagoletis 326, 331, 332, 333, 334, 351, 360 361 ribicola group, Rhagoletis 326, 338, 361 Richardiidae 18 rohweri Foote, Rhagoletotrypeta 366, 366, 367, 368 rose hip maggot 333 rotundipennis (Loew), Eutreta 179, 182, 183, 184, 186, 192, 193, 199 rubidum (Coquillett), Chetostoma 54, 127, 129,

,

,

130 ruficauda (Fabricius), Orellia 254, 255, 256, 257, 258 rufipennis Doane, Tephritis 389, 390, 397, 398, 400, 407 rufipes (Curran), Urophora 459, 460, 466, 471 rugosum Stoltzfus, Strauzia 374, 383, 385 sabroskyi Novak, Paroxyna 279, 280, 291, 296, 297 sabroskyi Steyskal, Aciurina 75 sagittata (Stone), Anastrepha 83, 86, 87, 90, 91, 92, 94, 100, 104 sarcinata (Loew), Peronyma 46, 301, 302 sarnia (Walker), Paracantha 264 Schistopterini 25, 28, 29, 30 Schistopterum Becker 29 Schizophora 17 scorzorterae (Robineau-Desvoidy), Orellia 253 Scriptotricha [error] 262, 265 seabrai Lima, Flexachaeta 49, 206, 207, 208,

209 semilucida (Bates), Aciurina 75 sepentaria (Say), Strauzia 380 septenaria [error], Strauzia 381 seriata (Loew), Icterica 54, 58, 208, 209, 210,

211 sericata [error], Icterica 211 serpentaria [error], Strauzia 381

serpentina (Wiedemann), Anastrepha 83, 84, 85, 86, 90, 103, 105 106 setigera (Coquillett), Procecidochares 307 Setigeresta Benjamin 142, 146 setosa (Doane), Rhagoletis 335, 336 setosa Foote, Urophora 459, 460, 461, 466, 469,

,

471 Setosigena Stoltzfus 179, 180, 182, 193, 200 sigma (Phillips), Trypeta 449, 450, 454, 456, 457 signalis [error], Euaresta 145 signata Foote, Trupanea 418, 422, 423, 426, 442, 444 445 signatipennis Foote, Tephritis 390, 391, 396, 397, 402, 404, 407 signifer [error], Neaspilota 240 signifera (Coquillett), Neaspilota 224, 226, 235, 239, 240 simplex Thomas, Eutreta 179, 182, 183, 189, 190, 194, 196, 200 sirunaseva (Hering), Urophora 460, 461, 462, 466, 470, 472 Sitarea Robineau-Desvoidy 126, 128, 253 snowberry maggot 364 snowi Hering, Paroxyna 276, 277, 297, 298 solanaceous fruit fly 118 Solaris (Loew), Trupanea 439, 440 solidaginis (Fitch), Eurosta 46, 56, 165, 166, 167, 168, 172, 175 177 solstitialis (Linnaeus), Urophora 460, 461, 462,

,

,

473 sonchi (Linnaeus), Ensina 54 sororcula (Wiedemann), Dioxyna 56, 61, 63, 133, 134, 134 135, 137 sourbush seed fly 68 South American fruit fly 95 Sp. A Huettel and Bush, Procecidochares 309 Sp. A Wangberg, Aciurina 77 Sp. A Wangberg, Procecidochares 314 Sp. B Wangberg, Aciurina 77 Sp. B Wangberg, Procecidochares 314 Sp. C Wangberg, Procecidochares 314 Sp. E Brown, Anastrepha 97 Sp. F Brown, Anastrepha 95 Sp. W Brown, Anastrepha 100 sparsa (Wiedemann), Eutreta 179, 187, 194, 197 Spathulina Rondani 27 spatulata Stone, Anastrepha 87, 88, 93, 97, 98,

,

107 speciosa Hendel, Xanthaciura 33, 482 Spilographa Loew 202, 448, 449 stagmatica [error], Tephritis 408 Stamnophora Munro 26 stecki Freidberg and Mathis, Neaspilota 227, 229, 239, 240 stellata (Fuessly), Trupanea 416 stelligera (Coquillett), Euaresta 149, 150 Stenopa Loew 15, 21, 28, 29, 52, 120, 122, 303, 318, 369 stenoparia Steyskal, Urophora 459, 460, 461, 466, 468, 471, 472, 473

562

Index of Insect Names

steyskali Foote, Valentibulla 278, 476, 477, 478,

480 steyskali Novak, Paroxyna 277, 291, 293, 298, 299 stigmatica Coquillett, Euaresta 143, 144, 145, 148, 150, 151, 153 stigmatica (Coquillett), Tephritis 389, 390, 401, 402, 408 stoltzfusi Steyskal, Strauzia 375, 376, 377, 385, 386 stonei Blanc and Foote, Procecidochares 304, 305, 308, 314, 317, 318 stonei Steyskal, Anastrepha 84 straminea (Doane), Orellia 254 Straussia Loew 373

Tepbritoides Benjamin 388 Tephritus [error] 74 Terellia Robineau-Desvoidy 2, 26, 253 Terelliini Hendel 21, 24, 25, 30, 223, 253, 270, 271 tessellata (Loew), Paroxyna 275 testudinea (Loew), Acrotaenia 46, 63, 81, 82 tetanops (Loew), Oedicarena 247, 248, 252 tetraspina (Phillips), Xanthaciura 481, 482, 483, 484, 486 Tetreuaresta Hendel 2, 26, 63, 64 texana Hering, Trupanea 440, 442 texana Malloch, Trupanea 418, 420, 421, 425, 428, 440, 442, 445 447 Thephritis [error] 246

,

Strauzia Robineau-Desvoidy 5, 20, 23, 24, 55, 162, 373, 449

thomae (Curran), Dioxyna 133, 134, 135, 136,

striata Schiner, Anastrepha 84, 85, 86, 90, 106,

thoracica Curran, Aciurina 71, 74, 76, 79, 80 thurmanae Foote, Valentibulla 60, 476, 477, 478,

108 striatella group, Rhagoletis 326

137

481

striatella Wulp, Rhagoletis 326, 328, 329, 331, 333, 337, 343, 345, 347, 348, 353, 359, 361, 364

timberlakei Blanc and Foote, Urophora 458, 459, 467, 468, 469, 475 Tomoplagia Coquillett 9, 21, 26, 27, 30, 59, 302, 410 412

,

Strobelia Rondani 30 Strumeta Walker 112

Tomoplagina Curran 301

stylata (Fabricius), Urophora 460, 461, 462, 465, 466, 474 Stylia Robineau-Desvoidy 275 suavis group, Rhagoletis 326, 328, 338, 348 suavis (Foew), Rhagoletis 326, 328, 337, 345, 348, 362 subfasciata Curran, Eurosta 46, 178 subfascipennis [error], Eurosta 178 subpura (Johnson), Tephritis 61, 388, 389, 390, 398, 399, 409 sunflower maggot 380

tortile Coquillett, Trypeta 449, 450, 453, 456 Toxotrypana Gerstaecker 5, 6, 15, 20, 22, 50, 84, 414 Toxotrypanea Curran 84, 414 Toxotrypanini Munro 20, 22 Toxytrypana [error] 414 tribulus (Say), Tephritis 406 tricolor (Doane), Gymnocarena 201, 203, 204,

205 tricolor Snow, Eutreta 190 trilitura Blanc and Foote, Aciurina 70, 71, 75, 76, 81

superba Foote, Paraterellia 271, 272, 273, 274 suspensa (Loew), Anastrepha 23, 84, 85, 88, 89, 90, 91, 96, 99, 101, 102, 109 111 symphoricarpi Curran, Rhagoletis 364

trimaculata (Macquart), Strauzia 380 trixa Curran, Aciurina 69, 74, 75 Trupanea Guettard 32, 416

Tabanomorpha 17

Trupanea Schrank 9, 21, 25, 32, 59, 138, 155, 216, 388, 402, 408, 409, 416

,

tabellaria (Fitch), Rhagoletis 324, 326, 327, 332, 346, 351, 354, 355, 360, 363 tabellaria group, Rhagoletis 326, 344 tabernella [error], Rhagoletis 363 Tachiniscidae 18 tapetis (Coquillett), Euaresta 143, 144, 151, 154 tapsetus [error], Euaresta 154 tenebrosa (Coquillett), Paroxyna 279, 284, 291

300 Tephrella Bezzi 69, 70 Tephrelliini 24, 25, 27, 33

tryoni (Froggatt), Bactrocera 112, 113, 114, 115,

118 Trypanaresta Hering 25, 32 Trypanea [emendation] 416 Trypeta Meigen 5, 20, 23, 24, 55, 65, 162, 248, 374, 448 trypete des cerises 340 trypete noire des cerises 347 Trypetina 18, 20, 23, 374, 449

Tephritinae 11, 14, 18, 21, 22, 24 27, 29, 30, 31, 33, 180, 411

Trypetinae 11, 15, 18, 20, 22, 25, 26, 29, 33, 217, 270, 271, 411 Trypetini 20, 22, 23 Tryreta [error] 253

Tephritini 14, 21, 25, 27, 28, 30, 31, 35

typica (Loew), Strauzia 374, 382, 383

Tephritis Latreille 9, 14, 21, 25, 28, 32, 62, 137, ^ 138, 142, 180, 259, 295, 387, 416 Tephritoidea 18

Uncaculeus Stoltzfus 29, 30, 31, 179, 180, 196, 214

,

563

Index of Insect Names

uncinata (Coquillett), Euleia 162, 163, 164, 165 undosa (Coquillett), Chaetostomella 53, 127, 128, 255 unifasciata Blanc and Foote, Myoleja 24, 217, 218, 220, 221 uniformis Steyskal, Rhagoletotrypeta 366, 367, 368 unipuncta Sein, Anastrepha 110 Urellia Robineau-Desvoidy 416 Urelliosoma Hendel 32 Urophora Robineau-Desvoidy 2, 5, 11, 26, 30, 39, 42, 46, 47, 333, 368, 369, 457 utahensis Quisenberry, Oxyna 259, 260, 261 utilis Stone, Procecidochares 304 uvedaliae Stoltzfus, Strauzia 374, 383, 386 Valentibulla Foote and Blanc 21, 29, 59, 69, 475 valida Lima, Hexachaeta 206 variabilis (Doane), Paroxyna 276, 277, 298, 299, 300 variegata Aczel, Zonosemata 497 varipennis (Coquillett), Paraterellia 271, 272, 273 verbesinae Steyskal, Strauzia 376, 377, 378, 387 vernoniae (Loew), Neaspilota 53, 223, 224, 229, 231, 234, 241 versatilis (Curran), Paraterellia 273 vicina (Wulp), Trupanea 418, 422, 423, 426, 429, 446 viciniformis Foote, Trupanea 418, 422, 423, 426, 430, 446 Vidalia Robineau-Desvoidy 23, 449 virens (Loew), Terellia 2 viridescens Quisenberry, Neaspilota 224, 225, 227, 229, 236, 242 vittigera (Coquillett), Zonosemata 49, 51, 492, 494, 495, 496 vittigera (Loew), Strauzia 374, 382, 383 vulnerata (Loew), Stenopa 53, 369, 370, 371, 372

walnut husk fly 341 walnut husk fly, northern 362 walnut husk maggot 362 webbii Doane, Tephritis 391, 392, 403, 406, 409 West Indian fruit fly 101 western cherry fruit fly 348 wheeleri Curran, Trupanea 58, 418, 420, 421, 422, 425, 426, 428, 440, 445, 447 whitebanded cherry fruit fly 340 Whiteina Korneyev 32 wilsoni Blanc and Foote, Neaspilota 224, 226, 233, 235, 242 wolffi (Cresson), Euaresta 151 worm, apple 355 worm, Morelos orange 98 worm, orange 98 worm, orange fruit 98 worm, railroad 355 Xanthaciura Hendel 2, 21, 33, 46, 59, 216, 217, 481 xanthogastra Aczel, Rhagoletotrypeta 365 Xanthomyia Phillips 21, 31, 57, 212, 487, 490 Xenochaeta Snow 21, 31, 57, 65, 180, 212, 323, 487, 490 yellow currant fly 140 ypsilon Foote, Paraterellia 271, 272, 273, 274 zephrina [error], Rhaqoletis 364 zephyria Snow, Rhagoletis 326, 329, 330, 332, 343, 344, 353, 359, 364 zonata (Saunders), Bactrocera 112, 113, 114, 115, 119 Zonosema Loew 334, 493 Zonosemata Benjamin 15, 20, 23, 50, 65, 252, 323, 324, 366, 493 zoqui Bush, Rhagoletis 326, 327, 328, 338 zuelaniae Stone, Anastrepha 88, 90, 93, 111

INDEX OF PLANT NAMES

Acamptopappus [Asteraceae] 230, 442, 448 Acamptopappus shockieyi 475

Ambrosia dumosa 160 Ambrosia ilicifolia 437 Ambrosia trifida 152, 385

Acamptopappus sphaerocephalus 241, 475 Acanthaceae 27, 33 acanthicarpa, Ambrosia 160 Achillea [Asteraceae] 448 Achillea ptarmica 394

Amphipappus [Asteraceae] 428, 442 Anacardiaceae 102

Achras [Sapotaceae] 100, 115 Achyrachaena [Asteraceae] 438 acradenius, Haplopappus 235, 471

Anacardium [Anacardiaceae] 96, 100, 102 Anaphalis [Asteraceae] 160, 433, 445 Anaphalis margaritacea 433

Acrolophus, Centaurea subg. [Asteraceae] 464

Annona [Annonaceae] 96, 100, 102, 107, 108 110

Actinospermum [Asteraceae] 136, 428 adenophorum, Ageratina 304 Aegiphila [Verbinaceae] 206 Aegle [Rutaceae] 115 affinis, Sorocea 206

annuus, Helianthus 204, 377, 383, 431 annuus ssp. lenticularis, Helianthus 267 annuus var. macrocarpus, Helianthus 267 Antennaria [Asteraceae] 438 Anthemideae 28, 30, 388 Apiaceae 23, 164 Apocynaceae 23, 416 apple 324 apple, rose 102

Ageratina adenophorum [Asteraceae] 304 Ageratina altissima 383 Ageratina nparia 304 Ageratum [Asteraceae] 485 Ageratum houstonianum 487 Ageratum littorale 428, 485, 486 Agoseris [Asteraceae] 438 Agoseris grandiflora 301 alba, Eclipta 435 albiflora, Lonicera 130 Alchornea [Euphorbiaceae] 96, 102, 107 altissima, Ageratina 383 altissima, Solidago 177, 198 Ambrosia [Asteraceae] 33, 122, 142, 150, 153, 160, 187, 188, 314, 400, 448 Ambrosia acanthicarpa 160 Ambrosia artemisiifolia 149 Ambrosia chamissonis 153, 160

564

Amelanchier [Rosaceae] 78, 360 americanus, Osmanthus 354

Aquifoliaceae 24 arbuscula ssp. nova, Artemisia 214 Arnica [Asteraceae] 160, 396, 400, 408, 438, 452 Arnica Arnica Arnica Arnica Arnica Aronia

cordifolia 288, 298 diversifolia 402 latifolia 288 rydbergi 288 sororia 282 [Rosaceae] 360

Artemisia [Asteraceae] 30, 75, 180, 192, 197, 259, 396, 438, 452 Artemisia arbuscula ssp. nova 214 Artemisia cana 196

565

Index of Plant Names

Artemisia filifolia 192 Artemisia ludoviciana 200 Artemisia tridentata 192, 199, 202, 261 Artemisia tridentata var. vaseyana 261 atemisiifolia, Ambrosia 149 artichoke 270 arvense, Cirsium 258, 465 arvensis, Sonchus 401 Asclepiadaceae 23, 416 Aster [Asteraceae] 136, 194, 229, 234, 237, 242, 281, 309, 314, 408, 414, 428, 436, 438, 444, 448 Aster lateriflorus 170 Aster occidentalis var. intermedius 394 Aster spinosus 242 Asteraceae 14, 23, 24, 27, 30, 33, 70, 82, 164, 180, 188, 223, 247, 271, 374, 388, 411, 452 Astereae 26, 28, 29, 30, 31, 32, 180, 275, 388, 458 Atalantia [Rutaceae] 110 Atrichoceris [Asteraceae] 444 aureus, Senecio 373 Averrhoa [Oxalidaceae] 96, 102, 110 Baccaurea motleyana [Euphorbiaceae] 118 Baccharis [Asteraceae] 30, 31, 32, 81, 389, 396, 398, 407, 436, 438, 442, 448 Baccharis emoryi 399 Baccharis glomulifera 409 Baccharis glutinosa 160, 398, 439 Baccharis halimifolia 405, 409 Baccharis neglecta 405 Baccharis pilularis 399, 407 Baccharis pilularis consanguinea 160, 399, 445 Baccharis salicifolia 398 Baccharis sarothroides 79, 397 Baccharis sergiloides 397 Baccharis viminea 161, 398 Baeria Fremontii [Asteraceae] 240 Baileya [Asteraceae] 438, 444 baldwini, Vernonia 231 Balsamorhiza [Asteraceae] 438, 444 Bebbia [Asteraceae] 75 Bebbia juncea 437, 442 beneolens, Gnaphalium 433 Berberidaceae 325, 326, 350 Berberis nervosa [Berberidaceae] 337 Berberis vulgaris 328, 352 berlanderi, Lycium 333 bicolor, Gnaphalium 433 Bidens [Asteraceae] 33, 136, 188, 210, 211, 485, 486 Bidens bipinnata 137 Bidens cernua 211 Bidens squarrosa 486 biennis, Helianthus 266 Bigelovia [Asteraceae] 75 bigelovii, Chrysothamnus nauseosus ssp. 479 bipinnata, Bidens 137

Bischofia [Euphorbiaceae] 110 black cherry 330, 341 blakei, Haplopappus ericoides ssp. 466 Blepharipappus [Asteraceae] 438 Blighia [Sapindaceae] 110 bloomeri, Haplopappus 293 blueberry 332, 354 Borrichia [Asteraceae] 267 Borrichia frutescens 123, 268 Brickellia [Asteraceae] 312, 314, 428, 436, 438, 442, 444, 448 Britoa [Myrtaceae] 110 Brosimum [Moraceae] 102 bull thistle 474 Bumelia [Sapotaceae] 107, 108 Bumelia celastrina 104 Bumelia spimflora 104 butternut 363 Byrsonima [Malpighiaceae] 107 Cacalia [Asteraceae] 452 calcitrapa, Centaurea 473 californica, Cirsium 269 californica, Solidago 307 californica, Trixis 413, 428, 433 californicum, Gnaphalium 433 calliosidea, Coreopsis 243 Callistephus [Asteraceae] 187 Calycoseris [Asteraceae] 444 camara, Lantana 180 campechiana, Pouteria 105 camphor daisy 311 Campomanesia [Myrtaceae] 96 cana, Artemisia 196 Canada thistle 465 canadensis, Conyza 310 canadensis, Erigeron 238 canadensis, Lactuca 212 canadensis, Solidago 166 canadensis var. hargeri, Solidago 177 Canella [Canellaceae] 110 canescens, Geraea 431 canescens, Machaeranthera 233, 408 Caprifoliaceae 326, 365 Capsicum [Solanaceae] 110 Cardueae 26, 458 Carduus [Asteraceae] 473, 474 Careya [Lecythidaceae] 115 Carica [Caricaceae] 100, 110 Caricaceae 23 Carissa [Apocynaceae] 110 Carthamus [Asteraceae] 442 Casimiroa [Rutaceae] 100, 110 celastrina, Bumelia 104 Celtis [Ulmaceae] 366, 367 Celtis laevigata 367 Centaurea [Asteraceae] 464, 469, 471 Centaurea calcitrapa 473 Centaurea diffusa 464, 471

566

Index of Plant Names

Centaurea idaea 472 Centaurea nigra 469 Centaurea solstitialis 470, 473 Centaurea sp. near maculosa 464, 471 Centaurea squarrosa 471 cernua, Bidens 211 Chaenactis [Asteraceae] 438, 442 chamissonis, Ambrosia 153, 160 cherry 324, 341 cherry, black 330, 341 cherry, fire 348 cherry, pin 348, 349 chilense, Gnaphalium 433, 445 chili pepper 118 Chionanthus [Oleaceae] 330 Chionanthus virginicus 340 Chromolaena odorata [Asteraceae] 485 Chrysanthemum [Asteraceae] 160, 187, 188, 452 Chrysobalanus [Rosaceae] 110 chrysophylloides, Schoepfia 97 Chrysophyllum [Sapotaceae] 95, 107, 108, 110 Chrysopsis [Asteraceae] 229, 237, 238, 240, 438, 448 Chrysopsis villosa 242 Chrysothamnus [Asteraceae] 75, 76, 78, 202, 234, 238, 242, 307, 314, 317, 396, 438, 442, 444, 448, 468, 475 Chrysothamnus nauseosus 75, 79, 80, 81, 475, 476, 478, 480 Chrysothamnus nauseosus hololeucus 402 Chrysothamnus nauseosus ssp. bigelovii 479 Chrysothamnus teretifolius 475 Chrysothamnus viscidiflorus 77, 78, 468, 475 ciliaris, Helianthus 431

Coreopsis gigantea 437 Corethrogyne [Asteraceae] 234, 314, 404, 436, 448 Corethrogyne filaginifolia 239, 318 Cornaceae 326 Cornus [Cornaceae] 324, 330, 332, 343, 363 Cornus drummondi 346 Cornus racemosa 346 Cotoneaster [Rosaceae] 360 crassulus, Senecio 409 Crataegus [Rosaceae] 96, 102, 359, 360 Crepis [Lactuceae] 31 creticum, Cirsium 465 Crofton weed 304 Cucumis melo [Cucurbitaceae] 133 Cucumis sativus 133 cucurbit 116, 133 Cucurbita pepo [Cucurbitaceae] 133 cuneatus, Haplopappus 474, 475 Cupressaceae 326 currant 333 Cydonia [Rosaceae] 96, 100, 107 Cynara [Asteraceae] 452 Cynara scolymus 266 Cynareae 30 cynoglossoides, Hieracium 493 Dahlia coccinea [Asteraceae] 486 daisy, camphor 311 decipiens, Juniperus 272 deppiana, Juniperus 273 devilwood 354 Dicoria [Asteraceae] 442 diffusa, Centaurea 471

Cirsium [Asteraceae] 128, 255, 257, 262, 266, 269, 270, 474

Diospyros [Ebenaceae] 96, 100, 102, 107, 108,

Cirsium Cirsium Cirsium Cirsium Cirsium

diversifolia, Arnica 402 dogwood 332 douglasii, Senecio 409

arvense 258, 465 californica 269 congdonii 269 creticum 465 vulgare 474

Citrofortunella [Rutaceae] 107 Citrofortunella X mitis 107, 110 Citrus [Rutaceae] 96, 100, 102, 107, 108, 110, 111, 115, 324 Clausena [Rutaceae] 110 Cnicus parryi [Asteraceae] 255 coccinea, Dahlia 486 Coccoloba [Polygonaceae] 110 cocklebur 142 Coffea [Rubiaceae] 96, 100, 102, 115 coffee 133 condensatus, Elymus 287 congdonii, Cirsium 269 consanguinea, Baccharis pilularis ssp. 399, 445 Conyza canadensis [Asteraceae] 310 cordifolia, Arnica 288, 298 Coreopsis [Asteraceae] 136, 311, 428, 438, 446 Coreopsis calliosidea 243

110

douglasii var. douglasii, Senecio 285 douglasii var. monoensis, Senecio 285 Dovyalis [Flacourtiaceae] 96, 102, 107, 110 drummondi, Cornus 346 Drypetes [Euphorbiaceae] 110 dumosa, Ambrosia 160 Dyssodia [Asteraceae] 442, 444, 446 Eclipta alba [Asteraceae] 435 Eclipta prostrata 435 eggplant 118, 496 eleagnifolium, Solanum 497 Elephantopus [Asteraceae] 27 Elymus condensatus [Gramineae] 287 emarginata, Manilkara 100 emarginata, Prunus 348, 349 emoryi, Baccharis 399 Encelia [Asteraceae] 438, 442, 448 Ericaceae 326, 330 ericoides ssp. blakei, Haplopappus 466

567

Index of Plant Names

Erigeron [Asteraceae] 229, 234, 237, 242, 307, 396, 404, 428, 438, 442, 444, 448 Erigeron canadensis 238 Erigeron peregrinus 287 Erigeron pusillus 310 Eriobotrya [Rosaceae] 96, 102, 110 Eriophyllum [Asteraceae] 291, 436, 438, 444, 448 Eugenia [Hydrophyllaceae] 95, 96, 102, 110, 115, 119 Eupatorieae 28 Eupatorium [Asteraceae] 485, 487 Eupatorium glandulosum 485 Eupatorium riparium 485 Eupatorium rugosum 322, 383, 386 exaltatus, Senecio integerrimus var. 285 Feijoa [Myrtaceae] 96 Ficus [Moraceae] 96, 107, 110, 115 filaginifolia, Corethrogyne 239, 318 filifolia, Artemisia 192 fire cherry 348 fistulosa, Solidago 173 Flacourtia [Flacourtiaceae] 110 Fortunella [Rutaceae] 96, 97, 110 Fragaria [Rosaceae] 97 Fremontii, Baeria 240 frutescens, Borrichia 123, 268 Gaillardia [Asteraceae] 160 Galactites [Asteraceae] 474 Garcinia [Guttiferae] 110 Gaylussacia [Ericaceae] 354 Geoffraea [Leguminosae] 102 Geraea [Asteraceae] 438, 442 Geraea canescens 431 gigantea, Coreopsis 437 gigantea, Solidago 177 giganteus, Helianthus 379 glandulosum, Eupatorium 485 glomulifera, Baccharis 409 glutinosa, Baccharis 160, 398, 439 Gnaphalium [Asteraceae] 428, 433, 438, 439 Gnaphalium beneolens 433 Gnaphalium bicolor 433 Gnaphalium californicum 433 Gnaphalium chilense 433, 445 Gnaphalium luteo-album 433, 445 Gnaphalium microcephalum 433 Gnaphalium obtusifolium 435 Gnaphalium palustre 433 Gnaphalium ramissimum 433 Godmania [Bignoniaceae] 102 goldenrod 166, 195 Gonolobus [Asclepiadaceae] 416 Goodingii, Haplopappus 230 gooseberry 333 gracile, Porophyllum 442 Gramineae 287

graminifolia, Heterotheca 441 graminifolia, Solidago 177 grandiflora, Agoseris 301 grape, Oregon 332, 337 grapefruit 100 Grindelia [Asteraceae] 238, 313, 466, 468 Grindelia nana var. nana 291 Grindelia squarrosa var. nuda 469 grindelioides, Haplopappus squarrosus ssp. 243, 468 grosseserratus, Helianthus 377, 383, 384 guajava, Psidium 103 guava 102, 108 guidonia, Zuelania 111 Gutierrezia [Asteraceae] 242, 475 Gutierrezia microcephala 474, 475 Gutierrezia sarothrae 74, 466, 474 hackberry 367 halimifolia, Baccharis 405, 409 Hamakua pamakani 304 Haplopappus [Asteraceae] 75, 76, 234, 242, 291, 428, 436, 438, 442, 444, 448, 468 Haplopappus acradenius 235, 471 Haplopappus bloomeri 293 Haplopappus cuneatus 474, 475 Haplopappus ericoides ssp. blakei 466 Haplopappus Goodingii 230 Haplopappus hartwigi 230 Haplopappus laricifolius 475 Haplopappus pinifolius 74, 285 Haplopappus squarrosus ssp. grindelioides 243, 468 Haplopappus venetus 468 Haplopappus venetus var. vernonioides 235 hartwigi, Haplopappus 230 Helenium [Asteraceae] 136, 160, 438 Heliantheae 28, 30, 31, 33, 247, 262 Helianthella [Asteraceae] 442 Helianthus [Asteraceae] 188, 204, 267, 383, 448 Helianthus annuus 204, 377, 383, 431 Helianthus annuus ssp. lenticularis 267 Helianthus annuus var. macrocarpus 267 Helianthus biennis 266 Helianthus ciliaris 431 Helianthus giganteus 379 Helianthus grosseserratus 377, 383, 384 Helianthus niveus ssp. tephrodes 431 Helicostylis [Moraceae] 206 Hemizonia [Asteraceae] 436, 448 Hemizonia pungens 240 Heterotheca [Asteraceae] 234, 237, 311, 408, 436, 441, 444 Heterotheca graminifolia 441 Heterotheca latifolia 310, 441 Heterotheca microcephala 435, 441 Heterotheca oligantha 441 Heterotheca oregana 311 Heterotheca subaxillaris 310

568

Index of Plant Names

Heterotheca trichophylla 302 Heterotheca villosa 311, 318 Heterotheca villosa var. sessiflora 311 Hieracium [Asteraceae] 67, 229, 428 Hieracium cynoglossoides 493 Holocarpa [Rubiaceae] 436 hololeucus, Chrysothamnus nauseosus ssp. 402 houstonianum, Ageratum 487 Hulsea [Asteraceae] 438 husk tomato 328, 361 Hymenoclea salsola [Asteraceae] 474 Hymenoxys [Asteraceae] 438, 442

lenticularis, Helianthus annuus ssp. 267 Lepidospartum [Asteraceae] 242, 436, 438, 439, 448 Lessingia [Asteraceae] 436, 439 Lessingia lemmonii 234 Leucelene [Asteraceae] 442 Liabeae 27, 411 littorale, Ageratum 428, 486 Lonicera albiflora [Caprifoliaceae] 130 Lucuma [Sapotaceae] 105 ludoviciana, Artemisia 200 luteo-album, Gnaphalium 433, 445 Lycium berlanderi [Solanaceae] 333

idaea, Centaurea 472 Ilex [Aquifoliaceae] 24, 217, 220 Ilex paraguaiensis 24 ilicifolia, Ambrosia 437 incanum, Solanum 118 indica, Mangifera 100 indicum, Solanum 118 Inga [Leguminosae] 95, 97, 100 integerrimus, Senecio 409 integerrimus var. exaltatus, Senecio 285 interior, Vernona 231, 241 intermedius, Aster occidentalis var. 394 Iva [Asteraceae] 160 Jacaratia mexicana [Caricaceae] 416 Jacea, Centaurea subg. [Asteraceae] 469 Jambosa [Myrtaceae] 102 Juglandaceae 326 Juglans [Juglandaceae] 97, 324, 328, 338, 343, 350, 362 juncea, Bebbia 437, 442 juncea, Solidago 170, 172 juniper 333 Juniperus [Pinaceae] 271, 274, 324, 333, 352 Juniperus decipiens 272 Juniperus deppiana 273 knapweed 458, 464, 471 Labiatae 33 laciniata, Microseris 301 laciniata, Rudbeckia 380 laciniata, Viguiera 317 Lacmellea [Apocynaceae] 107 Lactuca canadensis [Asteraceae] 212 Lactuceae 31, 275 laevigata, Celtis 367 Lagenaria [Cucurbitaceae] 115 Lantana camara [Verbenaceae] 180 laricifolius, Haplopappus 475 Lasthenia [Asteraceae] 438, 444 lateriflorus, Aster 170 latifolia, Arnica 288 latifolia, Heterotheca 310, 441 Layia [Asteraceae] 291, 438 lemmonii, Lessingia 234

Machaeranthera [Asteraceae] 230, 234, 242, 436, 444 Machaeranthera canescens 233, 408 Machaeranthera phyllocephala 311 Machaeranthera tortifolia 233 macrocarpus, Helianthus annuus ssp. 267 maculosa, Centaurea 464, 471 Madia [Asteraceae] 291, 439, 442 Mahonia [Berberidaceae] 332, 337 Malacothrix [Asteraceae] 444 Malpighia [Malpighiaceae] 97, 102, 110 Malus [Rosaceae] 97, 100, 102, 107, 110, 360 Mammea [Guttiferae] 100, 107 Mangifera [Anacardiaceae] 95, 97, 100, 102, 107, 108, 110, 115 Mangifera indica 100 mango 102, 119, 416 Manihot [Euphorbiaceae] 108 Manilkara [Sapotaceae] 97, 102, 107, 110, 115 Mamlkara emarginata 100 Manilkara zapota 100, 103 Marcetia [Melastomataceae] 24 margantacia, Anaphalis 433 Mastichodendron [Sapotaceae] 97, 100, 107 Maui pamakani 304 melanocerasum, Solanum 496 Melanthera [Asteraceae] 139 Melastomataceae 24 melo, Cucumis 133 mexicana, Jacaracia 416 microcephala, Gutierrezia 474, 475 microcephala, Heterotheca 435, 441 microcephala, Perezia 413, 442, 448 microcephalum, Graveolens 433 Micropholis [Sapotaceae] 107 Microseris [Asteraceae] 291, 444 Microseris laciniata 301 Mimusops [Sapotaceae] 100, 107, 110 missouriensis, Solidago 172 Momordica [Cucurbitaceae] 110, 111, 116, 133 monoensis, Senecio douglasii var. 285 Monolopia [Asteraceae] 436, 439, 442 Monoptilon [Asteraceae] 444 Moraceae 24, 206 Morus [Moraceae] 206, 207

569

Index of Plant Names

motleyana, Baccaurea 118 mulberry 207 multilobatus, Senecio 371 Muntingia [Elaeocarpaceae] 111 Murraya [Rutaceae] 111 Mutisieae 27, 411, 413 Myrcia [Myrtaceae] 97, 108 Myrcianthes [Myrtaceae] 111 Myrciaria [Myrtaceae] 111 nana var. nana, Grindelia 291 nauseosus, Chrysothamnus 75, 79, 80, 81, 475, 476, 478, 480 nauseosus hololeucus, Chrysothamnus 402 nauseosus ssp. bigelovii, Chrysothamnus 479 neglecta, Baccharis 405 nervosa, Berberis 337 Nicolletia [Asteraceae] 444, 446 nigra, Centaurea 469 niveus ssp. tephrodes, Helianthus 431 nodding thistle 473 nuda, Grindelia squarrosa var. 469 obtusifolium, Gnaphalium 435 occidentalis var. intermedius, Aster 394 occidentalis, Verbesina 387 odorata, Chromolaena 485 Oleaceae 326, 330 oligantha, Heterotheca 441 orange 100, 120 oregana, Heterotheca 311 Oregon grape 332, 337 Osmanthus [Oleaceae] 330, 354 Osmanthus americanus 354 Otopappus [Asteraceae] 82 Palafoxia [Asteraceae] 444 palustre, Gnaphalium 433 papaya 416 paraguaiensis, Ilex 24 parryi, Cnicus 255 parsnip 164 Passiflora [Passifloraceae] 93, 100, 102, 116, 119 Passiflora quadrangularis 98 peach 120, 343, 362 Pectis [Asteraceae] 446 pennsylvanica, Prunus 348 pepo, Cucurbita 133 pepper 116, 118, 494, 496 pepper, chili 118 peregrinus, Erigeron 287 Perezia [Asteraceae] 436, 442, 448 Perezia microcephala 413, 442, 448 Perityle [Asteraceae] 439, 444, 448 Persea [Lauraceae] 97, 100, 107 persica, Prunus 343, 413 Petasites [Asteraceae] 452 Peucephyllum [Asteraceae] 436 Phoenix [Palmae] 111

phyllocephala, Machaeranthera 311 Physalis [Solanaceae] 328, 361, 496 pilularis, Baccharis 399 pilularis consanguinea, Baccharis 399, 407 pilularis ssp. consanguinea, Baccharis 160, 445 Pimenta [Myrtaceae] 111 pin cherry 348 pin cherry, wild 349 pineapple 119 pinifolius, Haplopappus 74, 285 Pleurocoronis [Asteraceae] 442, 444 Pluchea [Asteraceae] 69, 237, 240 plum 366 plumeless thistle 473 Poa [Gramineae] 396 Pomoideae 359 Ponderosa tomato 118 Porophyllum gracile [Asteraceae] 442 Pouteria [Sapotaceae] 97, 102, 107, 108, 111 Pouteria campechiana 105 Prenanthes [Asteraceae] 229 prostrata, Eclipta 435 Prunoideae 359 Prunus [Rosaceae] 97, 100, 102, 107, 108, 111, 115, 328, 341, 348, 349, 360 Prunus emarginata 348, 349 Prunus pennsylvanica 348 Prunus persica 343, 413 Prunus serotina 341 Pseudanamomis [Myrtaceae] 111 pseudaureus, Senecio 409 Psidium [Myrtaceae] 97, 100, 102, 107, 108, 111, 115 Psidium guajava 103 ptarmica, Achillea 394 pungens, Hemizonia 240 pungens, Xanthium 147 Punica [Punicaceae] 97, 100, 111 pusillus, Erigeron 310 Pyracantha [Rosaceae] 360 Pyrus [Rosaceae] 97, 100, 102, 107, 111, 360 quadrangularis, Passiflora 98 racemosa, Cornus 346 Rafinesquia [Asteraceae] 444 ragweed 142 Raillardella [Asteraceae] 439 ramissimum, Gnaphalium 433 Ratibida [Asteraceae] 187 Rheedia [Guttiferae] 111 Ribes [Saxifragaceae] 22, 140, 141, 333, 361 Ricinus [Euphorbiaceae] 115 riparia, Ageratina 304 riparium, Eupatorium 485 Rosa [Rosaceae] 324, 328, 330, 336, 496 Rosaceae 70, 325, 326, 350, 359, 365 rose apple 102 rubiginosus, Sideranthus 238

570

Index of Plant Names

Rubus [Rosaceae] 97, 111 Rudbeckia laciniata [Asteraceae] 380 rugosa, Solidago 170, 177 rugosum, Eupatorium 383, 386 rust, wheat stem 352 rydbergi, Arnica 288 salicifolia, Baccharis 398 salsola, Hymenoclea 474 Santalum [Santalaceae] 115 Sapotaceae 107 Sargentia [Rutaceae] 100 sarmentosum, Solanum 118 sarothrae, Gutierrezia 74, 466, 474 sarothroides, Baccharis 79, 397 sativus, Cucumis 133 Saxifragaceae 326 Schoepfia chrysophylloides [Olacaceae] 97 Schoepfia schreberi 97 schreberi, Schoepfia 97 scolymus, Cynara 266 sempervirens, Solidago 172 Senecio [Asteraceae] 78, 187, 291, 399, 439, 442, 444, 452 Senecio aureus 373 Senecio crassulus 409 Senecio douglasii 409 Senecio douglasii var. douglasii 285 Senecio douglasii var. monoensis 285 Senecio integerrimus 409 Senecio integerrimus var. exaltatus 285 Senecio multilobatus 371 Senecio pseudaureus 409 Senecio triangularis 298, 409 Senecioneae 28, 275, 388 sergiloides, Baccharis 397 Sericocarpus [Asteraceae] 229 serotina, Prunus 341 serotina, Solidago 177 serotina var. salebrosa, Solidago 177, 178 sessiflora, Heterotheca villosa var. 311 Severinia [Rutaceae] 111 shockleyi, Acamptopappus 475 Sicyos [Cucurbitaceae] 116 Sideranthus [Asteraceae] 237 Sideranthus rubiginosus 238 Sitilias [Asteraceae] 428 Smallanthus uvedalia [Asteraceae] 383, 386 Solanaceae 326, 333, 361, 494 Solanum [Solanaceae] 97, 108, 248, 252, 326, 333, 494, 496 Solanum eleagnifolium 497 Solanum incanum 118 Solanum indicum 118 Solanum melanocerasum 496 Solanum sarmentosum 118 Solanum verbascifolium 118 Solidagininae 26, 458

Solidago [Asteraceae] 29, 76, 172, 174, 178, 309, 314, 317, 410, 428, 442, 448 Solidago altissima 177, 198 Solidago californica 307 Solidago canadensis 166 Solidago canadensis var. hargeri 177 Solidago fistulosa 173 Solidago gigantea 177 Solidago graminifolia 177 Solidago juncea 170, 172 Solidago missouriensis 172 Solidago rugosa 170, 177 Solidago sempervirens 172 Solidago serotina 177 Solidago serotina var. salebrosa 177, 178 Solidago ulmifolia 177 solstitialis, Centaurea 470, 473 Sonchus [Asteraceae] 401 Sonchus arvensis 401 Sorbus [Rosaceae] 360 Sorocea [Moraceae] 206 Sorocea affims 206 sororia, Arnica 282 sow thistle 401 sphaerocephalus, Acamptopappus 241, 475 spiniflora, Bumelia 104 spinosum, Xanthium 152 spinosus, Aster 242 Spondias [Anacardiaceae] 97, 100, 102, 107, 108, 111 squarrosa, Bidens 486 squarrosa, Centaurea 471 squarrosa var. nuda, Grindelia 469 squarrosus ssp. grindelioides, Haplopappus 243, 468 star thistle 473 star thistle, yellow 470 stem rust, wheat 352 Stephanomeria [Asteraceae] 439 Stephanomeria virgata 229, 296 strawberry 119 strumarium, Xanthium 147, 152 subaxillaris, Heterotheca 310 sunflower 262, 316, 374 Symphoricarpos [Caprifoliaceae] 330, 360, 365 Synsepalum [Sapotaceae] 111 Syzygium [Myrtaceae] 97, 100, 102, 108, 111, 115 Tabernaemontana [Apocynaceae] 416 Tagetes [Asteraceae] 136, 446 talayote 416 Tanacetum [Asteraceae] 396, 400 tea olive, wild 354 tephrodes, Helianthus niveus ssp. 43 teretifolius, Chrysothamnus 475 Terminalia [Combretaceae] 97, 108, 111 Tessaria [Asteraceae] 69

571

Index of Plant Names

Tetradymia [Asteraceae] 439, 442 Theobroma [Sterculiaceae] 97 thistle 262, 458 thistle, bull 474 thistle, Canada 465 thistle, nodding 473 thistle, plumeless 473 thistle, sow 401 thistle, star 473 thistle, yellow star 470 tomato 116, 118, 496 tomato, husk 328, 361 tomato, Ponderosa 118 tortifolia, Machaeracantha 233 Trevesia [Araliaceae] 111 triangularis, Senecio 298, 409 trichophylla, Heterotheca 302 Trichoptilium [Asteraceae] 428 tridentata, Artemisia 192, 199, 202, 261 tridentata var. vaseyana, Artemisia 261 trifida, Ambrosia 152, 385 Trilisa [Asteraceae] 229 Triphasia [Rutaceae] 111 Trixis californica [Asteraceae] 413, 428, 433 Turpinia [Staphyleaceae] 97

Verbesininae 247

ulmifolia, Solidago 177 uvedalia, Smallanthus 383, 386

Xanthium [Asteraceae] 33, 142, 160 Xanthium pungens 147 Xanthium spinosum 152 Xanthium strumarium 147, 152 Ximenia [Olacaceae] 97

Vaccinium [Ericaceae] 332, 354, 363 vaseyana, Artemisia tridentata var. 261 Venegasia [Asteraceae] 291 venetus, Haplopappus 468 venetus var. vernonioides, Haplopappus 235 verbascifolium, Solanum 118 Verbenaceae 24, 30, 33, 180, 206 Verbesina occidentalis [Asteraceae] 387

Vernoma [Asteraceae] 187, 188, 223, 231, 232, 237, 241, 414 Vernonia Baldwini 231 Vernonia interior 231, 241 Vernonieae 27, 223, 411 Viguiera [Asteraceae] 314, 436, 442 Viguiera laciniata 317 villosa, Chrysopsis 242 villosa, Heterotheca 311, 318 villosa var. sessiflora, Heterotheca 311 viminea, Baccharis 161, 398 virgata, Stephanomeria 229, 296 virginicus, Chionanthus 340 viscidiflorus, Chrysothamnus 77, 78, 468, 475 Vitis [Vitaceae] 97 vulgare, Cirsium 474 vulgaris, Berberis 328, 352 walnut 338, 339, 363 Wedelia [Asteraceae] 82, 137, 435 wheat stem rust 352 wild pin cherry 349 wild tea olive 354 Wyethia [Asteraceae] 439

yellow star thistle 470 zapota, Manilkara 100, 103 Zizyphus [Rhamnaceae] 115 Zuelania guidonia [Flacourtiaceae] 111