Neotropical Ceratopogonidae, Diptera, Insecta (Aquatic Biodiversity in Latin America) 9546423017, 9789546423016, 9781435630352

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ABLA ol. 4

Neotropical Ceratopogonidae (Diptera: Insecta)

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

Aquatic Biodiversity in Latin America Biodiversidad Acuática en América Latina

Volume 4

Neotropical Ceratopogonidae (Diptera: Insecta) Volumen 4

Neotropical Ceratopogonidae (Diptera: Insecta)

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Volumen 4

CERATOPOGONIDAE NEOTROPICALES (DIPTERA: INSECTA) por

Art Borkent & Gustavo R. Spinelli Lector Científico a los Editores: William L. Grogan Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2007

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

AQUATIC BIODIVERSITY IN LATIN AMERICA Volume 4

NEOTROPICAL CERATOPOGONIDAE (DIPTERA: INSECTA) by

Art Borkent & Gustavo R. Spinelli Scientific Reader to the Editors: William L. Grogan Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2007

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

AQUATIC BIODIVERSITY BIODIVERSIDAD ACUÁTICA

IN

LATIN AMERICA

EN

AMÉRICA LATINA

Series Editors/Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen VOLUMEN 4. CERATOPOGONIDAE NEOTROPICALES (DIPTERA: INSECTA) por Art Borkent & Gustavo R. Spinelli Lector Científico a los Editores: William L. Grogan

VOLUME 4. NEOTROPICAL CERATOPOGONIDAE (DIPTERA: INSECTA) by Art Borkent & Gustavo R. Spinelli Scientific Reader to the Editors: William L. Grogan Front cover:

Female Forcipomyia (Microhelea) sp. feeding on phasmid antenna (Chlorosphasma sp.) in Costa Rica; Amazon River near Manaus (photos: P. Naskrecki, WJ. Junk; design: Zheko Aleksiev & Elke Bustorf). Aquatic Biodiversity in Latin America Vol. 4 ISSN: 1312-7276 First published 2007 ISBN-13: 978-954-642-301-6

The book should be cited as follows: Art Borkent & Gustavo R. Spinelli. 2007. Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.): Aquatic Biodiversity in Latin America (ABLA). Vol. 4. Pensoft, Sofia-Moscow, 198 pp.

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria [email protected] www.pensoft.net Printed in Bulgaria, June 2007

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

Habitus of female Culicoides variipennis COQUILLETT (Nearctic Region; from McALPINE et al., 1981).

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

AQUATIC BIODIVERSITY IN LATIN AMERICA BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen Biodiversity is a key word in science and global management schemes; however very few people are able to identify the species and their ecology that make up “biodiversity”. For many years, researchers and students from numerous countries complain about the lack of “tools” to identify aquatic invertebrates from Latin America. Keys found in accepted entomological textbooks are mostly highly limited, superficial and rarely cover Neotropical biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology is scattered throughout the literature in many single publications. An international team of editors have combined their efforts with Pensoft Publishers to launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Their goal was to find experts who combine the current state of knowledge in taxonomy and ecology, in order to produce a concise and affordable handbook for each group. About 15 separate monographs, written by reference scientists from various countries will offer a new, unrivalled view on the aquatic fauna of South America. Information on the ecology and status of the taxa (written in English) is combined with illustrated identification keys to families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists, hydrobiologists, biogeographers, conservationists and students interested in aquatic biodiversity. The series will be an essential tool for any biological library. Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and photos (15 plates in color); publication date: March 2006. Volume 2: Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-259-2, ISBN-13: 978-954-642259-0, 165x240 mm, 646 pp., including 234 plates of figures in line drawings and photos (16 plates in color); publication date: May 2006.

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Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarón & Cecilia L. Coscarón Arias. Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-293-4, 165x240 mm, 685 pp., including 135 plates of figures in line drawings and 19 maps of distribution; publication date: March 2007. Volume 4: Neotropical Ceratopogonidae (Diptera: Insecta) by Art Borkent & Gustavo R. Spinelli. Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-301-6, 165x240 mm, 198 pp., including 26 plates of figures in line drawings and photos; publication date: June 2007. Ceratopogonidae are found in virtually every habitat with even a small amount of sustained moisture, are notorious as severe pests of man and domestic animals, and are one of the most common fly families in traps. Yet they are by far the most poorly understood of all the biting flies. In part, this is due to their remarkably diversity, both in numbers of species and in their various adaptations. Larvae range from being detritivores and scavengers to predators and adult females of various species feed on a wide array of vertebrates and invertebrates. Some females draw blood from conspecific males while they are mating. Adults are also important pollinators of a wide array of plants. This book brings together, for the first time, all known information on the 51 genera and 1095 described Neotropical species of biting midges. An overview of the group includes a discussion of bionomics, adaptations of the various life stages, habitats, their ecological roles (including disease transmission), collecting and preparation techniques, zoogeography and classification. A key to adults (including males) provides the first key to all the genera of this region. Limited keys to eggs and larvae are presented. Keys are written in both English and Spanish. A table of all previous descriptions of immatures and a catalog of all Neotropical species is included. This book will be of interest to systematists, ecologists, medical entomologists and students wishing to undertake innovative research in a remarkably interesting and instructive group of insects.

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

ADDRESSES Authors: DR. ART BORKENT 691-8th Avenue SE Salmon Arm, British Columbia V1E 2C2 Canada e-mail: [email protected] DR. GUSTAVO R. SPINELLI Divisíon Entomologia Museo de La Plata Paseo Del Bosque s/n 1900 La Plata, Argentina e-mail: [email protected] Series Editors: PROF. DR. JOACHIM ADIS Max-Planck-Institute for Evolutionary Biology Tropical Ecology Working Group Postfach 165 24302 Plön, Germany e-mail: [email protected] DR. JORGE R. ARIAS 5870 Colfax Avenue Alexandria, Virginia 22311, USA e-mail: [email protected]

M.Sc. GUILLERMO RUEDA-DELGADO Grupo de Investigación en Cuencas y Humedales Tropicales GICHT UNIMAGADALENA Universidad de Bogotá Jorge Tadeo Lozano Laboratorio de Limnología Carrera 4 No. 22-61 Bogotá, D.C., Colombia e-mail: [email protected] DR. KARL MATTHIAS WANTZEN Institute of Limnology University of Konstanz Postfach M 659 78457 Konstanz, Germany e-mail: [email protected] Scientific Reader to the Editors: DR. WILLIAM L. GROGAN Department of Biological Sciences Salisbury University Salisbury, Maryland 21801-6837 e-mail: [email protected]

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FOREWORD Neotropical Ceratopogonidae (Diptera: Insecta) by Art Borkent & Gustavo R. Spinelli The Ceratopogonidae are a large and diverse family of small, mostly crepuscular flies commonly known as biting midges. Though biting midge species are known as vicious biters of humans and domestic animals in many areas, some of the most vivid descriptions of these encounters have come from the Neotropical region. Even the intrepid Victorian naturalist Alfred Russell Wallace, writing near the headwaters of the Rio Negro on the Brazil-Venezuelan border, found them to be intolerable: “I was…exposed to the pest of the sand-flies, which, every afternoon, from four to six, swarm in millions, causing by their bites on the face, ears and hands, the most painful irritation.” The sheer numbers of biting midges in many areas continue to circumvent the advance of agriculture and tourism in the Neotropics. In addition, some ceratopogonid species are known to vector important viruses (e.g., Oropouche) and filarial nematodes (e.g., Dipetalonema spp.) to humans. Though it comes as little consolation to those who have experienced an onslaught by biting midges, most species in the Neotropics feed exclusively on the blood of other insects. Nevertheless, to their credit, some ceratopogonids (species in the genus Forcipomyia) serve as an important cacao pollinator, making a direct contribution to the enjoyment of chocolate worldwide. Although there have been many taxonomic papers published on Neotropical Ceratopogonidae from the nineteenth century to the present, this volume represents the first truly comprehensive work for the region because it includes all described taxa. The significance of this work can not be overstated since approximately 20 percent of the World’s described ceratopogonid species are treated in its 198 pages. Both authors have collected extensively in Latin America and have authored or coauthored taxonomic and ecological papers about ceratopogonid species from the region, many of which included descriptions of new species. I have had the distinct pleasure of meeting Art Borkent and Gustavo Spinelli. Both are professional systematists with an infectious enthusiasm for the study of ceratopogonids. Both have extensive contacts in many countries of Central and South America and have trained parataxonomists to assist them (e.g., through the National Biodiversity Institute (INBio) in Costa Rica). Both are advocates of continued biodiversity studies in the Neotropics towards the conservation and sustainable use of the numerous species which inhabit the region. In addition to their comprehensive, well-illustrated keys to the adults, eggs and larvae of Neotropical genera, the authors have included important chapters on the bionomics of

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species in this region, methods of collection and rearing ceratopogonids, phylogenetic relationships among taxa including references to the fossil record and a preliminary discussion of the zoogeography of selected species. A synopsis of the medical importance of ceratopogonids in Latin America is also included as are useful syntheses of information about Neotropical ceratopogonids (e.g., Table 1 includes references for all species whose immature stages have been described). The color photographs in the bionomics chapter of biting midges feeding on other insect species are particularly striking. Another useful feature found in the Catalog of the Ceratopogonidae of the Neotropical Region is the designation of species in the genera Culicoides and Leptoconops which are known to bite humans. Borkent and Spinelli acknowledge more than once that many more ceratopogonid species will be described from Latin America following expeditions to poorly collected areas. Their foundational work will most certainly facilitate future taxonomic work. The authors have provided an invaluable resource for scientists who investigate aquatic biodiversity in the Neotropics.

STEVE MURPHREE Editor, Ceratopogonidae Information Exchange Belmont University Nashville, Tennessee, USA

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PREFACE The Ceratopogonidae are commonly known as biting midges, no-see-ums or punkies, in Spanish as manta blanca, polvorines, or jejenes (which also may refer to Simuliidae), and in Portuguese as maruim or mosquito pólvora. They have a bad reputation as being nasty biters that pester humans and domestic animals and, in some instances, transmit harmful diseases. Because of their small size, the females of some species can pass through screen and mesh that keeps other biting pests outside and these can make life insufferable. Biting midges may occur in such huge numbers that in some areas people are driven indoors (or complain loudly and then suffer numerous itchy bites!). Few people, however, realize that this group of flies provides important services in a wide array of ecosystems. Some species are important pollinators of such plants as cacao (without them we wouldn’t enjoy chocolate!) and rubber trees, and the larvae of many are important predators of other organisms in semiaquatic and aquatic habitats. The adults of most biting midges actually suck blood from other insects and may even be important vectors of viruses that kill caterpillars. The biting midges are a diverse group in both numbers of species and in their habits and there are some intriguing stories associated with the way they obtain their food, mate and the kinds of different habitats they occupy. This contribution provides the basis for understanding the family in the Neotropical Region, allows for their identification to the generic level, gives a synopsis of each genus, details some aspects of their behaviour and ecology, discusses various collecting and preparation techniques, and provides a catalog of the Neotropical species. The first author thanks his wife and dearest friend Annette BORKENT for providing moral and financial support throughout the writing of this contribution and for her steadfast faith in his work. Pablo DELLAPÉ and Maria M. RONDEROS provide substantial advice and assistance in preparing the plates and we express our gratitude to them. A number of the figures were skillfully produced by Ana BRENES (INBio) and we appreciate permission from the Instituto Nacional de Biodiversidad in Costa Rica to reproduce these here. Nelida CALIGARIS redrew some of the illustrations of previously published figures. The Series Editors and the Scientific Reader are thanked for valuable suggestions, improving the content and the final shape of the book. Many of the figures reproduced here are from Chapter 28 of Volume 1 of the Manual of Nearctic Diptera (MCALPINE et al. (eds), Agriculture Canada Monograph 27, Agriculture & Agri-Food Canada, 1981) and we are grateful for permission to do so from the Minister of Public Works and Government Services Canada (2006). Annia PICADO (INBio) has slide mounted tens of thousands of Ceratopogonidae from Costa Rica with tremendous skill and this has helped our understanding of Neotropical biting midges immeasurably. Heron HUERTA provided a careful review of this work and we thank him for his detailed critique.

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

We gratefully acknowledge permission to use photographs from several colleagues who are noted in the figure captions. THE AUTHORS

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

Table of Contents 1 B References to species of Neotropical Ceratopogonidae IONOMICS

described as eggs, larvae and/or pupae (Table 1) Number of named species with described immatures for genera of Neotropical Ceratopogonidae (Table 2) Organisms transmitted by Neotropical Ceratopogonidae (Table 3)

2C 3R 4P

APTURE AND STUDY OF

EARING

CERATOPOGONIDAE

CERATOPOGONIDAE

15 25 32 33 34 38

HYLOGENETIC RELATIONSHIPS AND FOSSIL

RECORD OF

CERATOPOGONIDAE

5Z 6 C Catalog of the Ceratopogonidae of the Neotropical OOGEOGRAPHY

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LASSIFICATION

42

Region (Table 4)

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39

DENTIFICATION

Numbers of named valid species in each genus of Neotropical Ceratopogonidae (Table 5) Diagnosis of the Ceratopogonidae

43 100 101 102

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Key to the genera of adult Ceratopogonidae in the Neotropical Region Key to the genera of eggs of Ceratopogonidae in the Neotropical Region Key to the genera of larvae of Ceratopogonidae in the Neotropical Region Clave para los géneros de adultos de Ceratopogonidae de la Región Neotropical Clave para los huevos de géneros de Ceratopogonidae de la Región Neotropical Clave para las larvas de géneros de Ceratopogonidae de la Región Neotropical

8 SN

YNOPSIS OF THE EOTROPICAL

9R 10 S

UBJECT INDEX

129 129 134 146 146

CERATOPOGONIDAE OF THE

REGION

EFERENCES

103

148 154 182

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1 BIONOMICS Ceratopogonidae are one of the common of fly families in the Neotropical Region (BROWN, 2005). Adults may be found from the coast (some are intertidal) to the highest elevations. We have records of Forcipomyia MEIGEN and Dasyhelea KIEFFER from 3700 to 4000 meters in northwestern Argentina and Forcipomyia and Atrichopogon KIEFFER from 3819 meters in Costa Rica (which is that country’s highest point). They are present in nearly all habitats where there is even a little long-lasting moisture. Adults of a few species are vicious biters of humans and domestic animals and some act as vectors of various diseases or parasites (BORKENT, 2004). There is little information on the specific life cycle of Ceratopogonidae in the Neotropical Region. Further to this, our survey of the literature (Tables 1, 2) indicates that fewer than 15 percent of the species named from this area are described in even one immature stage. Nineteen genera are entirely unknown. Some descriptions, especially those that are older, are likely based on misidentified species and these immatures need to be restudied. Aware that many species remain to be described even as adults, there is a rich field available for investigation. Undoubtedly, many of the larvae of Ceratopogonidae play an important ecological role that is nearly entirely uninvestigated. Rearings of larvae will certainly provide important information regarding the structure of tropical ecosystems but clearly, the first step is to have these described and to have keys made available. At the present time there are no reliable keys to the larvae or pupae to even the generic level although these stages are presently being studied at a worldwide level (BORKENT, in prep.). All members of the family have an egg, four larval instars, a pupal and an adult stage. Eggs may be laid on moist substrate or, in the case of fully aquatic species, in water. The fact that adult ceratopogonids, as a group, are present throughout the year in tropical areas suggests that many species have multiple life cycles, without the third or fourth instar experiencing a diapause as do nearly all species in northern temperate and arctic regions. In some tropical areas, it appears that adult Ceratopogonidae may be somewhat more diverse during the rainy season. It is unknown how tropical Ceratopogonidae respond in areas with a pronounced dry season. The life cycle of Ceratopogonidae in the southern temperate zone of South America is entirely unstudied but the presence of both numbers of species and adults in late spring in this area indicates that many species likely overwinter in the late larval stage, as they do in the northern temperate region. Larvae are important detritivores or predators in semiaquatic and aquatic systems of all sizes, from tiny tree holes and the water held by tropical flowers, to large rivers and lakes, where they may be an important source of food for other invertebrates and fish. In tropical areas of Latin America, the immatures of Ceratopogonidae form an important compo-

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nent of arboreal aquatic systems (such as in bromeliads and tree holes). The most diverse group, both in terms of numbers of taxa and structurally, are those in the subfamily Ceratopogoninae. Members of this group have predaceous larvae which are like small snakes, working their way slowly through the wet substrate and, depending on the instar and/or taxon, feeding on microorganisms or lashing out at prey. Some prey are swallowed whole (e.g. nematodes) but in the case of larger prey, which are mostly other insect larvae, the biting midge larvae of some taxa burrow through the cuticle of the host and eat their insides. It seems certain that in many habitats, biting midge larvae play an important role in the biological control of other insects, but this has not been studied in much detail. The immatures of each of the four subfamilies differ substantially. The Leptoconopinae includes just one genus, Leptoconops SKUSE in the Neotropical Region. The group is an ancient one and fossil Leptoconops are known from Lebanese amber, 121 million years old. Their larvae live in wet sand (especially on marine beaches) or in cracked alkaline, clay soils. The sluggish larvae burrow through wet sand or loose clay soil and feed on microorganisms by raking their heads over the substrate. Larvae of Forcipomyiinae are aquatic, semiaquatic or terrestrial. When they are in aquatic habitats, they are generally restricted to lentic (standing) waters although a few are known from streams and small rivers and are associated with mosses and algae. They retain the primitive anterior and posterior prolegs and therefore can walk on surfaces beneath or above water. As they move, they graze the substrate surface for microorganisms, looking somewhat like cattle eating grass. Some species feed on decaying vegetation and a few species may be found associated with manure. The larvae of most species have specialized setae along the length of their bodies that secrete a sticky fluid that repels ants. When the larvae pupate, most retain the larval exoskeleton on the posterior end of their bodies. Larvae of Dasyheleinae are aquatic and semiaquatic and tend to be sluggish. They have posterior hooks on their abdomens and move slowly over the substrate in which they are found. Most species live in small water bodies such as those in epiphytes, tree holes, broken or damaged bamboo, rotten banana stems, fruit husks and small rock pools on the margins of streams and rivers. A few species are common in mangrove swamps and salt marshes. Larvae may also be found in wet vegetation. One species in Costa Rica is a leafminer of the floating leaves of Salvinia and another is found in the wet refuse piles of some ants (BORKENT, pers. obs.). Larvae feed on detritus, algae, and, in a few species, on carrion (dead insects). There are some species whose adults are associated with rocky areas in the upper tidal zone of coastal Costa Rica but the immatures of these are not known. Larvae of the Ceratopogoninae, the most species-rich group of biting midges, are prognathous, lack prolegs and hooks (at least in instars 2-4), and their abdominal segments are not secondarily divided. Larvae of most species swim in a characteristic, very rapid, serpentine motion through the water column. When on or in substrate they assume a slower motion, working their way through wet substrate, eating microorganisms or lashing out at larger prey, looking very much like small snakes. Some smaller invertebrates are swallowed whole but in the case of larger prey, which are mostly other insect larvae, the biting midge larva penetrates the cuticle of the prey and devours the internal contents. The earlier phylogenetic lineages tend to feed on microorganisms and/or smaller invertebrates, often ingesting these whole. The more derived lineages, especially in the tribes Heteromyiini, Sphaeromiini, and Palpomyiini,

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feed on larger invertebrates such as chironomid larvae and sometimes burrow into the body of their host. In many habitats, biting midge larvae probably exert an important biological control on other insects. Larvae of the species-rich genus Culicoides LATREILLE may be found in decaying vegetation, fruits, and manure, epiphytes, tree holes, wet mud, mangrove swamps and a large array of other aquatic habitats. In streams and rivers, Ceratopogoninae larvae may be an important source of food for other invertebrates and fish. In general, the most primitive lineages of the subfamily are in smaller habitats, whereas those in the Heteromyiini, Sphaeromiini and Palpomyiini may be found in larger bodies of water: ponds, springs, and larger lotic and lentic habitats such as rivers and lakes. Pupae are generally sluggish and move their abdomens slowly in more or less circular movements until they find an appropriate resting position in the midst of the substrate or at the water surface where their paired respiratory organs can obtain air. The pupae of some Sphaeromiini move above the water line and have specialized membranous pads on the venter of the abdomens which they use to stick to the dry substrate. BORKENT & CRAIG (2001) and CAZORLA & MARINO (2004) described unusual pupae of Stilobezzia KIEFFER from Costa Rica and Peru respectively, which use their respiratory organs to obtain oxygen by piercing the submerged leaves of aquatic plants. Adult females of Leptoconops, Culicoides and Forcipomyia (only in the subgenus Lasiohelea KIEFFER) are haematophagous on vertebrates, needing the blood to develop their eggs. Female Leptoconops and Culicoides (Fig. 1B, frontispiece) may occur in large numbers in places where there is appropriate and abundant habitat (Table 4). Their bite (sometimes unnoticed at the time) can produce burning and itching welts completely out of proportion to their small size. In some areas of the Neotropical Region (northern South America, Central America and tropical Mexico) species of Leptoconops are generally restricted to marine beaches and adult females are daytime biters, generally on the lower legs of humans. The adults of some species (including L. bequaerti (KIEFFER) in Central America) have an interesting behaviour of resting under a thin layer of sand (LINLEY, 1968). Adult Culicoides occur, in varying numbers, in virtually every region of the Neotropical Region, from Mexico to southern Chile (Puerto Natales, at almost 52º S), from the coast to high elevations (at least 3320 m in the Colombian Andes (WIRTH & LEE, 1967)). Although the females of a few species may bite during the day and may occur in huge numbers, most are crepuscular or nocturnal feeders. Of the 266 species of Culicoides known from the Neotropical Region, 70 (26%) have been recorded feeding on humans (Table 4) but of these only the following occur in sufficient numbers to be considered serious pests of humans: Culicoides furens (POEY) and C. phlebotomus (WILLISTON) in coastal Central and northern South America, C. maruim LUTZ in coastal Brazil, C. insinuatus ORTÍZ & LEON, C. puracensis WIRTH & LEE at high elevations in the Andes of Colombia, C. pseudodiabolicus FOX and C. paraensis (GOELDI) in the Amazon basin, C. lahillei (ICHES) and C. debilipalpis LUTZ in Paraguay and northeastern Argentina, C. caridei (BRÈTHES) in Uruguay and the province of Buenos Aires, Argentina, and C. patagoniensis RONDEROS & SPINELLI in Argentinean and Chilean Patagonia. Further pestiferous species will likely be discovered in the largely unknown Andes. Females of some other species feed on domestic animals. For example, females of C. debilipalpis feed on both humans and horses and C. arubae FOX & HOFFMAN, C. foxi ORTÍZ and C. barbosai WIRTH & BLANTON have been collected from mules or

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horses. Culicoides insignis LUTZ is typically associated with cattle. Culicoides phlebotomus, a well known species in the Neotropics biting humans and dogs on marine beaches, is also known to suck blood from leatherback turtles that are on the beach laying their eggs (BORKENT, 1996). The hosts of the remaining species of Culicoides are largely unknown. The biting habits of female adult Forcipomyiinae, including only two genera, Forcipomyia and Atrichopogon, are remarkably diverse. In general, female adults act as ectoparasites of large insects, with some sucking blood from their hosts for extended periods of time. In particular, those Forcipomyia in the ixodoides species group of the subgenus Microhelea KIEFFER attack phasmids (Fig. 2B) and katydids (Figs. 2A, 4A) and when replete and developing their eggs, look like small ticks (CLASTRIER & WIRTH, 1995). One female of this species group we examined in Costa Rica had developed 2778 eggs. Other species of Forcipomyia suck hemolymph

Culicoides furens Det. G.R. Spinelli

‡

2 km N. Montezeuma, Playa Grande, Costa Rica, 21-XI-2000, A. Borkent Biting humans CD5099

A

B Figure 1. A: Dissected adult Ceratopogonidae on a microscope slide. B: Female Culicoides sp., biting human (taken by S.A. Marshall in South Carolina, USA).

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A

B Figure 2. Female Forcipomyia (Microhelea) spp., feeding. A: On katydid abdomen (Orophus sp.) (taken by M. Zumbado in Costa Rica). B: On phasmid antenna (Chlorophasma sp.) (taken by P. Naskrecki in Costa Rica).

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ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales

from caterpillars (Fig. 3B), sawfly larvae (WIRTH, 1972), spiders, crane flies, the wing veins of dragonflies and damselflies (Fig. 3A), lacewings and butterflies (WIRTH, 1956a; WIRTH & MESSERSMITH, 1971; MARINO & VON ELLENRIEDER, 1999). The feeding habits of female Atrichopogon are more poorly known but those of at least one subgenus feed on the blood of adult Meloidae beetles (BORKENT & ROCHA-FILHO, 2006). Species in the subgenus Forcipomyia (Lasiohelea) are the only Forcipomyiinae that are vertebrate feeders. Their hosts are poorly known and females have been recorded feeding only on frogs and lizards in the New World (SPINELLI et al., 2002). The males and females of many adult Forcipomyia and Atrichopogon feed on nectar to fuel their flight and are often common on flowers, particularly those that are small and white or green. One unusual example of consuming sugars is Forcipomyia adults from the backs of pentastomid bugs in Costa Rica (Fig. 4B).

A

B Figure 3. A: Female Forcipomyia (Pterobosca) sp. feeding on damselfly wing (Argia hinei) (taken by D. Huntington in New Mexico, USA). B: Female Forcipomyia (Microhelea) sp. feeding on caterpillar (Automeris zugana) (taken by D.H. Janzen in Costa Rica).

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

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A

B Figure 4. A: Female Forcipomyia (Microhelea) spp., feeding on katydid (Pristonotus latistylus) (taken by P. Naskrecki in Costa Rica). B: Forcipomyia sp. females likely feeding on surface sugars on a pentatomid (taken by M. Moraga in Costa Rica).

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Adult females of Dasyhelea have reduced mouthparts and do not suck blood from either vertebrates or invertebrates. Some species are common at flowers where they, like many other biting midges, obtain nectar from flowers. Adult females in the subfamily Ceratopogoninae, other than those of Culicoides, are predators and feed on other flying insects that are approximately the same size as themselves. In general, a female flies into a swarm of male non-biting midges (Chironomidae), grabs a male, and either settles to the ground or surrounding vegetation. The female injects an enzyme that dissolves the internal organs and tissues of the prey, sucking up the liquefied contents as food. In some members of the Heteromyiini, Sphaeromiini and Palpomyiini the female enters a male swarm of her own species and, while mating occurs, pierces the body of the male and sucks out its contents (DOWNES, 1978). Such females may later be found with the dried male genitalia still attached to their own abdomen, after the bulk of the dried male has broken off. The males of these species contribute not only their sperm and body tissues but also an effective mating plug to the female with which they mate. BORKENT (2004) provided a list of all diseases and parasites known to be transmitted by biting midges worldwide. The females of Leptoconops, Forcipomyia (Lasiohelea) and Culicoides are the only genera which include species which bite vertebrates (from frogs to birds to mammals) in the Neotropical Region. Worldwide, Ceratopogonidae are known to transmit 66 viruses, 15 species of protozoa and 26 species of filarial nematodes. It is highly likely that many more remain undetected. This strongly suggests that Ceratopogonidae will be found to transmit many more organisms in the Neotropical Region than the six viruses and eight nematodes currently known (Table 3). For example, it would be quite remarkable if there were no protozoan parasites of birds in the entire Neotropical Region when at least nine species of protozoans are known from North America. The adults of those biting midges that suck hemolymph from large insects (some Atrichopogon and Forcipomyia) suggest the possibility that they may be involved in transmitting viruses from one insect host to another, particularly among caterpillars. Bluetongue is a viral disease of cattle, sheep, goats, and wild ruminants. It is particularly damaging in sheep, where up to half of infected animals may die. In cattle and goats the disease primarily affects the animal’s reproductive capabilities. Although it was known that the virus occurred in Central America (MO et al., 1994) and antibodies were detected in cattle in the late 1980s in southern Brazil (CUNHA, 1990), there were no published data of animal illness in the Neotropics until recently. The Office International des Epizooties, World Health Organization (OIE, 2001) reported the presence of the disease in the state of Paraná, Brazil, in which eight sheep and one goat showed classical symptoms, including the death of four of these animals, and 70 cattle showed susceptibility as well. More recently, the virus was isolated and characterized from asymptomatic bovines from northeastern Argentina (GORCH et al., 2002). Bluetongue occurs virtually worldwide and has different, phylogenetically unrelated vectors in different regions of the globe, although all are in the genus Culicoides. In the Neotropical Region the primary vectors are likely C. insignis, C. pusillus LUTZ (WIRTH & DYCE, 1985; SÁENZ & GREINER, 1994; GORCH et al., 2002) and possibly C. filarifer HOFFMAN (SÁENZ & GREINER, 1994). There are other diseases of humans and domestic animals that occur elsewhere in the Neotropical Region. One of the more significant is Oropouche fever, a viral infection pro-

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

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A

B Figure 5. A: Female Palpomyia sp. (taken by S.A. Marshall in Ontario, Canada). B: Female Sphaeromias sp. (taken by K. van der Krieke in the Netherlands).

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ducing severe flu-like symptoms, mostly in Brazil. It has been responsible for at least 27 epidemic outbreaks in humans, including one in the state of Pará, Brazil in 1978-1980 which afflicted at least 130,000 people. The disease is primarily transmitted by species of Culicoides (and particularly by C. paraensis) but has also been isolated (at low levels) from some mosquitoes. Outbreaks occur primarily in urban areas, where populations of C. paraensis may occur at great densities (HOCH et al., 1990; MELLOR et al., 2000). MERCER et al. (2003) reported decaying platano (plantain: Musa parasidiaca) stems, stumps, flowers, fruits and debris beneath platano trees, as well as soil beneath a fruiting mamey (Pouteria sapota) tree and organic-rich mud along a lake shoreline, as larval substrates of C. paraensis in Iquitos, Peru, a city which has suffered several outbreaks of Oropouche fever. LEDUC & PINHEIRO (1989) provide an excellent overview of Oropouche fever. The disease has also been studied in Peru (CHAVES et al., 1992; WATTS et al.,1997; PINHEIRO et al.,1998) and Panama (PINHEIRO et al.,1998). Ceratopogonids provide some important services in ecosystems, although often thought of in negative terms. The adults of many genera, primarily Forcipomyia and Dasyhelea, are important pollinators of some commercial trees such as cacao (Theobroma cacao) and rubber (Hevea brasiliensis). Species of these ceratopogonid genera, and those of Atrichopogon, Culicoides, and Stilobezzia, are likely to be pollinators for a number of other plants in tropical America (possibly including mango (Mangifera indica) and a number of palms). Further information about pollination by ceratopogonids may be found in a series of papers by WARMKE (1951, 1952), SAUNDERS (1959), SAUNDERS & Bowman (1956), WINDER (1977a, b), WIRTH (1956c) and YOUNG (1982, 1983, 1986a, b). Although it is unpleasant for many, the vicious attacks on humans and domestic animals by the females of some species of Culicoides serve to restrict human encroachment in some important and threatened habitats, helping to preserve these for the future. Ceratopogonidae larvae, pupae and adults harbour a variety of their own parasites and diseases, including viruses, bacteria, fungi, protozoa, nematodes, mites, and even parasitic wasps (WIRTH, 1977; FRANA et al., 2001). Although nearly all reports are from elsewhere, VILLALOBOS & RONDEROS (2003) recently experimentally infected immatures stages of Dasyhelea necrophila SPINELLI & RODRÍGUEZ from Argentina with larvae of Nematomorpha, causing a high level of parasitization and death of the host.

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Table 1. References to species of Neotropical Ceratopogonidae described as eggs, larvae and/or pupae. ‘E’ refers to egg; ‘L’ refers to larva and ‘P’ to pupa. Theses are cited only when their results have not been published. Duplicated figures or repeated descriptions by subsequent authors are not listed here. Some descriptions, especially those that are older, may be based on incorrectly identified species. Taxon

Reference

Leptoconops bequaerti (as Leptoconops sp.)

PAINTER, 1927 MAYER, 1934 LINLEY, 1965a LINLEY, 1968

Atrichopogon bifidus A. caribbeanus A. fusculus

Stages Described L P P E L

EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958 MALLOCH, 1915a THOMSEN, 1937 NIELSEN, 1951 EWEN & SAUNDERS, 1958 DOWNES & WIRTH, 1981 WIRTH, 1994a

L L L L L L L L

P P P P P P

A. fusculus or rostratus (as Kempia haesitans) A. incultus A. longicornis A. obscurus A. remigatus A. saundersi A. tuberculatus

LENZ, 1934 EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958 EWEN & SAUNDERS, 1958

L L L L L L L

P P P P P P P

Forcipomyia (Caloforcipomyia) caerulea F. (C.) glauca (as monilis) F. (C.) varicolor

SAUNDERS, 1957 KRIVOSHEINA, 1968 SAUNDERS, 1957

L P L P L P

Forcipomyia (Euprojoannisia) blantoni

F. (E.) setigera F. (E.) spatulifera

SORIA & BYSTRAK, 1975 BYSTRAK & WIRTH, 1978 SAUNDERS, 1957 WIRTH, 1952a SAUNDERS, 1957 SAUNDERS, 1957 SAUNDERS, 1957 BYSTRAK & WIRTH, 1978 SAUNDERS, 1957 SAUNDERS, 1957

L L L L L L L L L L

P P P P P P P P P P

F. (Forcipomyia) argenteola F. (F.) genualis (as raleighi) (as raleighi) F. (F.) harpegonata

LANE, 1947c SAUNDERS, 1957 GUTSEVICH & GLUKHOVA, 1970 WIRTH & SORIA, 1975

L L L L

P P P P

(as polydactylus) (as polydactylus) (as polydactylus)

F. (E.) bromeliae F. (E.) calcarata F. (E.) falcifera F. (E.) longispina F. (E.) mortuifolii

P

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Table 1. Continued. Taxon

Reference

F. (F.) pictoni F. (F.) rioplatensis

WIRTH, 1976 SPINELLI et al., 2005a

L P L P

F. (Lasiohelea) attenuata F. (L.) cornuta F. (L.) intermedia F. (L.) quasicornuta F. (L.) stylifer

SAUNDERS, 1964 SAUNDERS, 1964 KRIVOSHEINA, 1968 SAUNDERS, 1964 SAUNDERS, 1964 LANE, 1947c

L L L L L L

P P P

F. (Lepidohelea) brasiliensis F. (L.) edmistoni F. (L.) herediae F. (L.) luteigenua F. (L.) seminole

WIRTH & SPINELLI, 1993a WIRTH & SPINELLI, 1993b WIRTH & SPINELLI, 1993a WIRTH & SPINELLI, 1992a WIRTH & SPINELLI, 1992a

L L L L

P P P P P

F. (Metaforcipomyia) cerifera

SAUNDERS, 1957

L P

F. (Microhelea) fuliginosa (as inornatipennis spp.ornaticrus)

LANE, 1947c

L P

F. (Phytohelea) antiguensis F. (P.) bromelicola (as a Apelma) F. (P.) caribbeana F. (P.) dominicana F. (P.) edwardsi (as a Apelma) F. (P.) jocosa F. (P.) keilini (as a Apelma) F. (P.) magna (as a Apelma) F. (P.) musae F. (P.) oligarthra

SAUNDERS, 1957 SAUNDERS, 1925 SAUNDERS, 1957 DE MEILLON & WIRTH, 1979 SAUNDERS, 1925 SAUNDERS, 1957 SAUNDERS, 1925 SAUNDERS, 1925 CLASTRIER & DELÉCOLLE, 1994 SAUNDERS, 1957 TOKUNAGA, 1961

L L L L L L L L

F. (Thyridomyia) nodosa

SAUNDERS, 1957

L P

F. (Trichohelea) trinidadensis

SAUNDERS, 1964

P

F. (Warmkea) aeria

SAUNDERS, 1957 WIRTH & SORIA, 1980 SAUNDERS, 1957 WIRTH & SORIA, 1980 SAUNDERS, 1957 WIRTH & SORIA, 1980 SAUNDERS, 1964 WIRTH & SORIA, 1980

F. (W.) lesliei (as bicolor) F. (W.) spinosa F. (W.) terrestris

Stages Described

P P

P P P P P P P P P L P L P

L L L L L L L L

P P P P P P P P

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Table 1. Continued. Taxon

Reference

F. (W.) tuberculata

SAUNDERS, 1957 WIRTH & SORIA, 1980 SAUNDERS, 1925 SAUNDERS, 1957 SAUNDERS, 1964 SAUNDERS, 1964

F. sp. (as an Apelma sp.) F. sp. (as species A) F. sp. (as species A) F. sp. (as species B) Dasyhelea atlantis D. bahamensis D. calvescens D. cincta (as penthesileae) D. correntina D. filibranchia (as a Ceratopogon) D. flavicauda D. lacustris D. mediomunda D. mutabilis

D. necrophila

D. paracincta D. paulistana D. pusilla Culicoides annettae C. annuliductus C. arubae (as sp. 1)

C. bambusicola (as sp. 2)

C. barbosai

C. bayano C. bermudensis

WAUGH & WIRTH, 1976 BORKENT, 1991 RONDEROS et al., 2003 WILLIAMS, 1944 SPINELLI, 1983b RONDEROS et al., 2004a LUTZ, 1914 SPINELLI & RONDEROS, 1987 SPINELLI & RONDEROS, 1987 MINAYA, 1978 THOMSEN, 1937 WIRTH, 1952a WAUGH & WIRTH, 1976 SPINELLI & RODRÍGUEZ, 1999 RONDEROS et al., 2003 RONDEROS et al., 2006 BORKENT, 1991 FORATTINI & RABELLO, 1957 FORATTINI & RABELLO, 1957 SPINELLI & BORKENT, 2004a VITALE et al., 1981 FOX, 1942 FORATTINI, 1957 MURPHREE & MULLEN, 1991 LUTZ, 1913 FOX, 1942 LANE, 1947c BARBOSA, 1952, 1953 FORATTINI, 1957 RONDEROS & SPINELLI, 2000 RONDEROS et al., 2000 LINLEY, 1965b LINLEY & DAVIES, 1971 BLANTON & WIRTH, 1979 MURPHREE & MULLEN, 1991 VITALE et al., 1981 JAMNBACK, 1965

Stages Described L P L P L P L L L

L E L

L

P P P P P P P P P P P P P P P

L L E L L P P P P P P P L L

P L P L P P L P L P P E P L P L P

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Table 1. Continued. Taxon

C. borinqueni C. cancer C. charruus C. chaverrii C. cochisensis C. crepuscularis

C. debilipalpis C. dicrourus C. edeni (as “possibly haematopotus”) C. fililductus C. floridensis C. furens (as dovei)

C. guyanensis C. haematopotus

C. hoffmani C. hylas (as sp. 3)

Reference BLANTON & WIRTH, 1979 MURPHREE & MULLEN, 1991 LINLEY, 1965b HOGUE & WIRTH, 1968 SPINELLI et al., 1993 SPINELLI & BORKENT, 2004a MURPHREE & MULLEN, 1991 THOMSEN, 1937 FOX, 1942 JAMNBACK, 1965 BLANTON & WIRTH, 1979 MURPHREE & MULLEN, 1991 BARBOSA, 1952, 1953 FORATTINI, 1957 WIRTH & SORIA, 1981 JONES, 1961 BLANTON & WIRTH, 1979 VITALE et al., 1981 LINLEY, 1970 BLANTON & WIRTH, 1979 PAINTER, 1927 DOVE et al., 1932 FOX, 1942 WIRTH, 1952d FORATTINI, 1957 JONES, 1961 LINLEY & KETTLE, 1964 JAMNBACK, 1965 LINLEY & DAVIES, 1971 BLANTON & WIRTH, 1979 HRIBAR & MULLEN, 1991 MURPHREE & MULLEN, 1991 FORATTINI & RABELLO, 1956 FORATTINI, 1957 THOMSEN, 1937 JONES, 1961 JAMNBACK, 1965 ATCHLEY & WIRTH, 1979 BLANTON & WIRTH, 1979 HRIBAR & MULLEN, 1991 MURPHREE & MULLEN, 1991 LINLEY & KETTLE, 1964 FOX, 1942 FORATTINI, 1957

Stages Described L P L P L P P L P L P P L P L P L P P L P P P P P P L P E L P P L P L P P L P L P E L P L L L P L P P P L P L P L P L L L P P P

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Table 1. Continued. Taxon

Reference

C. insignis

FORATTINI et al., 1956 FORATTINI, 1957 LINLEY, 1965b BLANTON & WIRTH, 1979 SPINELLI et al., 1993 LAMBERSON et al., 1992 WIRTH, 1952a WIRTH & SORIA, 1981 LUTZ, 1913 BARBOSA, 1952, 1953 FORATTINI, 1957 SPINELLI et al., 1993 WIRTH, 1952d JAMNBACK et al., 1958 JONES, 1961 JAMNBACK, 1965 BLANTON & WIRTH, 1979 HRIBAR & MULLEN, 1991 MURPHREE & MULLEN, 1991 JONES, 1961 MURPHREE & MULLEN, 1991 LAMBERSON et al., 1992 MURPHREE & MULLEN, 1991 LAMBERSON et al., 1992 PAINTER, 1927 FOX, 1942 FORATTINI, 1957 LUTZ, 1913 FORATTINI et al., 1960 RONDEROS & SPINELLI, 1989 DÍAZ et al., 2005 PAINTER, 1927

C. lahillei C. luteovenus C. macieli C. maruim

C. melleus

C. neopulicaris C. oklahomensis C. paraensis C. phlebotomus

C. reticulatus C. venezuelensis C. sp. Alluaudomyia bella (as splendida)

Stages Described L P L P P P P P L P L P E P P P P L P L P P L P L P L L P L P L P L P P P L L P P E L P L P

A. biestroi A. caribbeana A. distispinulosa A. schnacki

THOMSEN, 1937 WIRTH & GROGAN, 1981 SPINELLI, 1988 SPINELLI & WIRTH, 1984c SPINELLI, 1997b SPINELLI, 1983b

L P L P P P P L P

Baeodasymyia christopheri B. michaeli

BORKENT & CRAIG, 1999 BORKENT & CRAIG, 1999

E L P L P

Echinohelea lanei

WIRTH, 1994b

P

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Table 1. Continued. Taxon

Reference

Parabezzia alexanderi

P. balseiroi

GROGAN & WIRTH, 1977 WIRTH & GROGAN, 1981 SPINELLI & GROGAN, 1987 SPINELLI & GROGAN, 1987

Schizonyxhelea forattinii

BORKENT, 2000b

L P

Stilobezzia antennalis

L P L P L P P P P

S. panamensis S. rabelloi S. wygodzinskyi

THOMSEN, 1937 WIRTH & GROGAN, 1981 HRIBAR & MULLEN, 1991 FORATTINI & RABELLO, 1956 LANE & FORATTINI, 1961 LANE & FORATTINI, 1961 WIRTH & GROGAN, 1981 LUTZ, 1914 LANE et al., 1955 FORATTINI & RABELLO, 1956 CAZORLA et al., 2006 LANE et al., 1955 WIRTH & GROGAN, 1981 FORATTINI et al., 1960 BORKENT & CRAIG, 2001 LANE et al., 1955

Clinohelea horacioi C. nigripes

SPINELLI & DURET, 1993 SPINELLI & DURET, 1993

P P

Heteromyia clavata H. wokei

WIRTH & GROGAN, 1977 WIRTH & GROGAN, 1977

P P

Mallochohelea termophila

SPINELLI, 1984

P

Neobezzia amnicola

MAYER, 1959

P

Nilobezzia schwarzii

WIRTH, 1962 PALCHICK, 1981

P P

Amerohelea sordidipes

LANE et al., 1955

P

Bezzia bivittata B. blantoni B. brevicornis

HRIBAR & MULLEN, 1991 SPINELLI & WIRTH, 1989b SPINELLI, 1983c SPINELLI & WIRTH, 1989b SPINELLI & WIRTH, 1989b THOMSEN, 1937

S. bicolor S. chaconi S. coquilletti S. dorsofasciata (as a Palpomyia) S. dubitans S. fiebrigi S. glauca

B. bromeliae B. glabra

Stages Described P P P P

E P L P P P P P P P

L P P P P E L P

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Table 1. Continued. Taxon

Reference

Stages Described

B. roldani B. snowi

WIRTH, 1952a CHAN & LEROUX, 1967 PALCHICK, 1981 WIRTH, 1983a JOHANNSEN, 1905 MALLOCH, 1915a THIENEMANN, 1928 MAYER, 1934 MAYER, 1934 LENZ, 1934 LENZ, 1934 THOMSEN, 1937 GLUKHOVA, 1979 PALCHICK, 1981 WIRTH, 1983b SPINELLI, 1983c KNAUSENBERGER, 1987 HRIBAR & MULLEN, 1991 HRIBAR, 1993 MAYER, 1959 SPINELLI & WIRTH, 1989b SPINELLI & WIRTH, 1981, 1989b SPINELLI & WIRTH, 1991

Clastrieromyia dycei

SPINELLI & GROGAN, 1986

P

Pachyhelea pachymera

SPINELLI, 1983c

P

Palpomyia guarani P. lacustris P. subaspera (as essigi) P. wirthi

RONDEROS et al., 2004b LANE et al., 1955 WIRTH, 1952a GROGAN & WIRTH, 1979c LANE et al., 1955

Paryphoconus angustipennis P. flavidus (as lanei) P. mayeri P. oliveirai

MAYER, 1952 MAYER, 1959 MAYER, 1959 WIRTH & RATANAWORABHAN, 1972b

E

Stenoxenus johnsoni

WIRTH & RATANAWORABHAN, 1972b

E

B. nobilis (as setulosa) (as setulosa) (as chrysocoma) (as (as (as (as

chrysocoma) setulosa) chrysocoma) setulosa)

(as setulosa) (as acanthodes)

B. pulchripes

L P E L L L L L

P P P P

P P P P L P L P L P P L L L P P L P P

L P P P P P

P P P

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Table 2. Number of named species with described immatures for genera of Neotropical Ceratopogonidae (taxa identified only as “sp.” are not included). When the percentage of the Neotropical fauna known is zero, at least one immature stage is known only from elsewhere. Genus

Egg

Larva

Pupa

Total # Species

Percentage of Neotropical fauna

Leptoconops Forcipomyia Atrichopogon Dasyhelea Culicoides Paradasyhelea Allohelea Alluaudomyia Austrohelea Baeodasymyia Brachypogon Echinohelea Monohelea Parabezzia Schizonyxhelea Stilobezzia Clinohelea Heteromyia Pellucidomyia Johannsenomyia Mallochohelea Neobezzia Nilobezzia Amerohelea Bezzia Clastrieromyia Pachyhelea Palpomyia Phaenobezzia Paryphoconus Stenoxenus

1 0 0 2 4 0 0 0 0 1** 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0** 1** 1**

1 42 10 5 22 0 0 2 0 2 0 0 0 0 1 2 0 0 0 0* 0 0 0 0 4 0 0 1 0 0 0

1 45 10 14 34 0 0 5 0 2 0 1 0 2 1 10 2 2 0 0 1 1 1 1 8 1 1 4 0 3 0

1 45 10 14 35 0 0 5 0 2 0 1 0 2 1 11 2 2 0 0 1 1 1 1 9 1 1 4 0 4 1

8.3 21.3 10.0 24.6 13.2 0 0 26.3 0 40.0 0 9.1 0 8.3 50.0 17.2 14.3 18.2 0 0 16.7 12.5 33.3 9.1 19.6 25.0 50.0 8.7 0 10.0 6.3

Total

12

92

150

155

14.2

* The first instars of Johannsenomyia are superficially described by JOHANNSEN 1905: 108 and THOMSEN 1937: 73. ** Eggs extracted from or observed within adult.

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Table 3. Organisms transmitted by Neotropical Ceratopogonidae. Vectors are those known for this area and are in the following genera: Leptoconops, Forcipomyia or Culicoides. Taxon

Vector

Vertebrate Hosts

Location

VIRUSES Bunyavirus Ananindeua

C. paraensis

Birds, marsupials, rodents, primates

Brazil

Oropouche

C. paraensis

Sloths, humans; antibodies Trinidad, Panama, in other mammals Brazil, Peru

Utinga

Ceratopogonidae

Sloths; antibodies in other mammals

South and Central America

C. insignis,C. filarifer, C. pusillus

Wild and domestic ruminants

New World, Southern Europe, Africa, Asia, Australia

C. spp.

Horses, cattle, pigs, humans New World

Orbivirus Bluetongue

Vesiculovirus Vesicular stomatitis – New Jersey

Unclassified Arboviruses Buritirana C. spp.

Unknown

Brazil

FILARIOIDEA Dipetalonema caudispina

C. hollensis (in lab)

Monkeys (Cebidae)

South America

Dipetalonema gracile

C. hollensis (in lab)

Monkeys (Cebidae, Callithricidae)

Mexico, Central and South America

Dipetalonema marmosetae

C. furens

Monkeys (Cebidae, Callithricidae)

South America

Dipetalonema ozzardi

C. lahillei, C. paraensis Humans, other primates? L. bequaerti, C. furens C. phlebotomus

Central and South America

Dipetalonema perstans

C. spp.

Humans, other primates

Central and South America, Africa

Icosiella neglecta

F. spp.

Frogs

South America, Europe, Asia, Africa

Onchocerca cervicalis

C. spp.

Equidae

Worldwide

Onchocerca gutturosa

C. spp.

Bovidae

Worldwide

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2 CAPTURE AND STUDY OF CERATOPOGONIDAE Adult biting midges may be collected by any one of the following techniques, with varying results. 1. Light trap. Although many specialized light traps exist that work very well, a good sample may even be obtained at a lit window pane when there is good surrounding habitat. A simple and inexpensive technique is to place a funnel, approximately 20 cm in diameter above a jar of alcohol, close to a source of light. Another productive method is to lay a UV light on metal mesh (1 cm square, to keep out larger insects) which is lain over a shallow pan with soapy water; the water must be screened with fine mesh in the morning and the specimens preserved in alcohol. Ultraviolet light works well to attract adult midges but regular florescent or incandescent light may also bring in large numbers. An advantage of this technique is that it collects both males and females of a given species so that they can be associated. 2. CO2 trap. These traps collect females of those species that feed on vertebrate hosts (generally only those of species of Culicoides). A serious limitation of this technique is that males are not collected and therefore cannot be studied. 3. Sweeping vegetation, especially in areas with wet habitats, and aspirating the adults from the net. This provides a good method to sample diverse habitats and often results in species which are not discovered otherwise. This method also results in samples of both males and females. 4. Malaise traps. These work very well to sample males and females of many species. Larvae and pupae are obviously restricted to specific habitats and may be sampled directly from those aquatic, semiaquatic, or moist terrestrial habitats where they are found. The variety of habitats in which biting midge larvae and pupae may be found is described above. Sampling methods include the following: 1. Careful examination of wet substrate with a large magnifying glass or under a dissecting microscope. 2. Samples of substrate from aquatic habitats (e.g. mud, silt, detritus, leaves) placed in a pan with additional water. Most Ceratopogoninae larvae swim with a rapid serpentine motion and these may be removed with an eyedropper. Pupae almost always float at the water surface. Detritus removed from the margins of ponds, streams and lakes and placed in a white bucket allows for manual removal of pupae as they float. Some workers add salt or sugar to float larvae and pupae from sample substrate and this is the only practical way to collect those few species in which the larvae do not swim or the pupae do not float at the surface.

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3. Berlese funnels are a good method for extracting larvae from semiaquatic or merely moist material (e.g. mosses growing on trees in cloud forest). Once specimens are preserved in 70% alcohol (or some other preservative), most need to be placed on microscope slides (Fig. 1A) to be identified beyond the generic level. Methods vary but the following gives excellent results for slide mounting adult specimens stored in ethyl alcohol and which are not too old (3-4 years or, if stored under dark and very cold conditions (below 0 °C), over 10 years). This method works particularly well for material to be used for taxonomic studies. A more rapid method is given further below. Prepare for slide mounting by putting the following solutions into a series of four stender dishes: 15% acetic acid, 2-propanol, 2-propanol layered over clove oil, clove oil (each dish about 3/4 full). Each specimen needs to go through this sequence of solutions. Preparation of five specimens at a time is convenient and manageable, so that 20 dishes are in use at a time. Placing all the dishes on a thin board so that they can be moved around as a group on a table top makes it easier to look at specimens while they are in the solutions and still put all the dishes out of harm’s way while they are soaking. A series of slides needs to be ready, by cleaning these well with a cloth or paper towel and placing a label on each slide. These slides should be placed in a slide tray. In making the following preparations, round, 10 mm in diameter coverslips work very well. 1. Place the specimen into a dish of ethyl alcohol under the microscope. Remove the wings by grabbing the very base of the wing with fine forceps. Place both wings in the dish with 15% acetic acid. 2. For most specimens, separate head and abdomen from thorax. For very small specimens, just tear the membrane between the head and thorax and between the abdomen and thorax so that all the parts remain attached to each other. Place parts, or torn specimen, into a three dram vial filled about 1/4 to 1/3 full with 8% KOH. Prepare a total of five specimens in this way. Each vial needs to be numbered with a grease pencil so that it can be associated with the wings that are in the acetic acid. 3. Place vials of KOH into a beaker partially filled with water. Place beaker on hot plate and heat water to boiling point. Specimens will clear after 2-5 minutes, depending on the size of the specimens. It is important that specimens of a similar size are “cooked” at the same time so that they all clear at the same rate. 4. The following step requires some speed because specimens are damaged by remaining in hot KOH for too long. Remove the specimen from the vial into a dish with some 8% KOH in it and put under the microscope to see if it is properly cleared. The muscles should have dissolved and freely flow out of the specimen if squeezed very, very gently with the forceps. If the specimens are over-cleared they will make for a poor microscope preparation. If cleared sufficiently, place specimen in acetic acid along with the associated wings. Leave for 15 minutes (or longer if necessary). 5. Move specimen to 2-propanol for 15 minutes (or longer if necessary). 6. Move specimen to 2-propanol layered over clove oil. Leave until parts have sunk to the bottom of the dish which generally takes about 20-60 minutes, depending on the size of the specimen (specimens can be left longer if necessary). After a day or two, the 2propanol will mix with the clove oil and some more 2-propanol should be added so that there is always a distinct layer between the 2-propanol and the clove oil. If the clove oil has

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too much 2-propanol mixed into it, leave the dish without a lid for a few hours in a well ventilated area and the 2-propanol will evaporate from the mixture. 7. Move specimen to pure clove oil and leave for at least 30 minutes (or longer if necessary). 8. Place small drop of Canada Balsam on the microscope slide; place the wings in the drop and cover with a clean coverslip. If Canada Balsam has already hardened too much to move the wings around easily, add a small drop of xylene, which will make the Canada Balsam more fluid. Use xylene for the following procedures as well; it is always needed because Canada Balsam dries faster than the time it takes for a good slide to be made. 9. Place three small drops of Canada Balsam on the microscope slide and put the head, thorax and abdomen into each. If the head, thorax and abdomen are still partially attached (small specimens), put specimen into the drop which will contain just the thorax and remove the head and abdomen; put the head and abdomen into their proper places. 10. Remove the right legs and place all these under a separate coverslip. 11. Arrange the thorax so that the left side is facing upward. Orient the head so that the front of the head is oriented upward and the antennae are splayed outward. Do not add a coverslip. 12. Details of the genitalia are often very important in species identification, so that the following step is a very important one. Arrange the abdomen so that the ventral part of the genitalia is facing upward. In some males, the genitalia may be only partially turned in relation to the rest of the abdomen and in this case, sometimes the genitalia will need to be removed from the abdomen (by tearing with the forceps and a pin) so that the separated genitalia lies with the ventral side pointed up. The rest of the abdomen should be straight, with the tergites dorsal, if possible. Do not add a coverslip. 13. Put the slide into the drying oven at 40-50 °C. After 1-2 days, the Canada Balsam with the head, abdomen and thorax will have dried somewhat. Add some more Canada Balsam to each of these and very gently put a coverslip over each, so they are not crushed. The partially dried Canada Balsam should keep these parts from being damaged. While the slide is drying, some of the Canada Balsam may leave a small air bubble under the coverslip. If so, add a tiny bit of Canada Balsam. Don’t worry if the added Canada Balsam traps a bubble under the coverslip; the bubble nearly always moves out by itself (after some time). 14. Put the slide back into the drying oven. Check for air bubbles for the next several days and add Canada Balsam as needed. Remove the slide after 1-2 weeks and put into a slide box. Keep the slide box upright, so the slides are still horizontal. Larvae and pupae may be treated with the same solutions and should be mounted dorsal side up. Larvae require no dissection but the operculum of the pupa as well as one of the respiratory organs should be removed and placed under a separate coverslip (leaving the Canada Balsam to dry for a day or two and then adding the coverslips). If the larva or pupa is represented by an exuvia, the specimens obviously do not require clearing and may be placed directly into 2-propanol and processed from there. A simpler method for mounting specimens, which is particularly useful for mass mounting material which is likely to include previously described species, is as follows: remove specimens from 70% ethanol (many can be done as one lot), very briefly blot excess alcohol, transfer to 100% ethanol saturated with phenol, leave overnight or longer, and mount

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specimens on microscope slides in a one to one mixture of Canada Balsam / phenolethanol solution. Depending on the stage, specimens may be dissected on the slide and one or more coverslips be placed on the parts. As the phenol-ethanol solution evaporates during drying, more Canada Balsam needs to be added. This method provides good specimens but does not remove internal tissues making the observation of some structures difficult to see clearly. Workers using this method should use a fume hood as both xylene and phenols are dangerous to human health.

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3 REARING CERATOPOGONIDAE The immatures may be reared using one or more methods. Field collected female adults of some species may be induced to lay eggs by keeping them in a vial with a loose-fitting lid or a cotton ball, or in a small Petri dish with wet mud (as described by BORKENT & CRAIG, 2004). In such instances, food for hatching larvae will be somewhat uncertain and, depending on the genus, larvae may be offered either rich substrate with abundant microorganisms or an agar plate with nematodes (HUNT & SCHMIDTMANN, 2004). Larvae collected from nature will often molt to the pupal stage if given some of the substrate in which they were discovered, along with additions of either a rich microbial soup or small prey items, such as nematodes (depending on the taxon collected). It is always an advantage to collect mature larvae of either the third or fourth instar to enhance the likelihood of maturation. Pupae may either be left in a Petri dish or other small container with some of the substrate in which they were living or with a piece of moist to wet filter paper. Emerging adults should be kept alive for at least 12-24 hours at room temperature before preservation to ensure complete hardening of the cuticle and maturation of pigmentation patterns.

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4 PHYLOGENETIC RELATIONSHIPS AND FOSSIL RECORD OF CERATOPOGONIDAE The phylogenetic and fossil record of the Ceratopogonidae is one of the best understood among insects and provides a good example of combining cladistic analysis of extant taxa with interpretation of a detailed fossil record (BORKENT, 2000a; BORKENT & CRAIG, 2004). The biting midges are an exemplary group for studying evolutionary relationships because of three outstanding features. First, they are diverse in numbers of species. Generally, when a group is species-rich, there are intermediate species or groups present which provide good clues on the direction of evolutionary change of a given structure. Second, the biting midges have three very different stages (larva, pupa, adult) each of which have diversified structurally. Unlike many other groups, the immatures of biting midges have evolved in many different morphological directions. Therefore, it is possible to compare conclusions about relationships based on one stage against the conclusions based on a different stage. Third, the biting midges have an incredible fossil record in amber that has been investigated in some detail. We now have 11 major deposits ranging from 17 to 121 million years of age, with thousands of specimens studied. The fossil biting midges are largely in excellent condition and most taxonomic features can be studied in detail. These fossils provide an excellent gauge as to the accuracy of our interpretation of evolutionary relationships based on living species. General conclusions show that the Leptoconopinae are the sister group of all remaining Ceratopogonidae, the Forcipomyiinae and Dasyheleinae are sister groups, and that the preceding subfamilies form the sister group of the Ceratopogoninae. Within the Ceratopogoninae, the Culicoidini are the sister group of the remaining taxa (BORKENT & CRAIG, 2004). Significant research remains to be completed to further interpret the relationships between most of the remaining genera of the Ceratopogoninae.

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5 ZOOGEOGRAPHY Our understanding of the zoogeography of Ceratopogonidae is rather limited. This is for two important reasons. One is that the phylogenetic relationships between many taxa are unknown and it is therefore impossible to interpret the significance of their distributions in light of those relationships. Secondly, in the Neotropical Region, large areas have not been sampled, including much of the Amazon Basin and nearly all of the Andes. Without doubt, there remain a great number of species to be discovered. Nevertheless, there are some broad geographic patterns which are apparent. Most notable is the distinctive fauna of Patagonia. The following genera are endemic to the area, with most of these present in forested habitats: Borkenthelea SPINELLI & GROGAN (three forest species), Diaphanobezzia INGRAM & MACFIE (four species, three from forest, one from steppe), Isthmohelea INGRAM & MACFIE (monotypic, from forest), Notiohelea GROGAN & WIRTH (two forest species), Physohelea GROGAN & WIRTH (two forest species), Austrosphaeromias SPINELLI (two forest species). In addition, the following are restricted within the Neotropical Region to Patagonia but are also present in other, mostly southern hemisphere areas: Austrohelea WIRTH & GROGAN (four species (three undescribed) from Patagonian forest; six others from Australia or New Zealand), Macrurohelea INGRAM & MACFIE (13 species (two undescribed) from Patagonian forest, except one species that extends northward to arid zones of Chile and Argentina, one undescribed species in the Somuncura Plateau in the Patagonian steppe; three described and two unnamed from Australia), and Paradasyhelea MACFIE (three Patagonian species, two from forest and one from steppe; seven others from New Caledonia, Australia, New Zealand; one Nearctic species). The latter three genera may have southern hemisphere connections that reflect patterns of continental drift but their phylogenetic relationships await analysis to test this possibility. Study of other ceratopogonids in Patagonia indicates that most species are endemic in this region including species of Forcipomyia, Dasyhelea, and Stilobezzia (all of which are members of the subgenus Acanthohelea KIEFFER and all restricted to Nothofagus forest). In addition to those endemic Patagonian genera discussed above, several genera or subgenera of Ceratopogonidae are restricted or nearly restricted to the Neotropical Region: Forcipomyia (Rhynchoforcipomyia WIRTH & DOW) (seven species), Culicoides (Anilomyia VARGAS) (17 species), C. (Cotocripus BRÈTHES) (five species), C. (Macfiella FOX) (two species), C. (Mataemyia VARGAS) (15 species), C. (Psychophaena PHILIPPI) (two species), Baeodasymyia CLASTRIER & RACCURT (five species), Baeohelea WIRTH & BLANTON (one species), Bahiahelea WIRTH (one species), Cacaohelea WIRTH & GROGAN (two species), Leptohelea WIRTH & BLANTON (one species), Parastilobezzia WIRTH & BLANTON (one species), Echinohelea MACFIE (Echi-

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noideshelea WIRTH) (one species), Fittkauhelea WIRTH & BLANTON (one species), Schizonyxhelea CLASTRIER (two species), Clinohelea KIEFFER (Ceratobezzia KIEFFER) (one species), Groganhelea SPINELLI & DIPPOLITO (one species), Lanehelea WIRTH & BLANTON (two species), Neobezzia WIRTH & RATANAWORABHAN (eight species), Sphaerohelea SPINELLI & FELIPPE-BAUER (one species), Clastrieromyia SPINELLI & GROGAN (four species), Pachyhelea WIRTH (one of the two species extends north into Texas), Amerohelea GROGAN & WIRTH (one of the 11 known species extends north into Texas), Paryphoconus ENDERLEIN (one of 40 species is Nearctic). Several of these groups are clearly monophyletic and this indicates that they have speciated within the Neotropical Region (i.e. Baeodasymyia + Baeohelea, Clastrieromyia, Paryphoconus). A few of these genera, each with only 1-2 species, are endemic to Central America and northern South America: Cacaohelea (Costa Rica), Leptohelea (Costa Rica, Colombia), Parastilobezzia (Costa Rica, Colombia). In Central America, the genus Ceratoculicoides WIRTH & RATANAWORABHAN is present in tropical Mexico (one described species) and Costa Rica (one undescribed species) and this is likely a southern extension of an otherwise Holarctic genus. Within Costa Rica we recognize a pattern, common in many groups of organisms, that lowland species tend to be broadly distributed in the Neotropical Region and those that occur higher than 1500 meters in elevation tend to be more restricted in distribution and are often endemic to Costa Rica or Costa Rica and Panama. This general pattern will almost certainly be true in other high elevation areas in the Neotropics once they are investigated more thoroughly.

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6 CLASSIFICATION Ceratopogonidae belong to the superfamily Chironomoidea and are the sister group of the Chironomidae. Worldwide, extant Ceratopogonidae are divided into four well established subfamilies, based on cladistic relationships (BORKENT, 2000a; BORKENT & CRAIG, 2004): Leptoconopinae (2 genera), Forcipomyiinae (2 genera), Dasyheleinae (1 genus) and Ceratopogoninae (104 genera). All subfamilies are known from the Neotropical Region (Table 4). REMM (1975) proposed recognizing the Forcipomyiinae and Dasyheleinae as tribes in the subfamily Forcipomyiinae and, as sister groups, this is conceivable but has not yet been widely accepted. The Ceratopogoninae, which make up the bulk of the family both generically and at the species level, is divided into 6 tribes: Culicoidini (3 genera), Ceratopogonini (57 genera), Heteromyiini (8 genera), Sphaeromiini (28 genera), Palpomyiini (6 genera) and the Stenoxenini (2 genera). The relationships among these tribes are poorly understood, although it is clear that the Culicoidini are the sister group of the remaining tribes, the Ceratopogonini are paraphyletic in relation to the remaining tribes and that the Palpomyiini and Stenoxenini form a monophyletic group. The relationships among most of the genera of Ceratopogonini and the relationships among and within the genera of Heteromyiini and Sphaeromiini are particularly in need of cladistic analysis. Although 5925 extant species have been described worldwide (as of May, 2007) it is clear that many more remain unnamed. It is certainly reasonable to suggest that at least 15,000 species exist on our planet. Within the Neotropical Region, the Andes are virtually uncollected, in spite of the fact that Ceratopogonidae are abundant and diverse there (BORKENT, pers. obs.). Many areas of Brazil, Peru, northern Chile and northwestern Argentina are also particularly poorly known. Limited portions of the tropical rain forests of Central America and Amazonia were extensively collected during the 1960s and 1970s, especially by F.S. BLANTON and E. FITTKAU, respectively, providing the basis for numerous taxonomic papers mainly by W.W. WIRTH and his coworkers. Nevertheless, recent collections of biting midges in Costa Rica and Peruvian Amazonia reveal the presence of many more undescribed species (BORKENT & SPINELLI, pers. obs.), strongly suggesting that further collecting will be richly rewarded even in areas which have been previously sampled. Revisions are needed for virtually all genera; even Culicoides, which has drawn the most taxonomic attention, has many undescribed species and its subgeneric classification is in great need of analysis.

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Table 4. Catalog of the Ceratopogonidae of the Neotropical Region. The names of those species of Leptoconops and Culicoides which are known to bite humans are underlined.

SUBFAMILY LEPTOCONOPINAE NOÈ, 1907: 143 Genus LEPTOCONOPS SKUSE LEPTOCONOPS SKUSE, 1889: 288. Type species: Leptoconops stygius SKUSE, by monotypy. TERSESTHES TOWNSEND, 1893: 370. Type species: Tersesthes torrens TOWNSEND, by original designation. MYCTEROTYPUS NOÈ, 1905: 114. Type species: Mycterotypus bezzii NOÈ, designation by CARTER, 1921: 3. PROTERSESTHES KIEFFER, 1921c: 107. Type species: Tersesthes brasiliensis LUTZ, by original designation. REFERENCES: RONDEROS, 1988 (key to females from Argentina); RONDEROS & SPINELLI, 1992 (key to females); WIRTH & ATCHLEY, 1973 (key to subgenera of females, other than Proleptoconops; key to Nearctic females). Subgenus BRACHYCONOPS WIRTH & ATCHLEY BRACHYCONOPS WIRTH & ATCHLEY, 1973: 15 (as subgenus of Leptoconops). Type species: Leptoconops californiensis WIRTH & ATCHLEY, by original designation. patagoniensis RONDEROS, 1990b: 423 (female; Argentina). Distr.- Southern Argentina. Subgenus HOLOCONOPS KIEFFER HOLOCONOPS KIEFFER, 1918: 135. Type species: Leptoconops kerteszi KIEFFER, by original designation. MICROCONOPS KIEFFER, 1921c: 108. Type species: Microconops vexans KIEFFER, by original designation. REFERENCE: CLASTRIER & WIRTH, 1978 (key to Nearctic males and females). bassoi RONDEROS & SPINELLI, 1993: 305 (male; Chile). Distr.- Northern Chile. bequaerti (KIEFFER), 1925: 405 (Holoconops, as becquaerti; female; Honduras). Distr.- USA (Texas) to Panama and Trinidad. hondurensis HOFFMAN, 1926: 135. Honduras. knowltoni CLASTRIER & WIRTH, 1978: 26 (male, female; USA, California). Distr.- USA (western) south to Mexico (Sinaloa), Brazil (Santa Catarina).

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Subgenus LEPTOCONOPS SKUSE brasiliensis (LUTZ), 1913: 66 (Tersesthes; female; Brazil). Distr.- Argentina (Salta), Brazil (Amazonas), Uruguay. casali CAVALIERI & CHIOSSONE, 1966: 45 (female; Argentina). Distr.- Argentina (Salta, San Luis), Uruguay. chilensis FORATTINI, 1958: 38 (female; Chile). Distr.- Bolivia, Chile, Argentina (Jujuy, Salta, La Pampa). panamensis RONDEROS & SPINELLI, 1993: 308 (male; Panama). Distr.- Panama. petrocchiae SHANNON & DEL PONTE, 1927: 734 (female; Argentina). Distr.- Northwestern Argentina. ricardoi RONDEROS & SPINELLI, 1992: 43 (female; Argentina). Distr.- Southwestern Argentina. venezuelensis ORTÍZ, 1952a: 165 (female; Venezuela). Distr.- Venezuela. Subgenus MEGACONOPS WIRTH & ATCHLEY MEGACONOPS WIRTH & ATCHLEY, 1973: 18 (as subgenus of Leptoconops). Type species: Leptoconops floridensis WIRTH, by original designation. floridensis WIRTH, 1951a: 282 (female; USA, Florida). Distr.- USA (Florida), Jamaica, Colombia. SUBFAMILY FORCIPOMYIINAE LENZ, 1934: 96 Genus ATRICHOPOGON KIEFFER ATRICHOPOGON KIEFFER, 1906a: 53 (as subgenus of Ceratopogon). Type species: Ceratopogon exilis COQUILLETT (= Ceratopogon levis COQUILLETT), designation by COQUILLETT, 1910: 512. REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE, 1931 (key males and females from Patagonia and southern Chile); MACFIE, 1939b (key to males and females from the Neotropical Region). Subgenus ATRICHOPOGON KIEFFER adamsoni MACFIE, 1937a: 4 (female; Trinidad). Distr.- Mexico (Chiapas), Trinidad. albinensis INGRAM & MACFIE, 1931: 228 (female; Argentina). Distr.- Argentina (Buenos Aires). altivolans MACFIE, 1949: 113 (female; Mexico). Distr.- Mexico (Chiapas). aridus SPINELLI & MARINO, in SPINELLI et al., 2006: 304 (male, female; Argentina). Distr.Argentina (Mendoza, Rio Negro). asuturus BORKENT & PICADO, 2004: 29 (male; Costa Rica). Distr.- Costa Rica. auricoma KIEFFER, 1917b: 300 (male, female; Colombia). Distr.- Colombia. balseiroi SPINELLI, 1982: 206 (male, female; Argentina). Distr.- Argentina (Buenos Aires). barbatus BORKENT & PICADO, 2004: 23 (male; Costa Rica). Distr.- Costa Rica.

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beccus BORKENT & PICADO, 2004: 31 (male; Costa Rica). Distr.- Costa Rica. bicuspis BORKENT & PICADO, 2004: 9 (male, female; Costa Rica). Distr.- Costa Rica. bifidus EWEN, in EWEN & SAUNDERS, 1958: 694 (larva, pupa, male, female; Brazil). Distr.Brazil (Rio de Janeiro). brasiliensis MACFIE, 1939b: 184 (female; Brazil). Distr.- Brazil (Santa Catarina). brevipalpis MACFIE, 1944: 298 (male, female; Trinidad). Distr.- Trinidad. caribbeanus EWEN, in EWEN & SAUNDERS, 1958: 683 (larva, pupa, male, female; Tobago). Distr.- Tobago. carnatus BORKENT & PICADO, 2004: 13 (male; Costa Rica). Distr.- Costa Rica. carpinteroi MARINO & SPINELLI, 2004c: 157 (male, female; Argentina). Distr.- Northern Argentina. casali CAVALIERI & CHIOSSONE, 1973: 153 (male, female; Argentina). Distr.- Northeastern Argentina. colossus BORKENT & PICADO, 2004: 10 (male, female; Costa Rica). Distr.- Costa Rica. columbianus KIEFFER, 1917b: 303 (female; Colombia). Distr.- Colombia, Brazil (Santa Catarina). comechingon SPINELLI & MARINO, in SPINELLI et al., 2006: 306 (male, female; Argentina). Distr.- Argentina (Córdoba, Rio Negro). costalis MACFIE, 1939b: 179 (female; Brazil). Distr.- Brazil (Santa Catarina). costaricae MACFIE, 1953: 98 (female; Costa Rica). Distr.- Costa Rica. cryptogamus MACFIE, 1939b: 182 (female; Brazil). Distr.- Brazil (Santa Catarina). delpontei CAVALIERI & CHIOSSONE, 1972: 121 (male, female; Argentina). Distr.- Northeastern Argentina. depilis MACFIE, 1939b: 178 (female; Brazil). Distr.- Brazil (Santa Catarina). didymothecae MACFIE, 1953: 99 (female; Costa Rica). Distr.- Costa Rica. domizii SPINELLI, 1982: 201 (male, female; Argentina). Distr.- Argentina (Buenos Aires). echinodes MACFIE, 1939b: 194 (female; Brazil). Distr.- Brazil (Santa Catarina). endemicus SPINELLI & MARINO, in SPINELLI et al., 2006: 304 (male, female; Argentina). Distr.- Argentina (Rio Negro). eucnemus MACFIE, 1939b: 185 (male, female; Brazil). Distr.- Brazil (Santa Catarina). fiebrigi KIEFFER, 1917b: 302 (male; Paraguay). Distr.- Paraguay. fimbriatus MACFIE, 1939b: 194 (as variety of gordoni; female). Distr.- Brazil (Santa Catarina). flavicaudae MACFIE, 1939b: 181 (female; Brazil). Distr.- Brazil (Santa Catarina). pilosior MACFIE, 1939b: 182 (as variety of flavicaudae). Brazil. flavipes LUTZ, 1914: 90 (female; Brazil). Distr.- Brazil (type locality not stated). flumineus MACFIE, 1935a: 52 (female; Brazil). Distr.- Brazil (Maranhão, Santa Catarina). gamboai BORKENT & PICADO, 2004: 26 (male; Costa Rica). Distr.- Costa Rica. glaber MACFIE, 1935a: 50 (female; Brazil). Distr.- Costa Rica, Trinidad, Brazil (Pará, Maranhão). globulifer MACFIE, 1939b: 191 (female; Brazil). Distr.- Brazil (Santa Catarina). gordoni MACFIE, 1938: 164 (female; Trinidad). Distr.- Costa Rica, Trinidad, Brazil (Santa Catarina). granditergitus BORKENT & PICADO, 2004: 27 (male; Costa Rica). Distr.- Costa Rica. granditibialis BORKENT & PICADO, 2004: 20 (male, female; Costa Rica). Distr.- Costa Rica. guianensis MACFIE, 1940b: 184 (female; Guyana). Distr.- Guyana. harrisi MACFIE, 1938: 163 (female; Trinidad). Distr.- Trinidad.

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homofacies SPINELLI, in SPINELLI et al., 1989: 734 (male, female; Argentina). Distr.- Argentina (Buenos Aires). inacayali SPINELLI & MARINO, in SPINELLI et al., 2006: 309 (male, female; Argentina). Distr.Argentina (Chuvut, Rio Negro). incultus EWEN, in EWEN & SAUNDERS, 1958: 699 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica. insignipalpis MACFIE, 1940a: 73 (male, female; Brazil). Distr.- Northeastern Brazil. insigniventris MACFIE, 1935a: 51 (female; Brazil). Distr.- Brazil (Maranhão), Trinidad. lacajae MACFIE, 1953: 100 (male; Costa Rica). Distr.- Costa Rica. lituratus (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- St. Vincent. lobatus BORKENT & PICADO, 2004: 14 (male, female; Costa Rica). Distr.- Costa Rica. longicornis EWEN, in EWEN & SAUNDERS, 1958: 700 (larva, male, female; Costa Rica). Distr.Costa Rica. maculipennis CLASTRIER, 1968: 89 (male, female; French Guiana). Distr.- French Guiana. magnus BORKENT & PICADO, 2004: 16 (male, female; Costa Rica). Distr.- Costa Rica. mendozae INGRAM & MACFIE, 1931: 229 (female; Argentina). Distr.- Argentina (Mendoza). mexicanus HUERTA, 2001: 373 (male, female; Mexico). Distr.- Mexico (Chiapas). nanus MACFIE, 1940b: 184 (male, female; Guyana). Distr.- Guyana. nebulosus MACFIE, 1939b: 192 (female; Brazil). Distr.- Brazil (Santa Catarina). novaeteutoniae MACFIE, 1939b: 189 (female; Brazil). Distr.- Brazil (Santa Catarina). nubeculosus MACFIE, 1949: 113 (female; Mexico). Distr.- Mexico (Chiapas). obfuscatus INGRAM & MACFIE, 1931: 173 (male, female; Argentina). Distr.- Southern Chile, southwestern Argentina south to Tierra del Fuego. obnubilus INGRAM & MACFIE, 1931: 175 (male, female; Argentina). Distr.- Southern Chile, southwestern Argentina. chilensis INGRAM & MACFIE, 1931: 175. Argentina. assimilis INGRAM & MACFIE, 1931: 176. Argentina. obscurus EWEN, in EWEN & SAUNDERS, 1958: 685 (larva, pupa, female; Puerto Rico). Distr.Puerto Rico. ocumare (ORTÍZ), 1952b: 254 (Monohelea; male; Venezuela). Distr.- Venezuela. ornatipennis CLASTRIER, 1987: 271 (male; French Guiana). Distr.- French Guiana. pachycnemus MACFIE, 1953: 101 (female; Costa Rica). Distr.- Costa Rica. pallidipes KIEFFER, 1917b: 301 (female; Paraguay). Distr.- Mexico (Chiapas), Costa Rica, Paraguay. palpalis MACFIE, 1939b: 196 (male, female; Brazil). Distr.- Brazil (Santa Catarina). pectinatus MACFIE, 1939b: 195 (female; Brazil). Distr.- Brazil (Santa Catarina). penicillatus DELÉCOLLE & RIEB, 1994: 274 (female; Guadeloupe). Distr.- Guadeloupe. peruvianus KIEFFER, 1917b: 302 (male; Peru). Distr.- Peru. piceiventris KIEFFER, 1917b: 301 (female; Paraguay). Distr.- Paraguay. pictipennis CLASTRIER, 1979: 30 (male; French Guiana). Distr.- French Guiana. quartibrunneus BORKENT & PICADO, 2004: 30 (male; Costa Rica). Distr.- Costa Rica. redactus BORKENT & PICADO, 2004: 32 (male; Costa Rica). Distr.- Costa Rica. remigatus EWEN, in EWEN & SAUNDERS, 1958: 687 (larva, pupa, male, female; Brazil). Distr.Brazil (Rio de Janeiro).

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rusticus MACFIE, 1939b: 190 (female; Brazil). Distr.- Brazil (Santa Catarina). sanctaeclarae MACFIE, 1949: 114 (male, female; Mexico). Distr.- Mexico (Chiapas). sanctilaurentii KIEFFER, 1917b: 303 (female; Paraguay). Distr.- Paraguay. saundersi EWEN, in EWEN & SAUNDERS, 1958: 691 (larva, pupa, male, female; Puerto Rico). Distr.- Puerto Rico. sequax (WILLISTON), 1896: 282 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent. setosilateralis BORKENT & PICADO, 2004: 33 (male; Costa Rica). Distr.- Costa Rica. seudoobfuscatus SPINELLI, 1982: 208 (male, female; Argentina). Distr.- Argentina (Buenos Aires). similis SPINELLI & MARINO, in SPINELLI et al., 2006: 316 (male, female; Argentina). Distr.Southern Chile, southwestern Argentina. spinosus BORKENT & PICADO, 2004: 12 (male; Costa Rica). Distr.- Costa Rica. taeniatus MACFIE, 1939b: 186 (female; Brazil). Distr.- Brazil (Santa Catarina). talarum SPINELLI, 1982: 204 (male, female; Argentina). Distr.- Argentina (Buenos Aires). tapantiensis BORKENT & PICADO, 2004: 34 (male; Costa Rica). Distr.- Costa Rica. tirzae BORKENT & PICADO, 2004: 28 (male; Costa Rica). Distr.- Costa Rica. thersites (WILLISTON), 1896: 280 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent. trinidadensis MACFIE, 1937a: 5 (male; Trinidad). Distr.- Trinidad. tuberculatus EWEN, in EWEN & SAUNDERS, 1958: 709 (larva, pupa, female, male; Trinidad). Distr.- Trinidad. umbratilis MACFIE, 1935a: 51 (female; Brazil). Distr.- Guyana, Brazil (Maranhão). utricularis MACFIE, 1953: 102 (female; Costa Rica). Distr.- Costa Rica. warmkei WIRTH, 1956c: 243 (male, female; Puerto Rico). Distr.- USA (Florida, Puerto Rico). woodfordi MACFIE, 1938: 162 (female; Trinidad). Distr.- Trinidad. yolancae BORKENT & PICADO, 2004: 24 (male; Costa Rica). Distr.- Costa Rica. Subgenus LOPHOMYIDIUM CORDERO LOPHOMYIDIUM CORDERO, 1929: 94. Type species: Lophomyidium uruguayense CORDERO, by original designation. REFERENCE: WIRTH 1994a (key to Nearctic males and females). archboldi WIRTH, 1994a: 27 (male, female; USA, Florida). Distr.- USA (California to Florida) to Argentina. fusculus (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, New Jersey). Distr.- Northern Eurasia, North America, Central America and Caribbean to Bolivia and Brazil (Pará). uruguayensis (CORDERO), 1929: 95 (Lophomyidium; male; Uruguay). Distr.- Paraguay, eastern Argentina, Uruguay. Genus FORCIPOMYIA MEIGEN FORCIPOMYIA MEIGEN, 1818: 73, 75. Type species: Tipula bipunctata Linnaeus, designation by WESTWOOD, 1840: 126. Generic name first published in synonymy with Ceratopogon but available under ICZN Code Article 11(e).

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TETRAPHORA PHILIPPI, 1865: 630. Type species: Tetraphora fusca PHILIPPI, by monotypy. REFERENCES: DEBENHAM, 1987 (key to males and females to subgenera from Australasian Region which includes all Neotropical subgenera other than Rhynchoforcipomyia and Saliohelea); INGRAM & MACFIE, 1931 (key to males and females from Patagonia and southern Chile); WIRTH & RATANAWORABHAN, 1978 (key to females to subgenera; their genus A near Lepidohelea = Pedilohelea). Subgenus BLANTONIA WIRTH & DOW BLANTONIA WIRTH & DOW, 1971: 289 (as subgenus of Forcipomyia). Type species: Forcipomyia caribbea WIRTH & DOW, by original designation. caribbea WIRTH & DOW, 1971: 291 (male, female; Jamaica). Distr.- USA (Florida) to Trinidad, Mexico (Yucatán), Belize, Guyana. Subgenus CALOFORCIPOMYIA SAUNDERS CALOFORCIPOMYIA SAUNDERS, 1957: 680 (as subgenus of Forcipomyia). Type species: Forcipomyia caerulea SAUNDERS, by original designation. REFERENCE: UTMAR & WIRTH, 1976 (key to New World males and females). caerulea SAUNDERS, 1957: 681 (male, female; Brazil). Distr.- Panama, Colombia, Dominica, Brazil (Pará, Rio de Janeiro). . copanensis UTMAR & WIRTH, 1976: 129 (male, female; Honduras). Distr.- Distr.- Honduras, Costa Rica, northern Argentina. eukosma MACFIE, 1939b: 148 (male, female; Brazil). Distr.- Colombia, Dominica, Brazil (Pará, São Paulo, Rio de Janeiro, Santa Catarina). furcifera MACFIE, 1940e: 920 (male; Brazil). Distr.- Brazil (Santa Catarina). glauca MACFIE, 1934: 144 (Great Britain). Distr.- Europe, North America (Alberta to Nova Scotia to Arkansas and Florida) to Colombia and southern Brazil. hatoensis UTMAR & WIRTH, 1976: 131 (male, female; Panama). Distr.- Mexico (Sinaloa), Costa Rica, Panama, Colombia. hermosa UTMAR & WIRTH, 1976: 117 (male, female; Brazil). Distr.- Costa Rica, Panama, Colombia, Jamaica, Panama, Brazil (Amazonas, Pará). nigrescens MACFIE, 1939b: 156 (male, female; Brazil). Distr.- Panama, Brazil (São Paulo, Rio de Janeiro, Santa Catarina). remigera UTMAR & WIRTH, 1976: 121 (male, female; Colombia). Distr.- Panama, Colombia, French Guiana, Brazil (Pará). sabalitensis UTMAR & WIRTH, 1976: 127 (male, female; Costa Rica). Distr.- Honduras to Colombia, Brazil (Paraná, Santa Catarina), northeastern Argentina. varicolor SAUNDERS, 1957: 683 (larva, pupa, male; Brazil). Distr.- Brazil (Rio de Janeiro).

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Subgenus EUPROJOANNISIA BRÈTHES EUPROJOANNISIA BRÈTHES, 1914: 155. Type species: Euprojoannisia platensis BRÈTHES, by original designation. REFERENCES: BYSTRAK & WIRTH, 1978 (key to Nearctic larvae, pupae, males, females); HUERTA, 1996 (key to males and females from Mexico). blantoni SORIA & BYSTRAK, 1975: 3 (larva, pupa, male, female; Brazil). Distr.- Eastern USA to Ecuador, Brazil (Bahia). bromeliae SAUNDERS, 1957: 665 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro). calcarata (COQUILLETT), 1905: 64 (Ceratopogon; male; Mexico). Distr.- Eastern USA, Mexico (Tabasco, Quintana Roo). dowi BYSTRAK & WIRTH, 1978: 21 (male, female; USA, Florida). Distr.- USA (Florida), Mexico (Quintana Roo). esteparia MARINO & SPINELLI, 2001b: 12 (male, female; Argentina). Distr.- Southern Argentina. falcifera SAUNDERS, 1959: 39 (larva, pupa, male, female; Trinidad). Distr.- Venezuela, Trinidad. galliarii MARINO & SPINELLI, 1999a: 5 (female; Argentina). Distr.-. Argentina (Misiones). longispina SAUNDERS, 1957: 669 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro). lota (WILLISTON), 1896: 282 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent. mortuifolii SAUNDERS, 1959: 35 (larva, pupa, male, female; Trinidad). Distr.- USA (Florida) to St. Lucia. navaiae BYSTRAK & WIRTH, 1978: 31 (male, female; USA, Florida). Distr.- USA (Florida), Mexico (Yucatán). platensis (BRÈTHES), 1914: 156 (Euprojoannisia; female; Argentina). Distr.- Argentina (Buenos Aires). quasiingrami MACFIE, 1939b: 164 (male; Brazil). Distr.- USA (Florida) south through Caribbean to Brazil (Bahia, Santa Catarina). setigera SAUNDERS, 1959: 38 (larva, pupa, male, female; Trinidad). Distr.- Trinidad. setosicrus (KIEFFER), 1906b: 357 (Ceratopogon; male; Chile). Distr.- Southern Argentina and Chile. spatulifera SAUNDERS, 1957: 667 (larva, pupa, female; Brazil). Distr.- Brazil (Bahia, São Paulo), Venezuela. unica BYSTRAK & WIRTH, 1978: 44 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas. Subgenus FORCIPOMYIA MEIGEN REFERENCES: HUERTA, 1996 (key to females from Mexico); MARINO & SPINELLI, 1999b (key to species groups); MARINO & SPINELLI, 2001a (key to males and females from Patagonia); MARINO & SPINELLI, 2002 (key to squamitibia group); WIRTH, 1982a (key to argenteola group). argenteola Species Group argenteola MACFIE, 1939b: 146 (female; Brazil). Distr.- Brazil (Rio de Janeiro, Santa Catarina). calatheae WIRTH, 1982a: 573 (larva, pupa, male, female; Dominica). Distr.- Colombia, Dominica, Brazil (Bahia, Santa Catarina). .

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marini SPINELLI & DIPPOLITO, 1995: 155 (male, female; Venezuela). Distr.- Venezuela. quatei WIRTH, 1952a: 142 (male, female; USA, California). Distr.- USA (California to Florida) to Brazil (Bahia). uramaensis SPINELLI & DIPPOLITO, 1995: 156 (female; Venezuela). Distr.- Venezuela. youngi WIRTH, 1982a: 579 (larva, pupa, male, female; Panama). Distr.- Costa Rica, Panama, Ecuador. fusca Species Group caliginosella WIRTH, 1974: 5. New name for caliginosa. Distr.- Southwestern Argentina. caliginosa INGRAM & MACFIE, 1931: 167 (preoccupied by Forcipomyia caliginosa (INGRAM & MACFIE), 1924). Argentina. fusca (PHILIPPI), 1865: 630 (Tetraphora; female; Chile). Distr.- Central and southern Chile south to Deceit Island, southwestern Argentina south to Tierra del Fuego. patagonica INGRAM & MACFIE, 1931: 165. Argentina. piroskyi CAVALIERI, 1961b: 172 (female; Argentina). Distr.- Argentina (Tierra del Fuego). wygodzinskyi CAVALIERI, 1961a: 17 (female; Argentina). Distr.- Argentina (Tierra del Fuego). delpontei CAVALIERI, 1961b: 169. Argentina. genualis Species Group genualis (LOEW), 1866: 128 (Ceratopogon; male; Cuba). Distr.- USA (Louisiana to Florida), Bahamas, Mexico (Yucatán, Chiapas) to Colombia, Galápagos Islands, Cayman Islands, Cuba to Argentina (Buenos Aires province); Sao Tomé. propinqua (WILLISTON), 1896: 279 (Ceratopogon). St. Vincent. raleighi MACFIE, 1938: 160. Trinidad. harpegonata WIRTH & SORIA, 1975: 19 (larva, pupa, male, female; Puerto Rico). Distr.Costa Rica, Bahamas, Jamaica, Puerto Rico, Trinidad to Brazil (Amazonas, Pará, Bahia). taragui MARINO, SPINELLI & CAZORLA, 2002: 7. New name for guarani. Argentina. Distr.Paraguay, Uruguay, Argentina (Corrientes, Misiones). guarani MARINO & SPINELLI, 1999: 448 (preoccupied by Forcipomyia guarani RONDEROS & SPINELLI, 1999). Argentina. poulaineae Species Group edwardsiana WIRTH, 1974: 5. New name for edwardsi. Distr.- Southwestern Argentina. edwardsi INGRAM & MACFIE, 1931: 160 (preoccupied by Forcipomyia edwardsi (SAUNDERS), 1925). Argentina. minitheca MARINO & SPINELLI, 2001a: 111 (male, female; Chile). Distr.- Southern Chile south to Deceit Island, southwestern Argentina. multipicta INGRAM & MACFIE, 1931: 157 (male, female; Argentina). Distr.- Southern Chile, southwestern Argentina. muzoni MARINO & SPINELLI, 2004a: 150 (male, female; Argentina). Distr.- Argentina (Rio Negro). nuncupata MACFIE, 1949: 110 (male, female; Mexico). Distr.- Mexico (Chiapas).

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ornatipennis MACFIE, 1939b: 151 (male, female; Brazil). Distr.- Costa Rica, Brazil (Santa Catarina). poulaineae INGRAM & MACFIE, 1931: 227 (male; Argentina). Distr.- Brazil (Bahia, Santa Catarina), Argentina (Buenos Aires). somuncurensis MARINO & SPINELLI, 2001a: 116 (male, female; Argentina). Distr.- Argentina (Rio Negro). townsendi KNAB, 1915: 111 (male, female; Peru). Distr.- Peru. utae KNAB, 1915: 109 (female; Peru). Distr.- Peru. yamana MARINO & SPINELLI, 2001a: 118 (female; Chile). Distr.- Chile (Deceit Island). zonogaster INGRAM & MACFIE, 1931: 168 (male; Chile). Distr.- Southern Chile. squamitibia Species Group catarinensis MARINO & SPINELLI, 2002: 309 (male, female; Brazil). Distr.- Mexico, Costa Rica, Colombia, Brazil. chilensis (PHILIPPI), 1865: 601 (Ceratopogon; male; Chile). Distr.- Central and southern Chile, southwestern Argentina south to Tierra del Fuego. pinamarensis SPINELLI, 1983a: 121 (male, female; Argentina). Distr.- Costa Rica, Panama, Venezuela, Brazil (Pará, Bahia, Rio de Janeiro), Uruguay, Argentina (Buenos Aires). quechua MARINO & SPINELLI, 2002: 312 (male, female; Argentina). Distr.- Argentina (Salta). rioplatensis MARINO & SPINELLI, 2002: 314 (male, female; Argentina). Distr.- Argentina (Buenos Aires), Uruguay. sexvittata WIRTH, 1956c: 248 (male, female; Costa Rica). Distr.- Costa Rica. spatuligera MACFIE, 1949: 111 (female; Mexico). Distr.- Mexico (Chiapas). squamitibia LUTZ, 1914: 88 (female; Brazil). Distr.- Colombia, Brazil (Rio de Janeiro, Santa Catarina), Puerto Rico, Trinidad. soriai WIRTH, 1991b: 168. Brazil. tenuisquamipes WIRTH, 1952b: 89 (female; Juan Fernández Islands). Distr.- Juan Fernández Islands. Unplaced to Species Group flava (WILLISTON), 1896: 280 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent. pictoni MACFIE, 1938: 161 (male; Trinidad). Distr.- USA (Louisiana to Florida) to Brazil (Amazonas, Pará, Bahia, São Paulo). Subgenus LASIOHELEA KIEFFER CENTRORHYNCHUS LUTZ, 1913: 62 (preoccupied by Centrorhynchus STEVEN or FISHER WALDHEIM, 1829). Type species: Centrorhynchus stylifer LUTZ, by original designation. LASIOHELEA KIEFFER, 1921c: 115. Type species: Atrichopogon pilosipennis KIEFFER (= Ceratopogon velox WINNERTZ), by original designation.

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REFERENCES: AITKEN et al., 1975 (species in Trinidad and Tobago); RONDEROS & SPINELLI, 1999 (revision of Neotropical species). anitae HUERTA & IBAÑEZ-BERNAL, 1996: 350 (male; Mexico). Distr.- Mexico (San Luis Potosi, Chiapas), Costa Rica, Brazil (Bahia, Itabuna), northeastern Argentina. attenuata SAUNDERS, 1964: 469 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica, Bolivia, Trinidad. cacaophila RONDEROS & SPINELLI, 1999: 152 (male; Venezuela). Distr.- Costa Rica, Venezuela. cornuta SAUNDERS, 1964: 464 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica, Trinidad, Brazil (São Paulo). guarani RONDEROS & SPINELLI, 1999: 153 (female; Paraguay). Distr.- Paraguay. insigniforceps MACFIE, 1939b: 165 (male; Brazil). Distr.- Brazil (Santa Catarina). intermedia SAUNDERS, 1964: 468 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica. multidentata RONDEROS & SPINELLI, 1999: 153 (female; Brazil). Brazil (Bahia, Itaguipe, Fazenda Almirante). quasicornuta SAUNDERS, 1964: 467 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica. saltensis (CAVALIERI), 1962: 360 (Lasiohelea; female; Argentina). Distr.- Northwestern Argentina. stylifer (LUTZ), 1913: 63 (Centrorhynchus; female; Brazil). Distr.- Belize to Ecuador, Trinidad, Venezuela to northeastern Argentina. Subgenus LEPIDOHELEA KIEFFER LEPIDOHELEA KIEFFER, 1917b: 364. Type species: Ceratopogon chrysolophus KIEFFER, by original designation. REFERENCES: WIRTH, 1991c (key to species groups; key to males and females of bicolor group); WIRTH & SPINELLI, 1992a (males and females of seminole subgroup); WIRTH & SPINELLI, 1993a (key to males and some females of annulatipes group). abercrombyi MACFIE, 1938: 161 (male, female; Trinidad). Distr.- Trinidad. annulatipes MACFIE, 1939b: 154 (male; Brazil). Distr.- Colombia, Brazil (Pará, Bahia, Santa Catarina). bahiensis WIRTH & SPINELLI, 1993a: 113 (male, female; Brazil). Distr.- Trinidad, Brazil (Bahia). basifemoralis WIRTH & SPINELLI, 1993a: 113 (male, female; Jamaica). Distr.- USA (Florida) to Trinidad, Panama. bicolor LUTZ, 1914: 89 (male; Brazil). Distr.- Costa Rica, Ecuador, Brazil (Rio de Janeiro, Santa Catarina). discoloripes MACFIE, 1939b: 159. Brazil. bifida WIRTH & SPINELLI, 1993a: 115 (male, female; Jamaica). Distr.- Jamaica. brasiliensis MACFIE, 1939b: 153 (male; Brazil). Distr.- Ecuador, Trinidad, Brazil (Santa Catarina). convexipenis WIRTH & SPINELLI, 1993a: 116 (male, female; Colombia). Distr.- Colombia. dubia MACFIE, 1939b: 162 (male; Brazil). Distr.- Brazil (Santa Catarina). edmistoni WIRTH & SPINELLI, 1993b: 624 (larva, pupa, male, female; USA, Maryland). Distr.- Eastern USA, Argentina (Buenos Aires). euthystyla WIRTH & SPINELLI, 1993a: 117 (male, female; Colombia). Distr.- Panama, Colombia.

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flavifemoris MACFIE, 1940d: 24 (male, female; Guyana). Distr.- Guyana. herediae WIRTH & SPINELLI, 1993a: 119 (male, female; Costa Rica). Distr.- Costa Rica. hobbsi WIRTH & SPINELLI, 1993a: 121 (male, female; Dominica). Distr.- Mexico (Tabasco) to Colombia, Jamaica to Grenada. kuanoskeles MACFIE, 1939b: 150 (male; Brazil). Distr.- Brazil (Santa Catarina). lacrimatorii MACFIE, 1939b: 161 (male, female; Brazil). Distr.- Brazil (Santa Catarina). luteigenua WIRTH & SPINELLI, 1992a: 353 (male, female; Costa Rica). Distr.- USA (Florida) to Colombia and Brazil (Bahia). seminole WIRTH, 1976: 81 (male, female; USA, Florida). Distr.- USA (Florida) to Brazil (Rio de Janeiro). squamithorax CLASTRIER, 1972: 170 (male, female; French Guiana). Distr.- French Guiana. varipennis WIRTH & WILLIAMS, 1957: 8 (female; Bermuda). Distr.- USA (Florida and Puerto Rico), Bermuda, Paraguay, northeastern Argentina. weemsi WIRTH & SPINELLI, 1993a: 123 (male, female; USA, Florida). Distr.- USA (Maryland to Florida, Arizona), Mexico (Baja California). winderi WIRTH, 1991b: 171 (male, female; Brazil). Distr.- Ecuador, Brazil (Bahia). Subgenus METAFORCIPOMYIA SAUNDERS METAFORCIPOMYIA SAUNDERS, 1957: 685 (as subgenus of Forcipomyia). Type species: Forcipomyia cerifera SAUNDERS, by original designation. REFERENCE: MARINO & SPINELLI, 2003 (key to males and females of New World species). cerifera SAUNDERS, 1957: 685 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro), northeastern Argentina. darwini MARINO & SPINELLI, 2003: 24 (male, female; Chile). Distr.- Chile (Deceit Island). maculosa INGRAM & MACFIE, 1931: 159 (female; Argentina). Distr.- Southwestern Argentina. mapuche MARINO & SPINELLI, 2003: 26 (male; Argentina). Distr.- Southwestern Argentina, southern Chile. morenoi MARINO & SPINELLI, 2003: 26 (male; Argentina). Distr.- Southwestern Argentina. williamsi MARINO & SPINELLI, 1999a: 5 (male, female; Argentina). Distr.- Argentina (Buenos Aires, Martin Garcia Island). Subgenus MICROHELEA KIEFFER MICROHELEA KIEFFER, 1917b: 364. Type species: Atrichopogon microtomus KIEFFER, designation by KIEFFER, 1921a: 7, with explanatory notation on erroneous previous type designation). PHASMIDOHELEA MAYER, 1937: 233. Type species: Phasmidohelea crudelis MAYER (= Ceratopogon fuliginosus MEIGEN), by original designation. REFERENCES: CLASTRIER & WIRTH, 1995 (key to females in ixodoides group); HUERTA, 1996 (key to males and females from Mexico); WIRTH, 1972 (key to females in fuliginosa group).

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alleni CLASTRIER & WIRTH, 1995: 117 (female; Costa Rica). Distr.- Costa Rica. amazonica WIRTH, 1971: 241 (female; Brazil). Distr.- Brazil (Amazonas). antioquiae CLASTRIER & WIRTH, 1995: 142 (female; Colombia). Distr.- Colombia. belemensis CLASTRIER & WIRTH, 1995: 140 (female; Brazil). Distr.- Colombia, Brazil (Pará). brasiliana CLASTRIER & WIRTH, 1995: 116 (female; Peru). Distr.- Peru. breelandi CLASTRIER & WIRTH, 1995: 114 (female; Panama). Distr.- Panama. brevilabellata CLASTRIER & WIRTH, 1995: 136 (female; French Guiana). Distr.- French Guiana. broadheadi CLASTRIER & WIRTH, 1995: 129 (female; Panama). Distr.- Panama. castneri CLASTRIER & WIRTH, 1995: 111 (female; Peru). Distr.- Peru, Brazil (Pará). catarina CLASTRIER & WIRTH, 1995: 131 (female; Brazil). Distr.- Brazil (Santa Catarina). coheni CLASTRIER & WIRTH, 1995: 132 (female; Ecuador). Distr.- Ecuador. colombiana CLASTRIER & WIRTH, 1995: 102 (female; Colombia). Distr.- Costa Rica, Colombia. cristata CLASTRIER & WIRTH, 1995: 135 (female; Brazil). Distr.- Brazil (Santa Catarina). desutterae CLASTRIER & WIRTH, 1995: 100 (female; French Guiana). Distr.- French Guiana, Brazil (Pará). donskoffi CLASTRIER & WIRTH, 1995: 125 (female; French Guiana). Distr.- French Guiana. dunklei CLASTRIER & WIRTH, 1995: 130 (female; Peru). Distr.- Peru. eriophora (WILLISTON), 1896: 279 (Ceratopogon; male; St. Vincent). Distr.- USA (Florida), Caribbean, Belize to Brazil (Bahia). felippebauerae CLASTRIER & WIRTH, 1995: 122 (female; Peru). Distr.- Peru. grandcolasi CLASTRIER & WIRTH, 1995: 112 (female; French Guiana). Distr.- French Guiana. franklini CLASTRIER & WIRTH, 1995: 141 (female; Costa Rica). Distr.- Costa Rica, Panama. fuliginosa (MEIGEN), 1818: 86 (Ceratopogon; Germany). Distr.- Widespread in the Palaearctic, Oriental, Afrotropical, Nearctic and Neotropical regions. In the Neotropics south to Argentina and Uruguay. crudelis KNAB, 1914: 66 (preoccupied by Forcipomyia crudelis (KARSCH), 1886). Mexico. tropica (KIEFFER), 1917b: 297 (Ceratopogon). Costa Rica. galapagensis (COQUILLETT), 1901b: 372 (Ceratopogon; male; Galápagos Islands). Distr.Galápagos Islands. guyana CLASTRIER & WIRTH, 1995: 124 (female; French Guiana). Distr.- French Guiana. insignipalpis MACFIE, 1949: 109 (female; Mexico). Distr.- Mexico (Chiapas), Costa Rica, Panama, Colombia, Brazil (Pará). iquitosensis CLASTRIER & WIRTH, 1995: 139 (female; Peru). Distr.- Peru. ixodoides (FIEBRIG-GERTZ), 1928: 290 (Ceratopogon; female; Paraguay). Distr.- Paraguay. kawensis CLASTRIER & WIRTH, 1995: 120 (female; French Guiana). Distr.- French Guiana, Peru. luteisquamosa WIRTH, 1972: 575 (female; Brazil). Distr.- Brazil (Santa Catarina). mayeri FORATTINI & LANE, 1955: 4. New name for crudelis. Distr.- Costa Rica. crudelis (MAYER), 1937: 233 (Phasmidohelea; female) (preoccupied by Forcipomyia crudelis (KARSCH), 1886). Costa Rica. menzeli CLASTRIER & WIRTH, 1995: 106 (female; Ecuador). Distr.- Ecuador. microtoma (KIEFFER), 1917b: 299 (Atrichopogon; female; Paraguay). Distr.- Belize and Dominica to Brazil (Amazonas, Santa Catarina), Paraguay. minisquamosa WIRTH, 1972: 570 (male, female; Belize). Distr.- Belize to Costa Rica, Brazil (Amazonas, Pará).

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moorei CLASTRIER & WIRTH, 1995: 133 (female; Brazil). Distr.- Brazil (Santa Catarina). neotropica CLASTRIER & WIRTH, 1995: 126 (female; Peru). Distr.- Peru. nigrimaxillata CLASTRIER & WIRTH, 1995: 118 (female; Peru). Distr.- Peru. novaeteutoniae CLASTRIER & WIRTH, 1995: 107 (female; Brazil). Distr.- Brazil (Santa Catarina). obesa COSTA LIMA, 1928: 85 (female; Brazil). Distr.- Brazil (Amazonas). penultimata WIRTH, 1972: 573 (male, female; Brazil). Distr.- Belize and Jamaica to Colombia, Brazil (Santa Catarina). peruviana CLASTRIER & WIRTH, 1995: 123 (female; Peru). Distr.- Peru. plaumanni CLASTRIER & WIRTH, 1995: 127 (female; Brazil). Distr.- Brazil (Santa Catarina). raposoensis CLASTRIER & WIRTH, 1995: 144 (female; Colombia). Distr.- Colombia. rettenmeyerorum CLASTRIER & WIRTH, 1995: 104 (female; Panama). Distr.- Panama. squamosa LUTZ, 1914: 87 (male, female; Brazil). Distr.- Colombia, Bolivia, Brazil (Amazonas, Bahia, Rio de Janeiro, Santa Catarina), Argentina (Mendoza). tettigonaris WIRTH & CASTNER, 1990: 159 (female; Peru). Distr.- Costa Rica, Dominican Republic, Peru. thomasi CLASTRIER & WIRTH, 1995: 115 (female; Brazil). Distr.- Brazil (Pará). valleensis CLASTRIER & WIRTH, 1995: 145 (female; Colombia). Distr.- Colombia. vernoni CLASTRIER & WIRTH, 1995: 138 (female; Colombia). Distr.- Colombia. wagneri (SÉGUY), 1941: 85 (Phasmidohelea; female; Brazil). Distr.- Brazil (Rio de Janeiro). willistoni WIRTH, 1971: 242 (male, female; Puerto Rico). Distr.- Jamaica, Puerto Rico, Colombia, Brazil (Santa Catarina). Subgenus PEDILOHELEA DE MEILLON & WIRTH PEDILOHELEA DE MEILLON & WIRTH, 1980: 9 (as subgenus of Forcipomyia). Type species: Forcipomyia clastrieri DESSART, by original designation. REFERENCE: DE MEILLON & WIRTH, 1980 (key to males, some females). aitkeni DE MEILLON & WIRTH, 1980: 12 (male, female; Brazil). Distr.- Colombia, Brazil (Pará). archboldi DE MEILLON & WIRTH, 1980: 13 (male, female; Dominica). Distr.- Jamaica, Puerto Rico, Brazil (Pará). raposoi DE MEILLON & WIRTH, 1980: 21 (male; Colombia). Distr.- Colombia. spangleri DE MEILLON & WIRTH, 1980: 21 (male, female; Guatemala). Distr.- Mexico (Yucatán), Guatemala, Panama, Colombia, Ecuador. spilmani DE MEILLON & WIRTH, 1980: 22 (male, female; Dominica). Distr.- Panama, Colombia, Dominica, Trinidad, Brazil (Santa Catarina). Subgenus PHYTOHELEA REMM PHYTOHELEA REMM, 1971: 189 (as subgenus of Forcipomyia). Type species: Ceratopogon bromelicola LUTZ, by original designation. REFERENCE: DE MEILLON & WIRTH, 1979 (key to females).

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antiguensis SAUNDERS, 1957: 700 (larva, pupa, male, female; Antigua). Distr.- Antigua. bromelicola (LUTZ), 1914: 84 (Ceratopogon; male, female; Brazil). Distr.- USA (Florida), Trinidad, French Guiana, Brazil (Rio de Janeiro). caribbeana SAUNDERS, 1957: 696 (larva, pupa, male, female; Trinidad and Tobago). Distr.Trinidad and Tobago, Guyana. dominicana DE MEILLON & WIRTH, 1979: 195 (larva, pupa, male, female; Dominica). Distr.Puerto Rico, Dominica. edwardsi (SAUNDERS), 1925: 260 (Apelma; larva, pupa, male, female; Brazil). Distr.- Trinidad, Brazil (Pernambuco, Rio de Janeiro). jocosa SAUNDERS, 1957: 701 (male, female; Trinidad). Distr.- Trinidad. keilini (SAUNDERS), 1925: 265 (Apelma; larva, pupa, male; Brazil). Distr.- Brazil (Pernambuco). magna (SAUNDERS), 1925: 266 (Apelma; larva, pupa, male, female; Brazil). Distr.- Brazil (Pernambuco). musae CLASTRIER & DELÉCOLLE, 1994: 51 (male, female; French Guiana). Distr.- French Guiana. oligarthra SAUNDERS, 1957: 698 (larva, pupa, male, female; Puerto Rico). Distr.- USA (Florida, Puerto Rico), Cuba, Dominica, Mexico, Guyana, also from Singapore and Micronesia. Subgenus PTEROBOSCA MACFIE PTEROBOSCA MACFIE, 1932: 266. Unavailable name; proposed after 1930 without type species designation. PTEROBOSCA MACFIE, 1940f: 16. Type species: Ceratopogon aeschnosuga DE MEIJERE, by original designation. farri WIRTH, 1966: 29 (female; Jamaica). Distr.- Jamaica. fusicornis (COQUILLETT), 1905: 63 (Ceratopogon; female; USA, Florida). Distr.- USA (Louisiana to Florida) and Caribbean to Brazil (Rio de Janeiro). fur (JOHNSON), 1913: 444 (Ceratopogon). Bermuda. macfiei (COSTA LIMA), 1937b: 616 (Pterobosca). Brazil. incubans (MACFIE), 1937b: 111 (Pterobosca; female; Belize). Distr.- Mexico (San Luis Potosi, Veracruz), Puerto Rico to Argentina (Buenos Aires). Subgenus RHYNCHOFORCIPOMYIA WIRTH & DOW RHYNCHOFORCIPOMYIA WIRTH & DOW, 1972: 863 (as subgenus of Forcipomyia). Type species: Forcipomyia messersmithi WIRTH & DOW, by original designation. REFERENCE: WIRTH & DOW, 1972 (key to females). brachyrhyncha WIRTH & DOW, 1972: 867 (male, female; Colombia). Distr.- Colombia, Trinidad. cylindrica WIRTH & DOW, 1972: 865 (male, female; El Salvador). Distr.- El Salvador. dorsalis WIRTH & DOW, 1972: 870 (male, female; Mexico). Distr.- Mexico (Sinaloa, Veracruz), Honduras, El Salvador.

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guamai WIRTH & DOW, 1972: 869 (female; Brazil). Distr.- Panama, Brazil (Pará). messersmithi WIRTH & DOW, 1972: 864 (male, female; El Salvador). Distr.- Mexico (Veracruz), Central America to Colombia, Jamaica. puracensis WIRTH & DOW, 1972: 867 (female; Colombia). Distr.- Colombia. zeteki WIRTH & DOW, 1972: 870 (female; Panama). Distr.- Panama, Colombia. Subgenus SALIOHELEA WIRTH & RATANAWORABHAN SALIOHELEA WIRTH & RATANAWORABHAN, 1978: 494 (as subgenus of Forcipomyia). Type species: Forcipomyia leei WIRTH & RATANAWORABHAN, by original designation. leei WIRTH & RATANAWORABHAN, 1978: 498 (male, female; Colombia). Distr.- Eastern USA and Caribbean to southern Brazil (Amazonas, Pará, Bahia, Rio de Janeiro). Subgenus SCHIZOFORCIPOMYIA CHAN & LEROUX SCHIZOFORCIPOMYIA CHAN & LEROUX, 1971: 271 (as subgenus of Forcipomyia). Type species: Forcipomyia petersoni CHAN & LEROUX ( = Forcipomyia borbonica CLASTRIER), by original designation. harpa SPINELLI & BORKENT, 2004b: 3 (male, female; Costa Rica). Distr.- Costa Rica. Subgenus SYNTHYRIDOMYIA SAUNDERS SYNTHYRIDOMYIA SAUNDERS, 1957: 688 (as subgenus of Forcipomyia). Type species: Lasiohelea acidicola TOKUNAGA, by original designation. floridensis DOW & WIRTH, 1972: 195 (male, female; USA, Florida). Distr.- USA (Florida), Mexico (Yucatán). sanctaeclarae WIRTH, 1952b: 90 (male, female; Juan Fernández Islands). Distr.- Juan Fernández Islands, central Chile, northwestern Argentina. soibelzoni MARINO & SPINELLI, 2001c: 14 (female; Argentina). Distr.- Argentina (Rio Negro). tenuiforceps MACFIE, 1939b: 167 (male; Brazil). Distr.- USA (California, New Mexico) to Brazil (Santa Catarina). Subgenus THYRIDOMYIA SAUNDERS THYRIDOMYIA SAUNDERS, 1925: 268. Type species: Thyridomyia palustris SAUNDERS (= Ceratopogon monilicornis COQUILLETT), by original designation. REFERENCE: WIRTH, 1970 (key to males and females). calchaqui SPINELLI & MARINO, 1997: 188 (female; Argentina). Distr.- Northwestern Argentina.

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colombiae WIRTH, 1970: 435 (male, female; Colombia). Distr.- Mexico (Veracruz), Costa Rica, Panama, Trinidad, Colombia. jamaicensis WIRTH, 1970: 436 (male, female; Jamaica). Distr.- Jamaica, Puerto Rico. jipajapae WIRTH, 1970: 437 (male, female; Panama). Distr.- Costa Rica to Ecuador, Trinidad. nana (MACFIE), 1939b: 171 (Lasiohelea; female; Brazil). Distr.- Mexico (Veracruz), Jamaica through Caribbean to southern Brazil (Santa Catarina). nodosa SAUNDERS, 1959: 43 (larva, pupa, male, female; Costa Rica). Distr.- USA (Arizona, Kansas, Mississippi), Mexico (Baja California, Sonora, Sinaloa, San Luis Potosi), Costa Rica, Colombia. riojana SPINELLI & MARINO, 1997: 188 (male; Argentina). Distr.- Western and southern Argentina. sinuosa DOW & WIRTH, 1972: 189 (male, female; Mexico). Distr.- Mexico (Sinaloa) (possibly Iowa in USA). univesicula MACFIE, 1939b: 170 (female; Brazil). Distr.- Brazil (Santa Catarina). Subgenus TRICHOHELEA GOETGHEBUER APELMA KIEFFER, 1919a: 64 (preoccupied by BILLBERG, 1820). Type species: Apelma auronitens KIEFFER (= Trichohelea tonnoiri GOETGHEBUER), designation by MACFIE, 1940f: 16. TRICHOHELEA GOETGHEBUER, 1920: 39. Type species: Trichohelea tonnoiri GOETGHEBUER, by original designation. REFERENCE: MARINO & SPINELLI, 2004b (key to males and females). aeronautica MACFIE, 1935b: 265 (female; Guyana). Distr.- Venezuela to French Guiana. baueri WIRTH, 1956a: 361 (female; USA, Arizona). Distr.- USA (Florida, Maryland, Arizona), Mexico (Baja California, Sonora, Sinaloa, Veracruz). danaisi (FLOCH & ABONNENC), 1949: 2 (1950a: 72) (Lasiohelea; female; Venezuela). Distr.Venezuela. intrepida MACFIE, 1936: 228 (female; Peru). Distr.- Peru. limnetis INGRAM & MACFIE, 1931: 169 (male; Argentina). Distr.- Southwestern Argentina. shannoni (INGRAM & MACFIE), 1931: 171 (Lasiohelea). Argentina. macheti CLASTRIER & LEGRAND, 1990: 168 (female; French Guiana). Distr.- French Guiana. opilionivora (LANE), 1947c: 159 (Lasiohelea; female; Brazil). Distr.- Brazil (São Paulo). roubaudi CLASTRIER & DELÉCOLLE, 1997: 379 (female; French Guiana). Distr.- French Guiana. sayhuequei MARINO & SPINELLI, 2004b: 2257 (female; Argentina). Distr.- Southwestern Argentina, southern Chile. tehuelche MARINO & SPINELLI, 2004b: 2260 (male, female; Argentina). Distr.- Southwestern Argentina. trinidadensis SAUNDERS, 1964: 476 (pupa, female; Trinidad). Distr.- Trinidad. Subgenus WARMKEA SAUNDERS WARMKEA SAUNDERS, 1957: 671 (as subgenus of Forcipomyia). Type species: Forcipomyia bicolor SAUNDERS (= Forcipomyia lesliei WIRTH), by original designation.

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REFERENCE: WIRTH & SORIA, 1980 (key to males and females). aeria SAUNDERS, 1957: 675 (larva, pupa, male, female; Puerto Rico). Distr.- Puerto Rico, Colombia, Argentina (Tucuman). galindoi WIRTH & SORIA, 1980: 146 (male, female; Panama). Distr.- Panama, Colombia, Ecuador, Brazil (Pará). lesliei WIRTH, 1974: 12. New name for bicolor. Distr.- Jamaica, Haiti to Venezuela, Colombia. bicolor SAUNDERS, 1957: 672 (preoccupied by Forcipomyia bicolor LUTZ, 1914). Puerto Rico. louriei (MACFIE), 1935a: 49 (Lasiohelea; female; Brazil). Distr.- Colombia, Bolivia, Trinidad, Brazil (Pará, Maranhão, Santa Catarina). narthekophora MACFIE, 1939b: 166. Brazil. spinosa SAUNDERS, 1957: 676 (larva, pupa, male, female; Puerto Rico). Distr.- Puerto Rico, Dominica, Trinidad to Colombia, Guyana, and Brazil (Pará, Bahia, Rio de Janeiro). terrestris SAUNDERS, 1964: 479 (larva, pupa, male, female; Trinidad). Distr.- Trinidad. tuberculata SAUNDERS, 1957: 677 (larva, pupa, male, female; Trinidad and Tobago). Distr.Costa Rica, Colombia, Trinidad and Tobago, Brazil (Amazonas, Pará), Argentina (Buenos Aires). SUBFAMILY DASYHELEINAE LENZ, 1934: 96 Genus DASYHELEA KIEFFER DASYHELEA KIEFFER, 1911c: 5. Type species: Dasyhelea halophila KIEFFER, by monotypy. PSEUDOCULICOIDES M ALLOCH , 1915a: 309. Type species: Ceratopogon mutabilis COQUILLETT, by original designation. REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE, 1931 (key males and females from Patagonia and southern Chile); SPINELLI & WIRTH, 1984d (key to males and females of cincta group). aegealitis SPINELLI & WIRTH, 1984d: 589 (male, female; Jamaica). Distr.- USA (Florida), Jamaica. albopicta INGRAM & MACFIE, 1931: 187 (male, female; Argentina). Distr.- Southern Chile, southwestern Argentina. ancora (COQUILLETT), 1902: 87 (Ceratopogon; female; USA, Florida). Distr.- USA (California, Arizona, New Mexico, Connecticut to Florida), Galápagos Islands. andensis INGRAM & MACFIE, 1931: 180 (male; Argentina). Distr.- Southwestern Argentina. argentinensis KIEFFER, 1925: 92 (male; Argentina). Distr.- Argentina (Cordoba). atlantis WIRTH & WILLIAMS, 1957: 11 (male, female; Bermuda). Distr.- Bermuda, USA (New York, Florida), Grand Cayman, Galápagos Islands. australis WIRTH, 1952b: 92 (male, female; Juan Fernandez Islands). Distr.- Juan Fernandez Islands. bahamensis (JOHNSON), 1908: 71 (Ceratopogon; female; Bahamas). Distr.- USA (Florida, Puerto Rico), Mexico (Yucatán), Bahamas, Cuba, Jamaica. bajensis WIRTH, 1978: 192 (male, female; Mexico). Distr.- USA (California), Mexico (Baja California, Sinaloa).

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bermudae WIRTH & WILLIAMS, 1957: 11 (male, female; Bermuda). Distr.- Bermuda. borgmeieri WIRTH & WAUGH, 1976: 225 (male, female; Trinidad). Distr.- Panama, Grenada, Trinidad, Galápagos Islands, Brazil (Bahia). cacaoi WIRTH & WAUGH, 1976: 231 (male, female; Trinidad). Distr.- Panama, Trinidad. calvescens MACFIE, 1938: 157 (male, female; USA, Hawaii). Distr.- Mexico (Baja California, Sonora), Panama, Hawaii. caribbeana SPINELLI & WIRTH, 1984d: 596 (female; Jamaica). Distr.- Mexico (Quintana Roo, Chiapas), Panama, Jamaica, Haiti. chilensis INGRAM & MACFIE, 1931: 186 (female; Chile). Distr.- Southern Chile cincta (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Florida). Distr.- USA (California to New York to Florida) to Brazil and Argentina. penthesileae MACFIE, 1935a: 55. Brazil. columbiana (KIEFFER), 1917b: 304 (Culicoides; female; Colombia). Distr.- Colombia. correntina RONDEROS & DÍAZ, in RONDEROS et al., 2004a: 194 (male, female, pupa; Argentina). Distr.- Northeastern Argentina. filibranchia (LUTZ), 1914: 85 (Ceratopogon; pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro). flavicauda MACFIE, 1939b: 201 (female; Brazil). Distr.- Northeastern Argentina, Brazil (Santa Catarina). griseola WIRTH, 1978: 193 (male, female; Mexico). Distr.- Mexico (Baja California), Panama, Trinidad. guadeloupensis DELÉCOLLE & RIEB, 1994: 272 (male, female; Guadeloupe). Distr.Guadeloupe. hippolytae MACFIE, 1935a: 54 (female; Brazil). Distr.- Brazil (Maranhão). hirtipes (KIEFFER), 1917b: 305 (Culicoides; male; Peru). Distr.- Peru. hondurensis SPINELLI & WIRTH, 1984d: 600 (male, female; Belize). Distr.- Belize. jamaicensis SPINELLI & WIRTH, 1984d: 602 (female; Jamaica). Distr.- Jamaica. koenigi DELÉCOLLE & RIEB, 1994: 267 (male, female; Guadeloupe). Distr.- Guadeloupe. lacustris INGRAM & MACFIE, 1931: 190 (female; Argentina). Distr.- Western Argentina. luteogrisea WIRTH & WILLIAMS, 1957: 10 (female; Bermuda). Distr.- USA (Florida to Texas), Bermuda, Bahamas. maculata MACFIE, 1943: 119 (male, female; Bahamas). Distr.- USA (Florida), Mexico (Quintana Roo) to Panama, Bahamas, Jamaica to Trinidad. mediomunda MINAYA, 1978: 79 (pupa, male, female; Peru). Distr.- Peru, Chile. monticola INGRAM & MACFIE, 1931: 188 (male, female; Chile). Distr.- Southern Chile. morrisoni Grogan and Wieners, 2006: 468 (male, female; Bahamas). Distr.- Bahamas. mutabilis (COQUILLETT), 1901a: 602 (Ceratopogon; male, female; USA, District of Columbia). Distr.- North America (Alaska to Newfoundland, to California to Florida), Galápagos Islands. necrophila SPINELLI & RODRÍGUEZ, 1999: 59 (male, female, pupa, larva; Argentina). Distr.Mexico (Coahuila), Argentina (Buenos Aires). paracincta WIRTH, 1969: 576 (male, female; Galápagos Islands). Distr.- Galápagos Islands. parahybae MACFIE, 1940a: 74 (male, female; Brazil). Distr.- Northeastern Brazil. patagonica INGRAM & MACFIE, 1931: 182 (male, female; Argentina). Distr.- Southwestern Argentina.

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paulistana FORATTINI & RABELLO, 1957: 245 (pupa, female; Brazil). Distr.- Brazil (São Paulo). philotherma MACFIE, 1953: 103 (male; Costa Rica). Distr.- Costa Rica. pollex BORKENT & FORSTER, 1986: 1286 (male; Bahamas). Distr.- USA (Florida), Bahamas, Mexico (Baja California). pumila MACFIE, 1939b: 202 (male, female; Brazil). Distr.- Brazil (Santa Catarina). pusilla (LUTZ), 1913: 65 (Centrorhynchus; female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro). pygmaea (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- St. Vincent. pyrsonota MACFIE, 1953: 104 (male; Costa Rica). Distr.- Costa Rica. reynoldsi INGRAM & MACFIE, 1931: 185 (male, female; Chile). Distr.- Chile (Magallanes, Tierra del Fuego). santaemarthae (KIEFFER), 1917b: 306 (Culicoides; female; Colombia). Distr.- Colombia. scissurae MACFIE, 1937a: 15 (male; Trinidad). Distr.- USA (Florida), Mexico (San Luis Potosi, Guerrero) to Costa Rica, Bermuda, Trinidad to northeastern Argentina. shannoni INGRAM & MACFIE, 1931: 183 (male, female; Chile). Distr.- Southern Chile. sinclairi BORKENT, 1991: 110 (male, female; Galápagos Islands). Distr.- Galápagos Islands. soriai WIRTH & WAUGH, 1976: 229 (male, female; Brazil). Distr.- Brazil (Bahia). spathicerca WIRTH, 1969: 577 (male, female; Galápagos Islands). Distr.- Galápagos Islands. suarezi SPINELLI & RONDEROS, 1987: 11 (male, female; Argentina). Distr.- Argentina (Buenos Aires). thalestris MACFIE, 1935a: 55 (female; Brazil). Distr.- Trinidad, Guyana, Brazil (Maranhão). villosipes (KIEFFER), 1917b: 305 (Culicoides; male; Paraguay). Distr.- Paraguay. viridans FORATTINI & RABELLO, 1957: 247 (male; Brazil). Distr.- Brazil (Rio de Janeiro). williamsi WIRTH & WAUGH, 1976: 228 (male, female; Trinidad). Distr.- Panama, Dominica, Trinidad, Brazil (Bahia). winderi WIRTH & WAUGH, 1976: 233 (male, female; Brazil). Distr.- Brazil (Bahia). SUBFAMILY CERATOPOGONINAE NEWMAN, 1834: 388 TRIBE CULICOIDINI KIEFFER, 1911c: 1, 1911b: 319 Genus CULICOIDES LATREILLE CULICOIDES L ATREILLE , 1809: 251. Type species: Culicoides punctatus L ATREILLE (= Ceratopogon punctatus MEIGEN), by monotypy. PADROSIA RAFINESQUE, 1815: 130 (unnecessary replacement name for Culicoides LATREILLE). Type species: Culicoides punctatus LATREILLE (= Ceratopogon punctatus MEIGEN), automatic. REFERENCES: AITKEN et al., 1975 (key to females from Trinidad and Tobago); BLANTON & WIRTH, 1979 (key to larvae, males and females from Florida (USA)); FORATTINI, 1957 (key to females of all species); HUERTA, 1996 (key to females from Mexico); ORTÍZ & LEÓN, 1955 (key to females from Ecuador); RONDEROS, 1988 (key to females from Argentina); RONDEROS & SPINELLI, 1998 (key to males and females from the Yacyretá Dam, between Paraguay and Argentina); SPINELLI, RONDEROS, DÍAZ & MARINO, 2005 (key to males and females from Argentina); SPINELLI & WIRTH, 1986b (key to

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females south of the Amazon Basin); WIRTH & BLANTON, 1959 (key to females and some males from Panama); WIRTH & BLANTON 1973 (key to females and some males from Amazon Basin); WIRTH & BLANTON, 1974b (key to females and some males from the Caribbean); WIRTH, DYCE & SPINELLI, 1988 (some meristic character states and photographs of wings of all species). There is no reliable key to the subgenera and species groups of Neotropical Culicoides. For species outside of local areas for which revisions are available (as listed above), the easiest guide to species identification is that of WIRTH, DYCE & SPINELLI (1988). From there readers may refer to keys to subgenera or species groups referred to below. Subgenus AMOSSOVIA GLUKHOVA AMOSSOVIA GLUKHOVA, 1989: 226 (as subgenus of Culicoides). Type species: Ceratopogon dendrophilus Amosova, by original designation. REFERENCE: WIRTH & BLANTON, 1967 (key to males and females) oklahomensis KHALAF, 1952: 355 (as subspecies of villosipennis; male; USA, Oklahoma). Distr.- USA (California to Mississippi) to Guatemala. Subgenus ANILOMYIA VARGAS ANILOMYIA VARGAS, 1960: 37 (as subgenus of Culicoides). Type species: Culicoides covagarciai ORTÍZ, by original designation. REFERENCES: WIRTH & BLANTON, 1956b (key to males and females of covagarciai group); WIRTH & BLANTON, 1970d (key to key to males and females of nigrigenus group). ameliae BROWNE, 1980: 543 (female; Colombia). Distr.- Colombia. chaverrii SPINELLI & BORKENT, 2004a: 373 (larva, pupa, male, female; Costa Rica). Distr.Costa Rica. chrysonotus WIRTH & BLANTON, 1956b: 226 (male, female; Panama). Distr.- El Salvador, Costa Rica, Panama. covagarciai ORTÍZ, 1950b: 457 (male, female; Venezuela). Distr.- Honduras to Colombia, Venezuela. beebei FOX, 1952: 366. Venezuela. decor (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- Dominica, St. Lucia, St. Vincent. dominicanus WIRTH & BLANTON, 1970d: 146 (male, female; Dominica). Distr.- Dominica. efferus FOX, 1952: 365 (female; Peru). Distr.- Guatemala to Peru and Bolivia. farri WIRTH & BLANTON, 1970d: 148 (male, female; Jamaica). Distr.- Jamaica. hayesi MATTA, 1967: 75 (larva, pupa, male, female; Honduras). Distr.- Mexico (Tamaulipas), Honduras.

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lutealaris WIRTH & BLANTON, 1956b: 225 (male, female; Panama). Distr.- Costa Rica, Panama. marshi WIRTH & BLANTON, 1956b: 220 (female; Panama). Distr.- Costa Rica, Panama, Colombia. metagonatus WIRTH & BLANTON, 1956b: 221 (male, female, Panama). Distr.- Nicaragua to Ecuador. monicae SPINELLI & BORKENT, 2004a: 372 (female; Costa Rica). Distr.- Costa Rica. nigrigenus WIRTH & BLANTON, 1956b: 222 (male, female; Panama). Distr.- Mexico (Veracruz) to Colombia, Trinidad, Argentina (Salta). popayanensis WIRTH & LEE, 1967: 10 (male, female; Colombia). Distr.- Colombia. rostratus WIRTH & BLANTON, 1956b: 218 (male, female; Panama). Distr.- Panama. trapidoi WIRTH & BARRETO, 1978: 554 (male, female; Colombia). Distr.- Costa Rica, Colombia, Brazil. Subgenus AVARITIA FOX AVARITIA FOX, 1955: 218 (as subgenus of Culicoides). Type species: Ceratopogon obsoletus MEIGEN, by original designation. REFERENCES: RODRÍGUEZ & WIRTH, 1986 (revision of andicola group); WIRTH & MULLENS, 1992 (key to males and females in pusillus group). andicola WIRTH & LEE, 1967: 5 (female; Colombia). Distr.- Colombia. hermani SPINELLI & BORKENT, 2004a: 363 (male, female; Panama). Distri. - Costa Rica, Panama. impusilloides SPINELLI & WIRTH, 1984a: 178 (male, female; Brazil). Distr.- Brazil (Santa Catarina), Uruguay. orjuelai WIRTH & LEE, 1967: 6 (female; Colombia). Distr.- Colombia. puracensis WIRTH & LEE, 1967: 7 (male, female; Colombia). Distr.- Colombia. pusilloides WIRTH & BLANTON, 1955a: 104 (male, female; Panama). Distr.- Guatemala and Belize to Panama. pusillus LUTZ, 1913: 52 (male, female; Brazil). Distr.- USA (Florida), Mexico (Chiapas) to northeastern Argentina. suarezi RODRÍGUEZ & WIRTH, 1986: 313 (female; Colombia). Distr.- Colombia. Subgenus BELTRANMYIA VARGAS BELTRANMYIA VARGAS, 1953a: 34 (as subgenus of Culicoides). Type species: Culicoides crepuscularis MALLOCH, by original designation. REFERENCE: BLANTON & WIRTH, 1979 (males and females included in key to species from Florida). bermudensis WILLIAMS, 1956: 298 (female; Bermuda). Distr.- USA (New York to Florida and Texas), Bermuda. crepuscularis MALLOCH, 1915a: 303 (male, female; USA, Illinois). Distr.- Canada (British Columbia to Nova Scotia) to Costa Rica. knowltoni BECK, 1956: 136 (male, female; USA, Florida). Distr.- Bahamas, USA (Florida).

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Subgenus COTOCRIPUS BRÈTHES COTOCRIPUS BRÈTHES, 1912: 451. Type species: Cotocripus caridei BRÈTHES, by monotypy. bambusicola LUTZ, 1913: 62 (female; Brazil). Distr.- Colombia, Venezuela, Brazil (Bahia, Espírito Santo, Rio de Janeiro, São Paulo), Argentina (Misiones, Buenos Aires). bahiensis BARBOSA, 1947: 11. Brazil. caridei (BRÈTHES), 1912: 451 (Cotocripus; female; Argentina). Distr.- Argentina (Buenos Aires province), southeastern Brazil, Uruguay. setifer (LUTZ), 1913: 64 (Centrorhynchus). Brazil. irwini SPINELLI & WIRTH, 1984a: 180 (female; Chile). Distr.- Chile, southwestern Argentina. patagoniensis RONDEROS & SPINELLI, 1997: 34 (male, female; Argentina). Distr.- Southern Chile, southern Argentina. raposoensis WIRTH & BARRETO, 1978: 561 (male, female; Colombia). Distr.- Colombia. Subgenus CULICOIDES LATREILLE elutus MACFIE, 1948: 75 (female; Mexico). Distr.- Mexico (Chiapas, Oaxaca) to Panama. hondurensis SPINELLI & BORKENT, 2004a: 369 (male, female; Honduras). Distr.- El Salvador, Honduras. luteovenus ROOT & HOFFMAN, 1937: 156 (male, female; Mexico). Distr.- USA (California, Utah, Washington) to Panama. neopulicaris WIRTH, 1955: 355 (male, female; USA,Texas). Distr.- USA (Texas and Louisiana) to Costa Rica. Subgenus DIPHAOMYIA VARGAS DIPHAOMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides baueri HOFFMAN, by original designation. blantoni VARGAS & WIRTH, 1955: 33 (male, female; Mexico). Distr.- USA (Texas), Mexico (Sinaloa, Tamaulipas, Morelos). edeni WIRTH & BLANTON, 1974a: 23 (male, female; notes, syn.; USA, Florida). Distr.Southeastern USA to Bahamas. evansi WIRTH & BLANTON, 1959: 342 (male, female; Panama). Distr.- Honduras, Costa Rica, Panama. freitasi WIRTH & BLANTON, 1973: 434 (female; Brazil). Distr.- Brazil (Pará). haematopotus MALLOCH, 1915a: 302 (male, female; USA, Illinois). Distr.- North America (British Columbia to Nova Scotia, entire USA) to Honduras. iriartei FOX, 1952: 368 (female; Venezuela). Distr.- Guatemala to Colombia, Venezuela, Tobagao, Brazil (Pará). vargasi WIRTH & BLANTON, 1953a: 74. Panama. jurbergi FELIPPE-BAUER, in FELIPPE-BAUER et al., 2005: 51 (female; Peru). Distr.- Peru. marinkellei WIRTH & LEE, 1967: 13 (male, female; Colombia). Distr.- Colombia.

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mckeeveri BRICKLE & HAGAN, 1999: 39 (female; Belize). Distr.- Belize. minasensis FELIPPE-BAUER, 1987: 147 (female; Brazil). Distr.- Brazil (Minas Gerais). mirsae ORTÍZ, 1953a: 801 (female; Venezuela). Distr.- Panama, Venezuela, Trinidad. ronderosae SPINELLI & BORKENT, 2004a: 377 (female; Costa Rica). Distr.- Costa Rica. tarapaca SPINELLI & WIRTH, 1984a: 184 (female; Chile). Distr.- Northern Chile. Subgenus DRYMODESMYIA VARGAS DRYMODESMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides copiosus ROOT & HOFFMAN, by original designation. REFERENCE: WIRTH & HUBERT, 1960: 646 (key to females and some males; as copiosus group). borinqueni FOX & HOFFMAN, 1944: 110 (male, female; Puerto Rico). Distr.- Jamaica to St. Croix. bredini WIRTH & BLANTON, 1970b: 41 (male, female; Dominica). Distr.- Dominica. chacoensis SPINELLI & WIRTH, 1984a: 174 (male, female; Argentina). Distr.- Bolivia, Paraguay, Argentina, Uruguay. haitiensis DELÉCOLLE, RACCURT & REBHOLTZ, 1986: 108 (male, female; Haiti). Distr.- Haiti. hitchcocki SPINELLI & WIRTH, 1984a: 176 (female; Peru). Distr.- Peru. jamaicensis EDWARDS, 1922: 165 (as variety of loughnani; female; Jamaica). Distr.- USA (Texas, Florida), Mexico (Yucatán), Central America and Caribbean to Colombia and Venezuela. loughnani EDWARDS, 1922: 165 (female; Jamaica). Distr.- USA (Texas to Georgia), Mexico (Yucatán), Bahamas, Jamaica, Cuba, Australia (as introduction with Opuntia). panamensis BARBOSA, 1947: 22 (male, female; Panama). Distr.- Mexico (Nayarit, Veracruz, Chiapas) to Costa Rica, Jamaica. alambiculorum MACFIE, 1948: 81. Mexico. pilosus WIRTH & BLANTON, 1959: 332 (male, female; Panama). Distr.- Costa Rica, Panama, Brazil (Pará). poikilonotus MACFIE, 1948: 82 (female; Mexico). Distr.- Mexico (Chiapas), Central America, Venezuela and Trinidad to Brazil (Bahia). cacozelus MACFIE, 1948: 85. Mexico. hertigi WIRTH & BLANTON, 1953b: 229. Panama. saltaensis SPINELLI & WIRTH, 1984a: 183 (female; Argentina). Distr.- Western Argentina. uruguayensis RONDEROS, 1990a: 117 (female; Uruguay). Distr.- Uruguay. wirthomyia VARGAS, 1953b: 227 (male; Mexico). Distr.- Mexico (Guerrero). Subgenus GLAPHIROMYIA VARGAS GLAPHIROMYIA VARGAS, 1960: 41 (as subgenus of Culicoides). Type species: Culicoides scopus ROOT & HOFFMAN, by original designation. parascopus WIRTH & BLANTON, 1978: 238 (male, female; Mexico). Distr.- Mexico (Michoacan). scopus ROOT & HOFFMAN, 1937: 170 (male, female; Mexico). Distr.- Mexico (DF), Costa Rica, Panama.

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Subgenus HAEMATOMYIDIUM GOELDI HAEMATOMYIDIUM GOELDI, 1905: 137. Type species: Haematomyidium paraense GOELDI, by original designation. REFERENCES: VITALE, WIRTH & AITKEN, 1981 (key to females of debilipalpis group); WIRTH & FELIPPE-BAUER, 1989 (key to paraensis group). annuliductus WIRTH, in VITALE et al, 1981: 150 (pupa, male, female; Panama). Distr.- Costa Rica, Panama. austroparaensis SPINELLI, in SPINELLI et al., 2005b: 141 (male, female; Argentina). Distr.northeastern Argentina. aragaoi TAVARES & LUNA DIAS, 1980: 393 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro). bachmanni SPINELLI, in SPINELLI et al, 2005b: 146 (female; Argentina). Distr.- Paraguay, northeastern Argentina. bayano WIRTH, in VITALE et al, 1981: 152 (pupa, male, female; Panama). Distr.- Costa Rica, Panama. crucifer CLASTRIER, 1968: 85 (male, female; French Guiana). Distr.- Trinidad, Guyana, French Guiana. darlingtonae WIRTH & BLANTON, 1971: 39 (male, female; Trinidad). Distr.- Costa Rica, Trinidad. debilipalpis LUTZ, 1913: 60 (female; Brazil). Distr.- Widespread from USA (Maryland, Kentucky, Nebraska south to Lousiana and Florida), Guatemala and Belize to Argentina. ichesi RONDEROS & SPINELLI, 1995b: 77. Argentina. denisae CLASTRIER, 1971: 290 (male, female; French Guiana). Distr.- French Guiana, Brazil (Amazonas). diversus FELIPPE-BAUER, in FELIPPE-BAUER et al. 2003: 1052 (female; Peru). Distr.- Peru. dureti RONDEROS & SPINELLI, 1995a: 59 (female; Paraguay). Distr.- Northeastern Argentina, Paraguay. eadsi WIRTH & BLANTON, 1971: 37 (male, female; USA, Texas). Distr.- USA (Florida, Texas), Cuba, Mexico (Nayarit, Sonora, San Luis Potosi, Yucatán), Guatemala. eldridgei WIRTH & BARRETO, 1978: 561 (male, female; Colombia). Distr.- Colombia. equatoriensis BARBOSA, 1952: 13 (as variety of debilipalpis; female; Ecuador). Distr.- Ecuador. espinolai FELIPPE-BAUER & LOURENCO-DE-OLIVEIRA, 1987: 149 (female; Brazil). Distr.- Brazil (Minas Gerais). filiductus WIRTH, in VITALE et al., 1981: 155 (pupa, male, female; Panama). Distr.- Belize to Panama. flinti WIRTH, 1982b: 251 (female; Argentina). Distr.- Northeastern Argentina, southeastern Brazil, Uruguay. germanus MACFIE, 1940d: 27 (female; Guyana). Distr.- Costa Rica, Guyana. ginesi ORTÍZ, 1951c: 586 (female; Venezuela). Distr.- El Salvador to Panama, Colombia, Venezuela, Trinidad, Brazil (Pará), northeastern Argentina. glabrior MACFIE, 1940d: 27 (as variety of debilipalpis; female; Guyana). Distr.- Honduras to Ecuador, Guyana, Surinam, Trinidad, Brazil (Pará). grahambelli FORATTINI, 1956b: 35. Panama.

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hoffmani FOX, 1946: 251 (female; Trinidad). Distr.- Belize, Costa Rica, Cayman Islands, Jamaica to Trinidad. horticola LUTZ, 1913: 61 (female; Brazil). Distr.- Paraguay, eastern Argentina, southeastern Brazil. imitator ORTÍZ, 1953b: 807 (male, female; Venezuela). Distr.- Guatemala to Panama, Venezuela. insinuatus ORTÍZ & LEÓN, 1955: 577 (female; Ecuador). Distr.- Colombia, Ecuador, Peru, Trinidad, Guyana, Brazil (Amazonas, Pará, (?) São Paulo). kampa FELIPPE-BAUER, VERAS, CASTELLON & MOREIRA, 2000: 35 (female; Brazil). Distr.Brazil (Acre). lahillei (ICHES), 1906: 264 (Ceratopogon, female; Argentina). Distr.- Northeastern Argentina, southeastern Brazil, Paraguay. limonensis ORTÍZ & LEÓN, 1955: 576 (female; Ecuador). Distr.- Ecuador, Venezuela, Brazil (Pará, São Paulo, Santa Catarina). neoparaensis TAVARES & SOUZA, 1978: 621 (male, female; Brazil). Distr.- Ecuador, Brazil (Amazonas, Rio de Janeiro, Santa Catarina). pampaensis SPINELLI & WIRTH, 1984a: 182 (female; Argentina). Distr.- Western Argentina. paraensis (GOELDI), 1905: 137 (Haematomyidium; female; Brazil). Distr.- USA (Colorado, Nebraska, Pennsylvania, Wisconsin to Louisiana and Florida) to Argentina. undecimpunctatus KIEFFER, 1917b: 307. Argentina. peruvianus FELIPPE-BAUER, in FELIPPE-BAUER et al., 2003: 1054 (female; Peru). Distr.- Peru. quasiparaensis CLASTRIER, 1971: 286 (male, female; French Guiana). Distr.- Honduras and El Salvador to Colombia, Peru, French Guiana, Brazil (Amazonas, Rondônia). rachoui TAVARES & SOUZA, 1978: 622 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro). spurius WIRTH & BLANTON, 1959: 433 (male, female; Panama). Distr.- Costa Rica, Panama. todatangae WIRTH & BLANTON, 1973: 447 (male, female; Brazil). Distr.- Trinidad, Brazil (Pará, Santa Catarina). youngi WIRTH & BARRETO, 1978: 562 (male, female; Colombia). Distr.- Colombia. Subgenus HOFFMANIA FOX HOFFMANIA FOX, 1948: 21 (as subgenus of Culicoides). Type species: Culicoides inamollae FOX & HOFFMAN (= Culicoides insignis LUTZ), by original designation. REFERENCES: SPINELLI, GREINER & WIRTH, 1993 (key to species groups; key to males and females of guttatus group); WIRTH & BLANTON, 1968a (key to males and females in hylas group). aitkeni WIRTH & BLANTON, 1968a: 214 (female; Trinidad). Distr.- Trinidad, Brazil (Amazonas). annettae SPINELLI & BORKENT, 2004a: 365 (male, female; Costa Rica). Distr.- Costa Rica. batesi WIRTH & BLANTON, 1973: 426 (male, female; Brazil). Distr.- Guatemala, Colombia, Ecuador, Bolivia, Brazil (Pará). sanmartini WIRTH & BARRETO, 1978: 553. Colombia. biestroi SPINELLI & RONDEROS, 1991: 86 (male, female; Argentina). Distr.- Eastern Argentina, Uruguay.

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bimaculatus FLOCH & ABONNENC, 1942b: 3 (female; French Guiana). Distr.- French Guiana, Brazil (Pará). brasilianum FORATTINI, 1956a: 81 (female; Brazil). Distr.- Northeastern Argentina, southeastern Brazil. brownei SPINELLI, in SPINELLI et al., 1993: 24 (male, female; Colombia). Distr.- Colombia. charruus SPINELLI & Martinez, 1992: 176 (as charrua; pupa, male, female; Uruguay). Distr.Paraguay, northeastern Argentina, Uruguay. coutinhoi BARRETTO, 1944: 96 (male; Brazil). Distr.- French Guiana, Brazil (Amazonas, Pará, São Paulo). davidi SPINELLI, in SPINELLI et al., 1993: 30 (female; Colombia). Distr.- Costa Rica, Colombia, Trinidad. diabolicus HOFFMAN, 1925: 294 (female; Panama). Distr.- Mexico to Venezuela and Ecuador. diffusus SPINELLI, in SPINELLI et al., 1993: 34 (female; Brazil). Distr.- Brazil (Espirito Santo). fernandoi TAVARES & SOUZA, 1979: 611 (male, female; Brazil). Distr.- Colombia, Brazil (Espírito Santo, Rio de Janeiro, Santa Catarina), northeastern Argentina, Uruguay. ferreyrai RONDEROS & SPINELLI, 1995a: 61 (female; Argentina). Distr.- Northeastern Argentina. filarifer HOFFMAN, 1939: 172 (female; Mexico). Distr.- Mexico (Veracruz, Chiapas) to northern Brazil. flavivenulus COSTA LIMA, 1937a: 418 (as flavivenula; female; Brazil). Distr.- Trinidad, French Guiana, Brazil (Pará, Pernambuco, Mato Grosso, Espírito Santo, São Paulo, Rio de Janeiro, Santa Catarina). foxi ORTÍZ, 1950c: 461 (male; Puerto Rico). Distr.- Mexico (Chiapas) to Bolivia, Puerto Rico to northeastern Argentina. franklini SPINELLI, in SPINELLI et al., 1993: 45 (male, female; Panama). Distr.- Mexico (Guerrero) to Bolivia, Brazil (Pará). fusipalpis WIRTH & BLANTON, 1973: 435 (male, female; Brazil). Distr.- El Salvador to Ecuador, Bolivia, French Guiana, Guyana, Brazil (Amazonas, Pará, Bahia, Rio de Janeiro). guttatus (COQUILLETT), 1904b: 35 (Ceratopogon; female; Brazil). Distr.- Paraguay, northeastern Argentina, southeastern Brazil. heliconiae FOX & HOFFMAN, 1944: 108 (male, female; Venezuela). Distr.- Belize to Ecuador, Venezuela, Grenada, Trinidad and Tobago. rozeboomi BARBOSA, 1947: 26. Trinidad. hylas MACFIE, 1940d: 26 (female; Guyana). Distr.- Mexico (Veracruz) to Peru, Brazil (Amazonas). ignacioi FORATTINI, 1957: 215 (male, female; Brazil). Distr.- Brazil (Minas Gerais, São Paulo, Rio de Janeiro), Paraguay. saintjusti TAVARES & RUIZ, 1980: 27. Brazil. insignis LUTZ, 1913: 51 (pupa, male, female; Brazil). Distr.- USA (Alabama, Georgia, Florida), Mexico (Yucatán, Chiapas), Central America and Caribbean to central Argentina. inamollae FOX & HOFFMAN, 1944: 110. Puerto Rico. painteri FOX, 1946: 257. Honduras. lutzi COSTA L IMA, 1937a: 419 (female; Brazil). Distr.- Colombia to French Guiana, northeastern Argentina, Brazil (Roraima, Amazonas, Pará, Mato Grosso, Goiás, São Paulo, Rio de Janeiro, Santa Catarina).

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maruim LUTZ, 1913: 48 (female; Brazil). Distr.- Venezuela to French Guiana, Trinidad, coastal Brazil. recifei BARBOSA, 1947: 25. Brazil. ocumarensis ORTÍZ, 1950b: 455 (male, female; Venezuela). Distr.- Mexico (Oaxaca, Tabasco) to northern Brazil (Pará, Rondônia). palpalis MACFIE, 1948: 78 (female; Mexico). Distr.- Mexico (Chiapas) to Peru, Brazil (Amazonas). paraignacioi SPINELLI, in SPINELLI et al., 1993: 66 (male, female; Colombia). Distr.- Belize to Colombia, French Guiana, Brazil (Amazonas, Pará). paramaruim WIRTH & BLANTON, 1973: 443 (male, female; Brazil). Distr.- Brazil (Pará). plaumanni SPINELLI, in SPINELLI et al., 1993: 69 (female; Argentina). Distr.- Bolivia, Brazil (Amazonas) northeastern Argentina. polypori WIRTH & BLANTON, 1968a: 212 (male, female; Panama). Distr.- Costa Rica to Colombia, Brazil (Amazonas). pseudodiabolicus FOX, 1946: 256 (female; Trinidad). Distr.- Mexico (Veracruz) to Peru and northern Brazil. ruizi FORATTINI, 1954a: 189 (male, female; Brazil). Distr.- Colombia, Brazil (Amazonas, Goiás). tidwelli SPINELLI, in SPINELLI et al., 1993: 74 (male, female; Colombia). Distr.- Honduras to Colombia, Ecuador. travassosi FORATTINI, 1957: 198 (male, female; Brazil). Distr.- Surinam, Brazil (Amazonas, Pará, Mato Grosso). trinidadensis HOFFMAN, 1925: 286 (female; Trinidad and Tobago). Distr.- Coastal; Honduras and El Salvador to Colombia, Ecuador to Surinam, Cuba and Cayman Islands to Trinidad. oliveri FOX & HOFFMAN, 1944: 108. Haiti. wokei BARBOSA, 1947: 28 (preoccupied by Culicoides wokei FOX, 1947). Panama. diminutus BARBOSA, 1951: 163. New name for wokei. verecundus MACFIE, 1948: 76 (male, female; Mexico). Distr.- Mexico (Chiapas) to Ecuador. contubernalis ORTÍZ & LEÓN, 1955: 574 (as variety of rozeboomi). Ecuador. xanifer WIRTH & BLANTON, 1968a: 210 (male, female; Panama). Distr.- Honduras to Panama. Subgenus MACFIELLA FOX MACFIELLA FOX, 1955: 217 (as subgenus of Culicoides). Type species: Ceratopogon phlebotomus WILLISTON, by original designation. phlebotomus (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- Coastal; Mexico (Sinaloa) to Ecuador, Jamaica to Brazil (Maranhão, Ceará, Pernambuco, Goiás). amazonius MACFIE, 1935a: 52. Brazil. willistoni WIRTH & BLANTON, 1953c: 116 (male, female; Panama). Distr.- Mexico (Sonora), Honduras, Panama. Subgenus MATAEMYIA VARGAS MATAEMYIA VARGAS, 1960: 43 (as subgenus of Culicoides). Type species: Culicoides mojingaensis WIRTH & BLANTON, by original designation.

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REFERENCE: WIRTH & SORIA, 1981 (key males and females to some species, as discrepans group). albuquerquei WIRTH & BLANTON, 1973: 424 (male, female; Brazil). Distr.- Brazil (Pará). avilaensis ORTÍZ & MIRSA, 1951: 593 (female; Venezuela). Distr.- Venezuela. azureus WIRTH & BLANTON, 1959: 377 (male, female; Panama). Distr.- Panama. barthi TAVARES & SOUZA, 1978: 619 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro). bricenoi ORTÍZ, 1951a: 445 (male; Venezuela). Distr.- Ecuador, Venezuela, Bolivia, Brazil (Amazonas, Pará). cuiabai WIRTH, 1982b: 250 (female; Brazil). Distr.- Brazil (Mato Grosso), northeastern Argentina. dalessandroi WIRTH & BARRETO, 1978: 556 (male, female; Colombia). Distr.- Costa Rica, Panama, Colombia. daviesi WIRTH & BLANTON, 1968b: 251 (female; Guyana). Distr.- Peru, Guyana. dicrourus WIRTH & BLANTON, 1955b: 123 (male, female; Panama). Distr.- Costa Rica to Ecuador. discrepans ORTÍZ & MIRSA, 1951: 595 (female; Venezuela). Distr.- Venezuela. lenti TAVARES & LUNA DIAS, 1980: 396 (male; Brazil). Distr.- Bolivia, Brazil (Rio de Janeiro). macieli TAVARES & RUIZ, 1980: 29 (male, female; Brazil). Distr.- Brazil (Bahia, Rio de Janeiro). mojingaensis WIRTH & BLANTON, 1953b: 232 (male, female; Panama). Distr.- Panama. volcanensis WIRTH & BLANTON, 1959: 389 (female; Panama). Distr.- Panama, Colombia. wallacei WIRTH & BLANTON, 1973: 449 (male, female; Brazil). Distr.- Brazil (Pará). Subgenus OECACTA POEY OECACTA POEY, 1853: 238. Type species: Oecacta furens POEY, by monotypy. alahialinus BARBOSA, 1952: 11 (1953: 12) (female; Ecuador). Distr.- Costa Rica, Panama, Colombia, Ecuador. barbosai WIRTH & BLANTON, 1956a: 161 (male, female; Panama). Distr.- USA (Florida) to Ecuador. cancer HOGUE & WIRTH, 1968: 2 (larva, pupa, male, female; Costa Rica). Distr.- Mexico (Sinaloa), El Salvador, Costa Rica. furens (POEY), 1853: 238 (Oecacta; female; Cuba). Distr.- USA (Massachusetts to Florida and Texas), Mexico (Campeche, Sinaloa, Yucatán, Veracruz) and Caribbean to Ecuador and coastal Brazil. maculithorax (WILLISTON), 1896: 277 (Ceratopogon). St. Vincent. birabeni CAVALIERI, 1966: 59. Venezuela. gorgasi WIRTH & BLANTON, 1953b: 232 (male, female; Panama). Distr.- Costa Rica to Colombia. Subgenus PSYCHOPHAENA PHILIPPI PSYCHOPHAENA PHILIPPI, 1865: 628. Type species: Psychophaena pictipennis PHILIPPI (= Culicoides venezuelensis ORTÍZ & MIRSA), by monotypy.

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lacustris RONDEROS, 1990a: 116 (male, female; Argentina). Distr.- Southern Argentina. venezuelensis ORTÍZ & MIRSA, 1950: 137 (Venezuela). Distr.- Costa Rica to Chile and Central Argentina. pictipennis (PHILIPPI), 1865: 628 (Psychophaena) (preoccupied by Culicoides pictipennis (STAEGER), 1839). Chile. ortizi FOX, 1952: 366. Venezuela. Subgenus unplaced, acotylus Species Group acotylus LUTZ, 1913: 69 (female; Brazil). Distr.- Mexico (DF), Honduras, Panama, Venezuela, Trinidad, Surinam, Brazil (Mato Grosso, Pará). panamericanus FOX, 1947: 90. Mexico. atripalpis WIRTH & BLANTON, 1973: 425 (female; Brazil). Distr.- Brazil (Pará). carsiomelas WIRTH & BLANTON, 1955a: 100 (male, female; Panama). Distr.- Panama, Colombia, Brazil (Pará). teretipalpis WIRTH & BARRETO, 1978: 557 (male, female; Colombia). Distr.- Colombia. Subgenus unplaced, carpenteri Species Group belemensis WIRTH & BLANTON, 1973: 427 (male, female; Brazil). Distr.- Colombia, Brazil (Amazonas, Pará). camposi ORTÍZ & LEÓN, 1955: 580 (female; Ecuador). Distr.- Costa Rica, Panama, Colombia, Ecuador. fairchildi WIRTH & BLANTON, 1955a: 102. Panama. carpenteri WIRTH & BLANTON, 1953a: 72 (male, female; Panama). Distr.- Costa Rica, Panama, Ecuador, Bolivia, Brazil (Amazonas). Subgenus unplaced, daedalus Species Group antefurcatus WIRTH & BLANTON, 1959: 315 (male, female; Panama). Distr.- Panama. beaveri WIRTH & BARRETO, 1978: 557 (female; Colombia). Distr.- Colombia. commatis WIRTH & BLANTON, 1959: 321 (male, female; Panama). Distr.- Panama. crescentis WIRTH & BLANTON, 1959: 317 (male, female; Panama). Distr.- Mexico (Chiapas) to Colombia, northeastern Argentina. cummingi SPINELLI & BORKENT, 2004a: 379 (female; Costa Rica). Distr.- Costa Rica. daedaloides WIRTH & BLANTON, 1959: 330 (male, female; Panama). Distr.- Panama, Colombia. daedalus MACFIE, 1948: 83 (male; Mexico). Distr.- USA (Arizona, New Mexico), Mexico (Chiapas) to Colombia. dunni WIRTH & BLANTON, 1959: 328 (female; Panama). Distr.- Costa Rica, Panama. pampoikilus MACFIE, 1948: 79 (female; Mexico). Distr.- USA (Arizona, New Mexico), Mexico (Chiapas, Oaxaca) to Venezuela. dominicii ORTÍZ, 1951b: 7. Venezuela. phaeonotus WIRTH & BLANTON, 1959: 326 (male, female; Panama). Distr.- Panama.

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picadoae SPINELLI & BORKENT, 2004a: 381 (male, female; Costa Rica). Distr.- Costa Rica. pseudocrescentis TAVARES & LUNA DIAS, 1980: 397 (male; Brazil). Distr.- Brazil (Rio de Janeiro), Paraguay, northeastern Argentina. Subgenus unplaced, dasyophrus Species Group dasyophrus MACFIE, 1940d: 27 (male, female; Guyana). Distr.- Colombia, Ecuador, Venezuela, Guyana, Brazil (Amazonas, Mato Grosso, Pará). estevezae RONDEROS & SPINELLI, 1994: 47 (female; Argentina). Distr.- Northern Argentina. guerrai WIRTH & BLANTON, 1971: 41 (male, female; Trinidad. Distr.- Trinidad, Brazil (Pará, Bahia). rodriguezi ORTÍZ, 1968: 67 (female; Venezuela). Distr.- Panama, Venezuela. Subgenus unplaced, eublepharus Species Group archboldi WIRTH & BLANTON, 1970b: 39 (male, female; Dominica). Distr.- Dominica, Trinidad. caldasi BROWNE, 1980: 535 (female; Colombia). Distr.- Colombia. caucaensis WIRTH & LEE, 1967: 19 (female; Colombia). Distr.- Colombia. eublepharus MACFIE, 1948: 86 (female; Guyana). Distr.- Mexico (Chiapas), Costa Rica to Ecuador, Venezuela, northern Brazil. transferrans ORTÍZ, 1953a: 801. Venezuela. florenciae MESSERSMITH, 1972: 167 (female; Colombia). Distr.- Colombia. guadeloupensis FLOCH & ABONNENC, 1950b: 2 (female; Guadeloupe). Distr.- Guadeloupe. guarani RONDEROS & SPINELLI, 1994: 48 (female; Argentina). Distr.- Northeastern Argentina. pabloi BROWNE, 1980: 541 (male, female; Colombia). Distr.- Colombia. propriipennis MACFIE, 1948: 84 (female; Mexico). Distr.- Mexico (Chiapas) to Panama, Ecuador, Venezuela and northern Brazil. rangeli ORTÍZ & MIRSA, 1952b: 126 (female; Venezuela). Distr.- Mexico (Oaxaca) to Ecuador, Bolivia, Venezuela, Trinidad, Brazil (Amazonas). donajii VARGAS, 1954: 28. Mexico. patulipalpis WIRTH & BLANTON, 1959: 421. Panama. tamboensis WIRTH & LEE, 1967: 20 (male, female; Colombia). Distr.- Colombia. zumbadoi SPINELLI & BORKENT, 2004a: 383 (male, female; Costa Rica). Distr.- Costa Rica. Subgenus unplaced, fluvialis Species Group balsapambensis ORTÍZ & LEÓN, 1955: 569 (as variety of pifanoi; female; Ecuador). Distr.Costa Rica to Ecuador, Brazil. castillae FOX, 1946: 251 (female; Honduras). Distr.- Guatemala to Ecuador, Venezuela, Trinidad. gibsoni WIRTH, 1952c: 246. Guatemala. flochabonnenci ORTÍZ & MIRSA, 1952a: 267. Venezuela. fernandezi ORTÍZ, 1954: 223 (female; Venezuela). Distr.- Venezuela. fluvialis MACFIE, 1940d: 25 (female; Guyana). Distr.- Honduras to Colombia, Venezuela, Trinidad, Guyana, Brazil (Amazonas, Pará). leopoldoi ORTÍZ, 1951c: 579 (female; Venezuela). Distr.- Guatemala and Belize to Bolivia and northeastern Argentina, Trinidad.

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lichyi FLOCH & ABONNENC, 1949: 1 (1950a: 69) (female; Venezuela). Distr.- Venezuela. pulchripennis MACFIE, 1939b: 200 (female; Brazil). Distr.- Brazil (Santa Catarina). tetrathyris WIRTH & BLANTON, 1959: 409 (male, female; Panama). Distr.- Honduras, Costa Rica, Panama, Ecuador, Trinidad, Surinam, northern Brazil. williamsi SPINELLI, in SPINELLI et al., 2005b: 147 (male, female; Paraguay). Distr.- Paraguay, northeastern Argentina. yaracuyensis ORTÍZ, 1959: 364 (as yaracuyanus; female; Venezuela). Distr.- Venezuela. Subgenus unplaced, leoni Species Group benarrochi ORTÍZ & MIRSA, 1952b: 126 (as benarrochei; female; Venezuela). Distr.- Brazil (Rio de Janeiro), Venezuela, Trinidad. fieldi WIRTH & BLANTON, 1956c: 50 (male, female; Honduras). Distr.- Honduras, Costa Rica, Panama. gabaldoni ORTÍZ, 1954: 221 (female; Venezuela). Distr.- Mexico (Tabasco) to Ecuador, Venezuela, Trinidad, Brazil (Bahia), Paraguay, northeastern Argentina. glabellus WIRTH & BLANTON, 1956c: 47 (male, female; Panama). Distr.- Honduras to Ecuador, Trinidad, Brazil (Bahia, Pará). leoni BARBOSA, 1952: 17 (1953: 19) (female; Ecuador). Distr.- Ecuador. trifidus SPINELLI & BORKENT, 2004a: 385 (male, female; Costa Rica). Distr.- Costa Rica. Subgenus unplaced, limai Species Group antunesi FORATTINI, 1954b: 315 (male, female; Brazil). Distr.- Brazil (Goiás, São Paulo). boliviensis SPINELLI & WIRTH, 1984a: 172 (female; Bolivia). Distr.- Bolivia, Brazil (Minas Gerais), northeastern Argentina. carvalhoi WIRTH & BLANTON, 1973: 429 (male, female; Brazil). Distr.- Brazil (Pará). duartei TAVARES & LUNA DIAS, 1980: 395 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro). galindoi WIRTH & BLANTON, 1953a: 73 (male, female; Panama). Distr.- Costa Rica, Panama. limai BARRETTO, 1944: 99 (male; Brazil). Distr.- El Salvador to Ecuador, Brazil (Pará, Mato Grosso, São Paulo, Rio de Janeiro, Santa Catarina) to northeastern Argentina. lobatoi FELIPPE-BAUER, in FELIPPE-BAUER & QUINTELAS, 1994: 25 (female; Brazil). Distr.Brazil (Rio de Janeiro). lopesi BARRETTO, 1944: 102 (male; Brazil). Distr.- Panama, Surinam, Brazil (São Paulo). santanderi BROWNE, 1980: 536 (female; Colombia). Distr.- Colombia. tenuilobus WIRTH & BLANTON, 1959: 354 (male, female; Panama). Distr.- Guatemala to Panama. vernoni WIRTH & BLANTON, 1973: 448 (male, female; Brazil). Distr.- Costa Rica, Colombia, Bolivia, Brazil (Pará). Subgenus unplaced, monticola Species Group andinus WIRTH & LEE, 1967: 17 (female; Colombia). Distr.- Colombia. magnipalpis WIRTH & BLANTON, 1953a: 76 (male, female; Panama). Distr.- Panama. monticola WIRTH & LEE, 1967: 15 (female; Colombia). Distr.- Costa Rica to Ecuador. pichindensis BROWNE, 1980: 538. Colombia.

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Subgenus unplaced, pachymerus Species Group almirantei WIRTH & BLANTON, 1959: 454 (male, female; Panama). Distr.- Costa Rica, Panama. atelis WIRTH, 1982b: 249 (female; Panama). Distr.- Panama. caprilesi FOX, 1952: 364 (female; Venezuela). Distr.- Panama, Colombia, Venezuela, Brazil (Pará, Mato Grosso). kintzi WIRTH & BLANTON, 1953a: 72. Panama. cylindricornis WIRTH & BLANTON, 1973: 430 (female; Brazil). Distr.- Brazil (Pará). obnoxius FOX, 1952: 365 (female; Venezuela). Distr.- Colombia, Venezuela. pachymerus LUTZ, 1914: 83 (female; Brazil). Distr.- Guatemala to Colombia, Brazil (Amazonas). uniradialis WIRTH & BLANTON, 1953a: 70 (male, female; Panama). Distr.- Panama, Colombia. Subgenus unplaced, reticulatus Species Group aureus ORTÍZ, 1951c: 585 (male; Venezuela). Distr.- Panama, Bolivia, Venezuela, Paraguay, Brazil (Amazonas), northeastern Argentina. miyamotoi WIRTH & BLANTON, 1953b: 231. Panama. fittkaui WIRTH & BLANTON, 1973: 432 (male, female; Brazil). Distr.- Brazil (Pará). forattinii ORTÍZ, 1961: 211 (female; Venezuela). Distr.- Venezuela. goeldii WIRTH & BLANTON, 1973: 437 (male, female; Brazil). Distr.- Brazil (Pará). guamai WIRTH & BLANTON, 1973: 438 (male, female; Brazil). Distr.- Brazil (Pará). guyanensis FLOCH & ABONNENC, 1942a: 4 (male, female; French Guiana). Distr.- Panama, Venezuela to French Guiana, Trinidad and Tobago, Brazil (Pará, Pernambuco, São Paulo). recifensis BARBOSA, 1943: 263. Brazil. stubalensis FOX, 1946: 254. Trinidad and Tobago. kuscheli WIRTH & BLANTON, 1978: 236 (female; Chile). Distr.- Northern Chile. lanei ORTÍZ, 1950a: 431 (male; Panama). Distr.- Mexico (Veracruz), Honduras, Costa Rica, Panama, Venezuela, Trinidad, Brazil (Pará). lyrinotatus WIRTH & BLANTON, 1955b: 126 (male, female; Panama). Distr.- Nicaragua, Panama, Brazil. macrostigma WIRTH & BLANTON, 1953b: 230 (male, female; Panama). Distr.- Costa Rica to Colombia. martinezi WIRTH & BLANTON, 1970b: 43 (male, female; Trinidad). Distr.- Trinidad. paucienfuscatus BARBOSA , 1947: 23 (male; Brazil). Distr.- Costa Rica to Peru and Bolivia,Venezuela, Trinidad, Brazil (Amazonas, Pará). pifanoi ORTÍZ, 1951c: 588 (male, female; Venezuela). Distr.- Belize to Colombia, Venezuela, Trinidad, Brazil (Pará, Bahia), Paraguay. tricoloratus WIRTH & BLANTON, 1953b: 233. Panama. reticulatus LUTZ, 1913: 49 (male, female; Brazil). Distr.- Honduras to Colombia, Brazil (Pernambuco, Bahia, São Paulo, Rio de Janeiro). tavaresi FELIPPE-BAUER & WIRTH, 1988: 261 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).

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Subgenus unplaced, stigmalis Species Group REFERENCE: FELIPPE-BAUER & WIRTH, 1987 (key to males and females). alvarezi ORTÍZ, 1957: 161 (female; Venezuela). Distr.- Ecuador, Venezuela. deanei FELIPPE-BAUER & WIRTH, 1987: 416 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro) fluviatilis (LUTZ), 1914: 82 (Johannseniella; female; Brazil). Distr.- Colombia, Ecuador, Bolivia, Venezuela, Trinidad, Brazil (Amazonas). scorzai ORTÍZ, 1956: 93. Venezuela. stigmalis WIRTH, 1952c: 245 (female; Guatemala). Distr.- Mexico (Oaxaca), Guatemala, Costa Rica, Panama. Subgenus unplaced, stonei Species Group melleus (COQUILLETT), 1901a: 604 (Ceratopogon; female; Florida). Distr.- Eastern Canada (New Brunswick) and USA (to Louisiana and Florida), Bahamas, Mexico (Baja California). Miscellaneous Unplaced Species arubae FOX & HOFFMAN, 1944: 109 (male, female; Aruba). Distr.- USA (Texas), Mexico (Yucatán), Aruba and Grenada, to Colombia and Venezuela. floridensis BECK, 1951: 135 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas, Bermuda. malariologiensis PERRUOLO, 1990: 28 (female; Venezuela). Distr.- Venezuela. pancensis BROWNE, 1980: 537 (male, female; Colombia). Distr.- Colombia. polystictus KIEFFER, 1921b: 181 (female; Paraguay). Distr.- Paraguay. propinquus MACFIE, 1948: 81 (male; Mexico). Distr.- Mexico (Chiapas). trilineatus FOX, 1946: 250 (female; Virgin Islands). Distr.- Guatemala to Panama, Puerto Rico, Virgin Islands, Dominica, Barbados, Paraguay. unetensis PERRUOLO, 2001: 35 (female; Venezuela). Distr.- Venezuela. wokei FOX, 1947: 90 (female; Panama). Distr.- Costa Rica, Panama. Genus PARADASYHELEA MACFIE PARADASYHELEA MACFIE, 1940f: 17. Type species: Dasyhelea brevipalpis INGRAM & MACFIE, by original designation. REFERENCES: SPINELLI, 1987 (description of female); SPINELLI & GROGAN, 2003 (key to males and females). brevipalpis (INGRAM & MACFIE), 1931: 178 (Dasyhelea; male; Argentina). Distr.- Southwestern Argentina south to Tierra del Fuego. ingrami SPINELLI & GROGAN, 2003: 572 (male, female; Argentina). Distr.- Southtern Argentina. macfiei SPINELLI & GROGAN, 2003: 574 (male, female; Argentina). Distr.- Southwestern Argentina, southern Chile.

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TRIBE CERATOPOGONINI Genus ALLOHELEA KIEFFER ALLOHELEA KIEFFER, 1917b: 364. Type species: Sphaeromias pulchripennis KIEFFER, by original designation. neotropica WIRTH, 1991a: 503 (male, female; Jamaica). Distr.- Belize, Panama, Colombia, Jamaica. Genus ALLUAUDOMYIA KIEFFER ALLUAUDOMYIA KIEFFER, 1913: 12. Type species: Alluaudomyia imparunguis KIEFFER, by monotypy. NEOCERATOPOGON MALLOCH, 1915b: 310. Type species: Ceratopogon bellus COQUILLETT, by original designation. ISOECACTA GARRETT, 1925: 9. Type species: Isoecacta poeyi GARRETT (= Ceratopogon bellus COQUILLETT), by original designation. REFERENCES: HUERTA, 1996 (key to males and females from Mexico); SPINELLI & WIRTH, 1984c (key to males and females). amazonica SPINELLI & WIRTH, 1984c: 676 (female, male; Brazil). Distr.- Brazil (Amazonas), northeastern Argentina. bella (COQUILLETT), 1902: 87 (Ceratopogon; USA, District of Columbia). Distr.- North America (Alaska to California and Nova Scotia and Florida), Mexico (Veracruz), Bahamas, Cayman Islands. biestroi SPINELLI, 1988: 130 (pupa, male; Argentina). Distr.- Northeastern Argentina, Uruguay. caribbeana SPINELLI & WIRTH, 1984c: 678 (pupa, male, female; Belize). Distr.- USA (California to Texas, Florida), Central America, circum-Caribbean from Mexico (Yucatán, Morelos, Sinaloa) to Colombia and Venezuela. catarinensis SPINELLI & WIRTH, 1984c: 681 (female; Brazil). Distr.- Brazil (Santa Catarina). distispinulosa SPINELLI & WIRTH, 1984c: 682 (male, female; Brazil). Distr.- Colombia, Bolivia, Venezuela, Brazil (Amazonas). estevezae SPINELLI & WIRTH, 1984c: 684 (male, female; Mexico). Distr.- Mexico (Morelos), Honduras, El Salvador. fittkaui SPINELLI & WIRTH, 1984c: 686 (male, female; Brazil). Distr.- Ecuador, Brazil (Amazonas, Pará). guarani SPINELLI, 1988: 131 (male, female; Uruguay). Distr.- Paraguay, northeastern Argentina, Uruguay. leei SPINELLI & WIRTH, 1984c: 687 (male, female; Colombia). Distr.- Colombia, Bolivia. nubeculosa SPINELLI & WIRTH, 1984c: 689 (male, female; Brazil). Distr.- Colombia, Brazil (Amazonas, Pará). plaumanni SPINELLI & WIRTH, 1984c: 691 (female; Brazil). Distr.- Brazil (Santa Catarina). prima CLASTRIER, 1976: 205 (female; French Guiana). Distr.- Panama, French Guiana.

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punctivenosa WIRTH & GROGAN, 1988: 20. New name for punctiradialis. Distr.- Brazil (Pará), Paraguay, northeastern Argentina. punctiradialis SPINELLI & WIRTH, 1984c: 692 (preoccupied by Alluaudomyia punctiradialis CHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Brazil. schnacki SPINELLI, 1983b: 403 (larva, pupa, male, female; Argentina). Distr.- Eastern Argentina, Uruguay. sexpunctata SPINELLI & WIRTH, 1984c: 695 (female; Colombia). Distr.- Colombia. spinellii WIRTH & GROGAN, 1988: 21. New name for tripunctata. Distr.- Colombia. tripunctata SPINELLI & WIRTH , 1984c: 697 (preoccupied by Alluaudomyia tripunctata CHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Colombia. tenuiannulata SPINELLI & WIRTH, 1984c: 696 (male, female; Guatemala). Distr.- Guatemala, Costa Rica. youngi SPINELLI & WIRTH, 1984c: 699 (male, female; Colombia). Distr.- Panama, Colombia. Genus AUSTROHELEA WIRTH & GROGAN AUSTROHELEA WIRTH & GROGAN, 1988: 22. Type species: Monohelea shannoni WIRTH & BLANTON, by original designation. REFERENCE: WIRTH & GROGAN, 1988 (description of male). shannoni (WIRTH & BLANTON), 1972b: 175 (Monohelea; female; Argentina). Distr.- Southern Chile, southwestern Argentina south to Tierra del Fuego. Genus BAEODASYMYIA CLASTRIER & RACCURT BAEODASYMYIA CLASTRIER & RACCURT, 1979b: 100. Type species: Baeodasymyia modesta CLASTRIER & RACCURT, by monotypy. REFERENCE: BORKENT & CRAIG, 1999 (keys to pupae, males, females). christopheri BORKENT, in BORKENT & CRAIG, 1999: 15 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica. gustavoi BORKENT, in BORKENT & CRAIG, 1999: 20 (female; Paraguay). Distr.- Paraguay, Argentina (Misiones). lydiae BORKENT, in BORKENT & CRAIG, 1999: 19 (male, female; Costa Rica). Distr.- Costa Rica. michaeli BORKENT, in BORKENT & CRAIG, 1999: 13 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica, Panama. modesta CLASTRIER & RACCURT, 1979b: 100 (male, female; Haiti). Distr.- Haiti. Genus BAEOHELEA WIRTH & BLANTON BAEOHELEA WIRTH & BLANTON, 1970c: 95. Type species: Baeohelea nana WIRTH & BLANTON, by original designation.

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nana WIRTH & BLANTON, 1970c: 95 (male, female; Dominica). Distr.- Costa Rica to Ecuador, St. Kitts, Dominica. Genus BAHIAHELEA WIRTH BAHIAHELEA WIRTH, 1992: 276. Type species: Bahiahelea brasiliensis WIRTH, by original designation. brasiliensis WIRTH, 1992: 280 (male, female; Brazil). Distr.- Brazil (Bahia). Genus BORKENTHELEA SPINELLI & GROGAN BORKENTHELEA SPINELLI & GROGAN, 1993: 321. Type species: Borkenthelea nothofagus SPINELLI & GROGAN, by original designation. REFERENCE: SPINELLI & GROGAN, 2001 (key to males and females species). harii SPINELLI & GROGAN, 2001: 148 (male, female; Argentina). Distr. - Southwestern Argentina, southern Chile. nerudai SPINELLI & GROGAN, 2001: 151 (male, female; Chile). Distr. - Southern Chile. nothofagus SPINELLI & GROGAN, 1993: 323 (male, female; Argentina). Distr.- Southwestern Argentina, southern Chile. quatei SPINELLI & GROGAN, 2001: 153 (male, female; Argentina). Distr.- Argentina (Chubut). Genus BRACHYPOGON KIEFFER BRACHYPOGON KIEFFER, 1899: 69. Type species: Ceratopogon vitiosus WINNERTZ, by original designation. REFERENCE: SPINELLI & WIRTH, 1993 (key to subgenera as part of larger key to genera). Subgenus BRACHYPOGON KIEFFER REFERENCE: SPINELLI & GROGAN, 1998 (key to males and females). apunctipennis SPINELLI & GROGAN, 1998: 64 (male; Brazil). Distr.- Brazil (Rondônia). bifidus SPINELLI & GROGAN, 1998: 66 (male, female; Dominica). Distr.- El Salvador, Dominica. bimaculatus SPINELLI & GROGAN, 1998: 67 (male, female; Colombia). Distr.- Mexico (Yucatán), Colombia, Brazil (Mato Grosso). bonaerensis SPINELLI, 1990: 744 (female; Argentina). Distr.- Argentina (Buenos Aires). calchaqui SPINELLI, 1990: 746 (male, female; Argentina). Distr.- Western Argentina. ecuadorensis SPINELLI & GROGAN, 1998: 68 (male, female; Ecuador). Distr.- Belize to Ecuador, Jamaica. ethelae SPINELLI & GROGAN, 1998: 62 (male, female; Panama). Distr.- Panama.

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fuscivenosus (LUTZ), 1914: 94 (Palpomyia; male, female; Brazil). Distr.- USA (Florida), Mexico (Yucatán, Quintana Roo), Panama, Jamaica, Virgin Islands, Guyana, Brazil (Rio de Janeiro). impar (JOHANNSEN), 1938: 223 (Ceratopogon; female; Puerto Rico). Distr.- Colombia, Venezuela, Puerto Rico, Brazil (Santa Catarina). insularis SPINELLI & GROGAN, 1998: 70 (male; Cuba). Distr.- Cuba. monicae SPINELLI & GROGAN, 1998: 70 (male, female; Colombia). Distr.- Colombia. paraensis WIRTH & BLANTON, 1970c: 99 (as parasensis; male, female; Brazil). Distr.- Brazil (Pará). pseudoparaensis SPINELLI & GROGAN, 1998: 64 (male; Brazil). Distr.- Brazil (Pará). ringueleti SPINELLI, 1990: 748 (male, female; Argentina). Distr.- Southern Argentina. schmitzi SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondônia). spatuliformis SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondônia). telesfordi SPINELLI & GROGAN, 1998: 71 (male, female; St. Vincent). Distr.- Costa Rica, St. Vincent, Grenada. woodruffi SPINELLI & GROGAN, 1998: 72 (male, female; Dominica Republic). Distr.- Mexico (Yucatán), Dominica Republic. Subgenus ISOHELEA KIEFFER ISOHELEA KIEFFER, 1917b: 295. Type species: Ceratopogon lacteipennis ZETTERSTEDT (misidentified, = Psilohelea sociabilis GOETGHEBUER), by original designation. REFERENCE: SPINELLI & GROGAN, 1994 (key to males and females). borkenti SPINELLI & CAZORLA, 2004: 3 (male, female; Argentina). Distr.- Western, southern Argentina. cuacuahuitlus HUERTA & BORKENT, 2005: 115 (male; Mexico). Distr.- Mexico (Jalisco). hugoi SPINELLI & GROGAN, 1994: 2 (male, female; Colombia). Distr.- Colombia. mapuche SPINELLI, 1990: 750 (male, female; Argentina). Distr.- Southwestern Argentina. misionensis SPINELLI, 1990: 752 (male; Argentina). Distr.- Northeastern Argentina, southeastern Brazil. pallidipennis SPINELLI & GROGAN, 1994: 6 (male, female; Honduras). Distr.- Honduras. wirthi SPINELLI, 1990: 753 (female; Argentina). Distr.- Northwestern Argentina. Genus CACAOHELEA WIRTH & GROGAN CACAOHELEA WIRTH & GROGAN, 1988: 33. Type species: Cacaohelea youngi WIRTH & GROGAN, by original designation. youngi WIRTH & GROGAN, 1988: 34 (female; Costa Rica). Distr.- Costa Rica. Genus CERATOCULICOIDES WIRTH & RATANAWORABHAN CERATOCULICOIDES WIRTH & RATANAWORABHAN, 1971a: 170. Type species: Helea longipennis WIRTH, by original designation.

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aliciae HUERTA & BORKENT, 2005: 112 (male, female; Mexico). Distr.- Mexico (Jalisco). Genus DIAPHANOBEZZIA INGRAM & MACFIE DIAPHANOBEZZIA INGRAM & MACFIE, 1931: 223. Type species: Diaphanobezzia pellucida INGRAM & MACFIE, by original designation. araucaria SPINELLI, 1996: 77 (female; Argentina). Distr.- Southwestern Argentina. patagonica SPINELLI & GROGAN, 1990: 127 (female; Argentina). Distr.- Southwestern Argentina. pellucida INGRAM & MACFIE, 1931: 223 (male; Argentina). Distr.- Southwestern Argentina. spinellii WIRTH & GROGAN, 1988: 48 (male, female; Argentina). Distr.- Southwestern Argentina. Genus DOWNESHELEA WIRTH & GROGAN DOWNESHELEA WIRTH & GROGAN, 1988: 50. Type species: Monohelea stonei WIRTH, by original designation. REFERENCE: LANE & WIRTH, 1964 (key to males and females). balboa (LANE & WIRTH), 1964: 225 (Monohelea; male, female; Panama). Distr.- Panama. bicornis FELIPPE-BAUER & QUINTELAS, 1993b: 185 (male; Brazil). Distr.- Brazil (Rio de Janeiro). blantoni (LANE & WIRTH), 1964: 217 (Monohelea; male, female; Panama). Distr.- Panama. carioca (TAVARES & SILVA PEREIRA), 1978: 157 (Monohelea; male, female; Brazil). Distr.- Brazil (Rio de Janeiro). castroi (TAVARES & SILVA PEREIRA), 1978: 159 (Monohelea; male, female; Brazil). Distr.- Brazil (Rio de Janeiro). cebacoi (LANE & WIRTH), 1964: 218 (Monohelea; male, female; Panama). Distr.- Panama, Brazil (Pará). charrua FELIPPE-BAUER & SPINELLI, 1994: 161 (male, female; Uruguay). Distr.- Uruguay. chiapasi (LANE & WIRTH), 1964: 219 (Monohelea; male, female; Nicaragua). Distr.- Nicaragua to Colombia. chirusi (LANE & WIRTH), 1964: 218 (Monohelea; male, female; Panama). Distr.- Nicaragua, Panama. colombiae (LANE & WIRTH), 1964: 220 (Monohelea; male; Colombia). Distr.- Colombia. deanei FELIPPE-BAUER & QUINTELAS, in FELIPPE-BAUER et al., 1995: 395 (male; Trinidad). Distr.- Trinidad. fluminensis FELIPPE-BAUER & QUINTELAS, 1993a: 33 (male; Brazil). Distr.- Brazil (Rio de Janeiro). fuscipennis (LANE & WIRTH), 1964: 221 (Monohelea). Colombia. guianae (WIRTH), 1953b: 150 (Monohelea; male; Guyana). Distr.- Brazil (Pará), Guyana. multilineata (LUTZ), 1914: 93 (Palpomyia; male, female; Brazil). Distr.- USA (Florida, Puerto Rico), Mexico (Yucatán), Panama, Guyana, Brazil (Mato Grosso, Rio de Janeiro). panamensis (LANE & WIRTH), 1964: 221 (Monohelea; male, female; Panama). Distr.- Mexico (Guerrero), Panama, Virgin Islands.

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quasidentica FELIPPE-BAUER & QUINTELAS, 1993a: 37 (male; Brazil). Distr.- Brazil (Rio de Janeiro). stonei (WIRTH), 1953b: 148 (Monohelea; male, female; USA, Louisiana). Distr.- Eastern North America (Ontario, Massachusetts to Florida), Caribbean to Colombia and Trinidad. Genus ECHINOHELEA MACFIE ECHINOHELEA MACFIE, 1940c: 187. Type species: Echinohelea ornatipennis MACFIE, by original designation. REFERENCE: WIRTH, 1994b (key to males and females). Subgenus ECHINOHELEA MACFIE blantoni WIRTH, 1994b: 232 (male, female; Panama). Distr.- Mexico (Yucatán, Oaxaca), Panama to Ecuador. jamaicensis WIRTH, 1994b: 234 (male, female; Jamaica). Distr.- Jamaica. lanei WIRTH, 1951b: 319 (male, female; USA, Virgina). Distr.- USA (Virginia, Michigan and Massachetts to Florida), Panama, Colombia, Trinidad, Brazil (Pará, Rondônia). leei WIRTH, 1994b: 237 (male; Colombia). Distr.- Colombia. macfiei LANE, 1948: 228 (female; Brazil). Distr.- Brazil (São Paulo). neotropica WIRTH, 1994b: 237 (male, female; Colombia). Distr.- Panama to Ecuador, Brazil (Amazonas, Pará, Rio de Janeiro). ornatipennis MACFIE, 1940c: 188 (male, female; Guyana). Distr.- Panama, Trinidad, Guyana, Brazil (Pará). panamensis WIRTH, 1994b: 239 (male, female; Panama). Distr.- Panama to Ecuador. richardsi MACFIE, 1940c: 189 (female; Guyana). Distr.- Guyana, Brazil (Pará, Rondônia, São Paulo). smarti MACFIE, 1940c: 190 (female; Guyana). Distr.- Colombia, Guyana. Subgenus ECHINOIDESHELEA WIRTH ECHINOIDESHELEA WIRTH, 1994b: 231 (as subgenus of Echinohelea). Type species: Echinohelea aitkeni WIRTH, by original designation. aitkeni WIRTH, 1994b: 231 (male, female; Brazil). Distr.- Brazil (Pará). Genus FITTKAUHELEA WIRTH & BLANTON FITTKAUHELEA WIRTH & BLANTON, 1970a: 7. Type species: Fittkauhelea amazonica WIRTH & BLANTON, by original designation. amazonica WIRTH & BLANTON, 1970a: 9 (male, female; Brazil). Distr.- Brazil (Amazonas).

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Genus ISTHMOHELEA INGRAM & MACFIE ISTHMOHELEA INGRAM & MACFIE, 1931: 208. Type species: Isthmohelea disjuncta INGRAM & MACFIE, by original designation. disjuncta INGRAM & MACFIE, 1931: 209 (male, female; Chile). Distr.- Southern Chile. Genus LEPTOHELEA WIRTH & BLANTON LEPTOHELEA WIRTH & BLANTON, 1970a: 12. Type species: Leptohelea micronyx WIRTH & BLANTON, by original designation. micronyx WIRTH & BLANTON, 1970a: 12 (female; Colombia). Distr.- Costa Rica, Colombia. Genus MACRUROHELEA INGRAM & MACFIE MACRUROHELEA INGRAM & MACFIE, 1931: 203. Type species: Macrurohelea caudata INGRAM & MACFIE, by original designation. REFERENCE: SPINELLI & GROGAN, 1990 (key to males and females). caudata INGRAM & MACFIE, 1931: 205 (male, female; Chile). Distr.- Southern Chile. fuscipennis SPINELLI & GROGAN, 1990: 128 (female; Argentina). Distr.- Southwestern Argentina. gentilii SPINELLI & GROGAN, 1984: 963 (male, female; Argentina). Distr.- Southwestern Argentina. irwini GROGAN & WIRTH, 1980b: 140 (male, female; Chile). Distr.- Central and southern Chile. kuscheli WIRTH, 1965c: 49 (female; Chile). Distr.- Central Chile. monotheca SPINELLI & GROGAN, 1984: 965 (female; Argentina). Distr.- Southwestern Argentina. paracaudata GROGAN & WIRTH, 1980b: 141 (male; Chile). Distr.- Central and southern Chile, southwestern Argentina. setosa WIRTH, 1965c: 49 (male, female; Chile). Distr.- Northern Chile. similis SPINELLI & GROGAN, 1990: 131 (male; Argentina). Distr.- Southwestern Argentina. thoracica INGRAM & MACFIE, 1931: 206 (male, female; Argentina). Distr.- Southwestern Argentina, southern Chile. wirthi SPINELLI & GROGAN, 1984: 965 (female; Argentina). Distr.- Southwestern Argentina. yamana SPINELLI & GROGAN, 1999: 710 (male, female; Argentina). Distr.- Argentina (Tierra del Fuego). Genus MONOHELEA KIEFFER MONOHELEA KIEFFER, 1917b: 294. Type species: Monohelea hieroglyphica KIEFFER, by original designation. REFERENCE: LANE & WIRTH, 1964 (key to males and females).

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affinis FELIPPE-BAUER & SPINELLI, 1991: 201 (male, female; Brazil). Distr.- Brazil (Amazonas). aguirrei TAVARES & SOUZA, 1980: 97 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro, Santa Catarina), Argentina (Corrientes, Buenos Aires province). archibaldoi TAVARES & SOUZA, 1980: 98 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro). bidentata FELIPPE-BAUER & SPINELLI, 1994: 163 (male; Argentina). Distr.- Argentina (Buenos Aires). brasiliensis LANE, 1948: 226 (female; Brazil). Distr.- Brazil (Rio de Janeiro). cunasi LANE & WIRTH, 1964: 232 (male, female; Panama). Distr.- Panama. fairchildi LANE & WIRTH, 1964: 214 (male, female; Panama). Distr.- Panama. guaimiesi LANE & WIRTH, 1964: 227 (male, female; Panama). Distr.- Panama. hieroglyphica KIEFFER, 1917b: 312 (male, female; Paraguay). Distr.- Dominica, Brazil (Goiás, Rio de Janeiro), Paraguay. lanei WIRTH, 1953b: 142 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas, Costa Rica. maculipennis (COQUILLETT), 1905: 64 (Ceratopogon; female; USA, Florida). Distr.- USA (Florida, Louisiana), Bahamas, Mexico (Yucatán, Tamaulipas), Guatemala, Panama. maya FELIPPE-BAUER, HUERTA & IBAÑEZ-BERNAL, 2000a: 815 (male, female; Mexico). Distr.Mexico (Yucatán). mayeri ORTÍZ, 1950d: 202 (female; Venezuela). Distr.- Costa Rica to Colombia, Venezuela, Trinidad. ornata WIRTH, 1953b: 144 (female; USA, Florida). Distr.- USA (Florida), Panama. poncai LANE & WIRTH, 1964: 228 (male, female; Panama). Distr.- Costa Rica, Panama, Brazil (Rio de Janeiro). roraimensis FELIPPE-BAUER & SPINELLI, 1991: 202 (male; Brazil). Distr.- Brazil (Roraima). urracaisi LANE & WIRTH, 1964: 231 (male, female; Panama). Distr.- Panama, Guyana. uruguayensis FELIPPE-BAUER & SPINELLI, 1998: 63 (male; Uruguay). Distr.- Uruguay. Genus NANNOHELEA GROGAN & WIRTH NANNOHELEA GROGAN & WIRTH, 1980a: 374. Type species: Ceratopogon bourioni CLASTRIER, by original designation. clastrieri GROGAN & WIRTH, 1980a: 381 (male; Colombia). Distr.- Colombia. Genus NOTIOHELEA GROGAN & WIRTH NOTIOHELEA GROGAN & WIRTH, 1979b: 283. Type species: Notiohelea chilensis GROGAN & WIRTH, by original designation. chilensis GROGAN & WIRTH, 1979b: 284 (female; Chile). Distr.- Southern Chile. pilosa SPINELLI & GROGAN, 1990: 133 (female; Argentina). Distr.- Southwestern Argentina. Genus PARABEZZIA MALLOCH PARABEZZIA MALLOCH, 1915a: 358. Type species: Parabezzia petiolata MALLOCH, by original designation.

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REFERENCE: SPINELLI & GROGAN, 1987 (key to females and some males). alexanderi WIRTH, 1965b: 219 (female; USA, Massachesetts). Distr.- Eastern North America (Ontario and Massachesetts to Florida), Mexico (Veracruz), Belize, El Salvador, Argentina (Misiones). arenosa CLASTRIER & RACCURT, 1979a: 172 (female; Haiti). Distr.- Haiti. balseiroi SPINELLI & GROGAN, 1987: 13 (pupa, male, female; Argentina). Distr.- Argentina (Mendoza, Corrientes, Entre Ríos). blantoni WIRTH, 1965b: 220 (female; Panama). Distr.- Panama. brasiliensis SPINELLI & GROGAN, 1987: 34 (female; Brazil). Distr.- Brazil (Mato Grosso). brunnea WIRTH, 1965b: 220 (female; Panama). Distr.- Belize, Panama. caribbeana CLASTRIER & RACCURT, 1979a: 173 (female; Haiti). Distr.- Haiti. cayoensis SPINELLI & GROGAN, 1987: 36 (female; Belize). Distr.- Belize. clastrieri SPINELLI & GROGAN, 1987: 23 (female; El Salvador). Distr.- Belize, El Salvador. costalis WIRTH, 1965b: 221 (male, female; Panama). Distr.- Panama. fuscipennis WIRTH, 1965b: 223 (female; Panama). Distr.- Panama. haitiensis CLASTRIER & RACCURT, 1979a: 170 (female; Haiti). Distr.- Haiti. hondurensis SPINELLI & GROGAN, 1987: 25 (male, female; Honduras). Distr.- Honduras. inaequalis SPINELLI & GROGAN, 1987: 38 (male, female; Brazil). Distr.- Brazil (Amazonas). jamaicensis WIRTH, 1965b: 224 (female; Jamaica). Distr.- Jamaica. neunguis GROGAN & WIRTH, 1977: 65 (female; USA, Virgina). Distr.- USA (Virginia, Maryland), El Salvador. pallida SPINELLI & GROGAN, 1987: 40 (female; Mexico). Distr.- Mexico (Oaxaca). panamensis WIRTH, 1965b: 225 (male, female; Panama). Distr.- Panama. pseudunguis SPINELLI & GROGAN, 1987: 30 (male, female; Panama). Distr.- Panama. raccurti SPINELLI & GROGAN, 1987: 42 (male, female; El Salvador). Distr.- El Salvador, northern Argentina. spangleri WIRTH, 1965b: 227 (female; Puerto Rico). Distr.- Puerto Rico. uncinata (JOHANNSEN), 1943: 761 (Stilobezzia; female; USA, Alabama). Distr.- USA (Alabama, Maryland, Virginia, Arkansas), Jamaica. unguis WIRTH, 1965b: 228 (male, female; USA, Arizona). Distr.- USA (Arizona, Texas) to Colombia. wirthi CLASTRIER & RACCURT, 1979a: 169 (female; Haiti). Distr.- Haiti. Genus PARASTILOBEZZIA WIRTH & BLANTON PARASTILOBEZZIA WIRTH & BLANTON, 1970a: 10. Type species: Parastilobezzia leei WIRTH & BLANTON, by original designation. leei WIRTH & BLANTON, 1970a: 10 (male, female; Colombia). Distr.- Costa Rica, Colombia. Genus RHYNCHOHELEA WIRTH & BLANTON RHYNCHOHELEA WIRTH & BLANTON, 1970c: 96. Type species: Rhynchohelea monilicornis WIRTH & BLANTON, by original designation.

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monilicornis WIRTH & BLANTON, 1970c: 98 (female; USA, Florida). Distr.- USA (Florida), Costa Rica. Genus SCHIZONYXHELEA CLASTRIER SCHIZONYXHELEA CLASTRIER, 1984: 1. Type species: Schizonyxhelea guyana CLASTRIER, by original designation. forattinii WIRTH & GROGAN, 1988: 81 (male, female; Brazil). Distr.- Costa Rica, Brazil (Santa Catarina). guyana CLASTRIER, 1984: 2 (female; French Guiana). Distr.- Mexico (Yucatán), French Guiana, Brazil (Mato Grosso). Genus STILOBEZZIA KIEFFER STILOBEZZIA KIEFFER, 1911a: 118. Type species: Stilobezzia festiva KIEFFER, by original designation. HARTOMYIA MALLOCH, 1915a: 339. Type species: Ceratopogon pictus COQUILLETT (= Stilobezzia coquilletti KIEFFER), by original designation. REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM &MACFIE, 1931 (key males and females from Patagonia and southern Chile); LANE & FORATTINI, 1961 (key to males and females from Neotropical Region north of Patagonia and southern Chile); WIRTH & GROGAN, 1988 (key to subgenera, as part of generic key). Subgenus ACANTHOHELEA KIEFFER ACANTHOHELEA KIEFFER, 1917a: 198. Type species: Acanthohelea pruinosa KIEFFER, by monotypy. amazonica CLASTRIER, 1991: 306 (male, female; French Guiana). Distr.- French Guiana, Peru. atrichopogon LANE & FORATTINI, 1956: 208 (male, female; Panama). Distr.- Panama. bicinctipes INGRAM & MACFIE, 1931: 195 (female; Chile). Distr.- Southern Chile. edwardsi INGRAM & MACFIE, 1931: 198 (male, female; Argentina). Distr.- Southwestern Argentina. furva INGRAM & MACFIE, 1931: 200 (female; Argentina). Distr.- Southern Chile, southwestern Argentina. guianae (MACFIE), 1940d: 28 (Acanthohelea; male, female; Guyana). Distr.- Guyana. hirsuta INGRAM & MACFIE, 1931: 201 (male, female; Argentina). Distr.- Southwestern Argentina. maia LANE & FORATTINI, 1958: 204 (male, female; Panama). Distr.- Panama, Colombia. manaosensis LANE & FORATTINI, 1958: 205 (male, female; Brazil). Distr.- Brazil (Amazonas). nigerrima INGRAM & MACFIE, 1931: 196 (female; Argentina). Distr.- Southwestern Argentina. ornata LANE & FORATTINI, 1958: 206 (female; Panama). Distr.- Panama. ornaticrus INGRAM & MACFIE, 1931: 194 (female; Chile). Distr.- Southern Chile. patagonica INGRAM & MACFIE, 1931: 196 (male, female; Argentina). Distr.- Southwestern Argentina.

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rava INGRAM & MACFIE, 1931: 203 (male, female; Chile). Distr.- Southern Chile. sanctibernardini KIEFFER, 1917b: 308 (female; Paraguay). Distr.- Paraguay. succinea INGRAM & MACFIE, 1931: 200 (male, female; Argentina). Distr.- Chile, southwestern Argentina south to Tierra del Fuego. tibialis LANE & FORATTINI, 1956: 209 (male, female; Panama). Distr.- Panama. varia INGRAM & M ACFIE , 1931: 191 (male, female; Chile). Distr.- Southern Chile, southwestern Argentina south to Tierra del Fuego. Subgenus EUKRAIOHELEA INGRAM & MACFIE EUKRAIOHELEA INGRAM & MACFIE, 1921: 347. Type species: Eukraiohelea africana INGRAM & MACFIE, designation by MACFIE, 1940f: 22. REFERENCE: WIRTH & SPINELLI, 1992b (key males and females). amnigena (MACFIE), 1935a: 56 (Eukraiohelea; female; Brazil). Distr.- USA (Florida, South Carolina), (?) Jamaica, (?) Puerto Rico, Brazil (Maranhão), northeastern Argentina. dorsofasciata (LUTZ), 1914: 96 (Palpomyia; female; Brazil). Distr.- Brazil (Rio de Janeiro). elegantula (JOHANNSEN), 1907: 109 (Bezzia; female; USA, Kansas). Distr.- USA (Kansas to Maryland to Louisiana and Florida), Mexico (Yucatán), Panama, (?) Puerto Rico, Paraguay, Argentina. maculitibia LANE & FORATTINI, 1956: 207. Panama. subsessilis KIEFFER, 1917b: 311. Paraguay. Subgenus STILOBEZZIA KIEFFER albicoxa LANE & FORATTINI, 1956: 210 (male, female; Panama). Distr.- Panama. albocincta, KIEFFER 1917b: 309 (female; Paraguay). Distr.- Paraguay. americana KIEFFER, 1917b: 310 (female; Paraguay). Distr.- Paraguay. antennalis (COQUILLETT), 1901a: 606 (Ceratopogon; female; USA, District of Columbia). Distr.- North America (British Columbia to Ontario to Texas and Florida), Panama. beckae WIRTH, 1953a: 69 (male, female; USA, Florida). Distr.- USA (Maryland to Mississippi, Florida), Mexico (Yucatán), Panama, Peru. bicolor LANE, 1947a: 208 (male; Brazil). Distr.- Panama, Brazil (São Paulo), Argentina (Buenos Aires). bimaculata LANE & FORATTINI, 1956: 211 (male, female; Panama). Distr.-Panama, Trinidad, Brazil (Rio de Janeiro). bispinosa KIEFFER, 1917b: 310 (female; Paraguay). Distr.- Brazil (São Paulo), Paraguay. blantoni LANE & FORATTINI, 1956: 211 (male, female; Panama). Distr.- Panama. caribe LANE & FORATTINI, 1958: 208 (male; Panama). Distr.- Panama. chaconi MACFIE, 1938: 166 (male; Trinidad). Distr.- Costa Rica, Trinidad, Brazil (Goiás, São Paulo). coquilletti KIEFFER, 1917b: 308. (New name for pictus COQUILLETT, female; USA, Virginia). Distr.- USA (Virgina, Illinois, Maryland), Mexico (Yucatán, Chiapas) to Panama, Trinidad, Brazil (São Paulo).

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picta (COQUILLETT), 1905: 60 (Ceratopogon) (preoccupied by Sphaeromias pictus (MEIGEN), 1818). USA (Virginia). coracina KIEFFER, 1917b: 311 (male; Paraguay). Distr.- Brazil (São Paulo), Paraguay, Argentina (Buenos Aires). diminuta LANE & FORATTINI, 1958: 209 (male, female; Panama). Distr.- Panama, Dominica. dryadum MACFIE, 1940b: 186 (female; Guyana). Distr.- Panama, Guyana. dubitans LANE, FORATTINI & RABELLO, 1955: 85 (pupa, male; Brazil). Distr.- Brazil (São Paulo). dureti LANE & FORATTINI, 1958: 210 (male; Brazil). Distr.- Brazil (São Paulo). esmeralda LANE & FORATTINI, 1958: 211 (male, female; Panama). Distr.- Panama. femoralis LANE & FORATTINI, 1956: 212 (male; Panama). Distr.- Mexico (Yucatán), Panama. fiebrigi KIEFFER, 1917b: 309 (female; Paraguay). Distr.- Brazil (São Paulo, Rio de Janeiro), Paraguay, Argentina (Buenos Aires province). glauca MACFIE, 1939b: 204 (male; Brazil). Distr.- USA (Maryland to Mississippi and Florida), Mexico (Chiapas) to southeastern Brazil. fluminensis LANE, 1947a: 210. Brazil. grandis LANE & FORATTINI, 1958: 213 (female; Panama). Distr.- Panama. kiefferi LANE, 1947a: 205 (female; Brazil). Distr.- USA (Florida), Mexico (Yucatán, Tamaulipas), Panama, Brazil (Minas Gerais). punctipes WIRTH, 1953a: 79. USA (Florida). macfiei LANE, 1947a: 213 (male, female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro). maculata LANE, 1947a: 207 (male; Brazil). Distr.- Mexico (Yucatán), Panama, Venezuela, Brazil (Rio de Janeiro). merceri CAZORLA & SPINELLI, in CAZORLA et al., 2005: 290 (male, female, pupa; Peru). Distr.- Peru. modesta LANE, 1947a: 206 (female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro). nigroflava LANE & FORATTINI, 1958: 215 (male, female; Costa Rica). Distr.- Costa Rica, Panama, Puerto Rico. obscura LANE & FORATTINI, 1958: 216 (male, female; Panama). Distr.- Panama. pallescens LANE & FORATTINI, 1958: 218 (male, female; Panama). Distr.- Panama, Brazil (São Paulo). panamensis LANE & FORATTINI, 1958: 218 (male, female; Panama). Distr.- Panama, Puerto Rico, Brazil (São Paulo, Rio de Janeiro). paulistensis LANE, 1947a: 200 (male, female; Brazil). Distr.- Panama, USA (Puerto Rico), Brazil (São Paulo, Rio de Janeiro). punctulata LANE, 1947a: 204 (female; Brazil). Distr.- Mexico (Tabasco), Panama, Perú, Brazil (Mato Grosso, Rio de Janeiro). rabelloi LANE, 1947a: 203 (male, female; Brazil). Distr.- USA (Maryland to Louisiana and Florida) to Argentina. silvicola MACFIE, 1940b: 185 (male; Guyana). Distr.- Guyana. similans LANE & FORATTINI, 1956: 214 (male, female; Panama). Distr.- Panama. simplex LANE & FORATTINI, 1958: 222 (male, female; Panama). Distr.- Panama. thomsenae WIRTH, 1953a: 83 (male, female; USA, Florida). Distr.- USA (Florida), Mexico (Yucatán), Panama. scutata LANE & FORATTINI, 1961: 92. Panama. transversa LANE & FORATTINI, 1958: 222 (male, female; Panama). Distr.- Panama.

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travassosi LANE, 1947a: 210 (female; Brazil). Distr.- Brazil (Espírito Santo). venezuelensis ORTÍZ, 1950d: 199 (as variety of glauca; female; Venezuela). Distr.- Venezuela. williamsi CAZORLA & SPINELLI, in CAZORLA et al., 2005: 292 (male, female; Peru). Distr.- Peru. wirthi LANE & FORATTINI, 1956: 214 (male, female; Panama). Distr.- Panama. wygodzinskyi LANE, 1947a: 212 (female; Brazil). Distr.- Panama, Brazil (São Paulo). TRIBE HETEROMYIINI WIRTH, 1962: 275 Genus CLINOHELEA KIEFFER CLINOHELEA KIEFFER, 1917b: 295. Type species: Ceratopogon variegatus WINNERTZ (= Ceratopogon unimaculata MACQUART), by original designation. Subgenus CERATOBEZZIA KIEFFER CERATOBEZZIA KIEFFER, 1917b: 326. Type species: Ceratobezzia fallax KIEFFER, by original designation. fallax KIEFFER, 1917b: 326 (female; Paraguay). Distr.- El Salvador to Colombia, Venezuela, Brazil (Amazonas, Pará, Mato Grosso), Paraguay, northeastern Argentina. barrettoi LANE & DURET, 1954: 249. Brazil. Subgenus CLINOHELEA KIEFFER REFERENCE: SPINELLI & DURET, 1993 (key to males and females). albopennis LANE, 1944: 259 (female; Brazil). Distr.- Brazil (Pará, São Paulo), Paraguay, northeastern Argentina. argentina LANE & DURET, 1954: 248 (female; Argentina). Distr.- Northeastern Argentina. damascenoi LANE & DURET, 1954: 250 (female; Brazil). Distr.- Peru, Bolivia, French Guiana, Brazil (Pará), Paraguay. horacioi LANE, 1944: 257 (male, female; Brazil). Distr.- Mexico (Yucatán, Tabasco, Morelos) to northeastern Argentina. muzoni SPINELLI & DURET, 1993: 46 (male, female; Colombia). Distr.- Colombia. neivai LANE, 1944: 252 (male, female; Brazil). Distr.- Panama, Brazil (Goiás, Minas Gerais, São Paulo, Rio de Janeiro), Paraguay. nigripes MACFIE, 1939b: 205 (female; Brazil). Distr.- Costa Rica to Colombia, Brazil (São Paulo, Santa Catarina), northeastern Argentina. rubriceps KIEFFER, 1917b: 318 (female; Paraguay). Distr.- Paraguay. saltanensis LANE & DURET, 1954: 252 (female; Argentina). Distr.- Brazil (Rondônia, Santa Catarina), Paraguay, northern Argentina. townsendi LANE, 1944: 256 (female; Brazil). Distr.- Panama to Argentina (Corrientes, Buenos Aires province). townesi LANE, 1944: 254. Brazil.

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wygodzinskyi LANE, 1948: 231 (female; Brazil). Distr.- Brazil (Rio de Janeiro). Genus HETEROMYIA SAY HETEROMYIA SAY, 1825: plate 35. Type species: Heteromyia fasciata SAY, by monotypy. PACHYLEPTUS WALKER, 1856: 426. Type species: Pachyleptus fasciatus WALKER (= Heteromyia nigra KIEFFER), by monotypy. REFERENCE: DURET & LANE, 1955 (key to males and females). antequerae (LYNCH ARRIBÁLZAGA), 1893: 227 (Pachyleptus; female; Argentina). Distr.Northeastern Argentina. bejaranoi DURET & LANE, 1955: 36 (male, female; Argentina). Distr.- Northeastern Argentina. castanea LANE, 1946a: 214 (female; Brazil). Distr.- Brazil (São Paulo). chaquensis DURET & LANE, 1955: 37 (female; Argentina). Distr.- Northeastern Argentina. clavata WILLISTON, 1900: 225 (female; Mexico). Distr.- Mexico (Veracruz) to Colombia, Brazil (Amazonas), northeastern Argentina. rufa KIEFFER, 1917b: 325. Colombia. caloptera KIEFFER, 1919b: 192. Unnecessary new name for H. rufa KIEFFER, not LOEW, 1861. correntina DURET & LANE, 1955: 39 (male, female; Argentina). Distr.- Panama, Brazil (Rondônia), northeastern Argentina. kiefferi LANE, 1946a: 213 (female; Brazil). Distr.- Brazil (Rio de Janeiro). lamprogaster EDWARDS, 1933: 87 (female; Argentina). Distr.- Northeastern Argentina. nigra KIEFFER, 1917b: 326 (female; Paraguay). Distr.- Brazil (São Paulo, Rio de Janeiro, Paraná), Paraguay. fasciatus (WALKER), 1856: 426 (Pachyleptus) (preoccupied by Heteromyia fasciata SAY, 1825). South America. orellana (ROBACK), 1957: 1 (Palpomyia; female; Peru). Distr.- Peru. wokei WIRTH & GROGAN, 1977: 181 (pupa, male, female; Nicaragua). Distr.- Nicaragua. Genus PELLUCIDOMYIA MACFIE PELLUCIDOMYIA MACFIE, 1939a: 99. Type species: Pellucidomyia ugandae MACFIE, by original designation. MACFIEHELEA LANE, 1946a: 208. Type species: Macfiehelea oliveirai LANE, by original designation. REFERENCE: WIRTH, 1960 (World key to males and females). blantoni (LANE), 1956b: 435 (Macfiehelea; female; Panama). Distr.- Panama. lanei WIRTH & RATANAWORABHAN, 1971b: 446 (female; Colombia). Distr.- Colombia, Jamaica. oliveirai (LANE), 1946a: 209 (Macfiehelea; female; Brazil). Distr.- Brazil (Mato Grosso, Minas Gerais). wirthi (LANE), 1956b: 437 (Macfiehelea; female; Panama). Distr.- USA (Texas) to Colombia.

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Genus PHYSOHELEA GROGAN & WIRTH REFERENCE: SPINELLI, 1994 (description of male P. turgidipes). PHYSOHELEA GROGAN & WIRTH, 1979a: 53. Type species: Neurohelea oedidactyla INGRAM & MACFIE, by original designation. oedidactyla (INGRAM & MACFIE ), 1931: 212 (Neurohelea; female; Argentina). Distr.Southwestern Argentina. turgidipes (INGRAM & MACFIE), 1931: 214 (Neurohelea; female; Chile). Distr.- Southern Chile. TRIBE SPHAEROMIINI NEWMAN, 1834: 388 Genus AUSTROSPHAEROMIAS SPINELLI AUSTROSPHAEROMIAS SPINELLI, 1997a: 224. Type species: Austrosphaeromias apricans (INGRAM & MACFIE), by original designation. apricans (INGRAM & MACFIE), 1931: 219 (Palpomyia; male, female; Argentina). Distr.Southwestern Argentina, southern Chile. sentior (INGRAM & MACFIE), 1931: 222 (Palpomyia, as variety of apricans). Chile. wirthi SPINELLI, 1997a: 228 (female; Argentina). Distr.- Southwestern Argentina. Genus GROGANHELEA SPINELLI & DIPPOLITO GROGANHELEA SPINELLI & DIPPOLITO, in SPINELLI et al., 1995: 166. Type species: Groganhelea rondoniensis SPINELLI & DIPPOLITO, by original designation. rondoniensis SPINELLI & DIPPOLITO, in SPINELLI et al., 1995: 167 (female; Brazil). Distr.Brazil (Rondônia), Uruguay. Genus JOHANNSENOMYIA MALLOCH JOHANNSENOMYIA MALLOCH, 1915a: 332. Type species: Johannsenomyia halteralis MALLOCH (= Ceratopogon argentata LOEW), designation by WIRTH, 1952a: 211. DICROHELEA KIEFFER, 1917b: 363. Type species: Palpomyia filicornis KIEFFER, designation by MACFIE, 1940f: 26. blantoni (LANE & WIRTH ), 1961: 81 (Dicrohelea; female; Panama). Distr.- Panama. lanei WIRTH, 1965a: 4 (female; Brazil). Distr.- Brazil (Santa Catarina). Genus LANEHELEA WIRTH & BLANTON LANEHELEA WIRTH & BLANTON, 1972a: 433. Type species: Lanehelea leei WIRTH and BLANTON, by original designation.

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leei WIRTH & BLANTON, 1972a: 434 (female; Colombia). Distr.- Colombia. spinifemur WIRTH & BLANTON, 1972a: 436 (female; Colombia). Distr.- Colombia. Genus MALLOCHOHELEA WIRTH MALLOCHOHELEA WIRTH, 1962: 278. Type species: Johannsenomyia albibasis MALLOCH, by original designation. aenipes (MACFIE), 1940c: 193 (Bezzia; female; Guyana). Distr.- Guyana limitrofe SPINELLI & FELIPPE-BAUER, 1990a: 87 (female; Argentina). Distr.- Northeastern Argentina, Uruguay. nemoralis (MACFIE), 1940c: 190 (Johannsenomyia; male; Guyana). Distr.- Guyana. nigripes (MACFIE), 1939b: 216 (Johannsenomyia; female; Brazil). Distr.- Brazil (Santa Catarina). pullata (WIRTH), 1952a: 213 (Johannsenomyia; female; USA, California). Distr.- USA (California, Arizona, Texas) to Brazil (Amapá). bicellii (LANE), 1961b: 449 (Nilobezzia). Brazil. termophila (SPINELLI), 1984: 197 (Neobezzia; male, female; Argentina). Distr.- Brazil (Rondônia), northern Argentina. Genus NEOBEZZIA WIRTH & RATANAWORABHAN NEOBEZZIA WIRTH & RATANAWORABHAN, 1972a: 477. Type species: Neobezzia clavipes WIRTH & RATANAWORABHAN, by original designation. REFERENCE: WIRTH & RATANAWORABHAN, 1972a (key to females). albitarsis WIRTH & RATANAWORABHAN, 1972a: 478 (female; Brazil). Distr.- Brazil (Pará). amnicola (MACFIE), 1940d: 30 (Bezzia; female; Guyana). Distr.- Belize to Brazil (Amazonas, Rondônia, Mato Grosso, São Paulo). blantoni WIRTH & RATANAWORABHAN, 1972a: 482 (female; Panama). Distr.- Nicaragua to Colombia, Brazil (Amazonas, Pará, Rondônia), northeastern Argentina. brasiliae (LANE), 1961a: 37 (Macropeza; female; Brazil). Distr.- Brazil (Brasilia, D.F.). clavipes WIRTH & RATANAWORABHAN, 1972a: 485 (female; Brazil). Distr.- Panama, Brazil (Amazonas). costaricae WIRTH & RATANAWORABHAN, 1972a: 486 (female; Costa Rica). Distr.- Costa Rica, Panama. fittkaui WIRTH & RATANAWORABHAN, 1972a: 489 (female; Brazil). Distr.- Brazil (Amazonas). wirthi SPINELLI & FELIPPE-BAUER, 1990a: 88 (male, female; Argentina). Distr.- Northeastern Argentina. Genus NILOBEZZIA KIEFFER NILOBEZZIA KIEFFER, 1921a: 24. Type species: Nilobezzia armata KIEFFER, by monotypy. REFERENCE: LANE, 1961b (key to males and females).

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neotropica (MACFIE), 1940a: 78 (Bezzia; female; Brazil). Distr.- Brazil (Rondônia, Bahia). paraensis (LANE), 1958: 33 (Bezzia; female; Brazil). Distr.- Brazil (Pará). schwarzii (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Texas). Distr.- USA (South Carolina to Texas and Florida), Mexico (Yucatán), Bahamas, Cuba, Panama, Brazil (Rio de Janeiro), northeastern Argentina. banksi (GERRY), 1933: 94 (Bezzia). Cuba. brasiliensis (LANE), 1958: 28 (Bezzia). Brazil. Genus SPHAEROHELEA SPINELLI & FELIPPE-BAUER SPHAEROHELEA SPINELLI & FELIPPE-BAUER, 1990b: 195. Type species: Sphaerohelea biestroi SPINELLI & FELIPPE-BAUER, by original designation. biestroi SPINELLI & FELIPPE-BAUER, 1990b: 197 (female; Argentina). Distr.- Northeastern Argentina. TRIBE PALPOMYIINI ENDERLEIN, 1936: 49 Genus AMEROHELEA GROGAN & WIRTH AMEROHELEA GROGAN & WIRTH, 1981: 1280. Type species: Amerohelea galindoi GROGAN & WIRTH, by original designation. REFERENCES: GROGAN & WIRTH, 1981 (key to females); HUERTA, 1996 (key to females from Mexico). dalcyi GROGAN & WIRTH, 1981: 1289 (female; Brazil). Distr.- Brazil (Amazonas). fasciata GROGAN & WIRTH, 1981: 1283 (male, female; Belize). Distr.- Mexico (Sonora, Morelos, Tabasco, Chiapas) to Colombia. frontispina (DOW & TURNER), 1976: 138 (Bezzia; female; USA, Texas). Distr.- USA (California to Texas) south to Colombia and Venezuela. galindoi GROGAN & WIRTH , 1981: 1294 (male, female; Colombia). Distr.- Panama, Colombia,Venezuela, northern Argentina. nelsoni GROGAN & WIRTH, 1981: 1290 (female; Brazil). Distr.- Brazil (Amazonas). pseudofasciata GROGAN & WIRTH, 1981: 1286 (female; Brazil). Distr.- Southeastern Brazil, Argentina (Misiones, Río Negro). ronderosi GROGAN & WIRTH, 1981: 1298 (male, female; Colombia). Distr.- Colombia. similis SPINELLI, 1989: 27 (female; Uruguay). Distr.- Northern Argentina, Uruguay. sordidipes (MACFIE), 1939b: 209 (Palpomyia; female; Brazil). Distr.- Bolivia, Brazil (Pará, Minas Gerais, São Paulo, Santa Catarina). spinellii GROGAN & WIRTH, 1981: 1292 (female; Colombia). Distr.- Colombia. vargasi GROGAN & WIRTH, 1981: 1296 (male, female; Belize) Distr.- Mexico (Veracruz, Oaxaca), Belize to Costa Rica.

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Genus BEZZIA KIEFFER BEZZIA KIEFFER, 1899: 69. Type species: Ceratopogon ornatus MEIGEN, by original designation. REFERENCES: HUERTA, 1996 (key to males and females from Mexico); SPINELLI & WIRTH, 1989a (key to subgenera and species groups for males and females, key to males and females of nobilis and punctipennis groups); SPINELLI & WIRTH, 1989b (key to females and some males of glabra and brevicornis groups); SPINELLI & WIRTH, 1990 (key to males and females of gibbera group); SPINELLI & WIRTH, 1991 (key to males and females of dentifemur and venustula groups). aitkeni SPINELLI & WIRTH, 1991: 5 (female; Brazil). Distr.- Brazil (Pará). albuquerquei LANE, 1961a: 43 (female; Brazil). Distr.- Brazil (Brasilia, D.F.). araucana SPINELLI & WIRTH, 1990: 14 (male, female; Argentina). Distr.- Southwestern Argentina. bivittata (COQUILLETT), 1905: 60 (Ceratopogon; female; USA, California). Distr.- North America (Alaska to Nova Scotia to California and Florida), Panama. blantoni SPINELLI & WIRTH, 1989b: 771 (pupa, male, female; Honduras). Distr.- Belize to Costa Rica, Puerto Rico, Argentina (Corrientes, Buenos Aires). brevicornis (KIEFFER), 1917b: 328 (Allobezzia; male, female; Paraguay). Distr.- Paraguay, Argentina (Corrientes, Entre Ríos, Buenos Aires, Río Negro), Uruguay. bromeliae SPINELLI & WIRTH, 1991: 9 (pupa, male, female; Panama). Distr.- Panama. capitata WIRTH & GROGAN, 1983: 503 (male; Honduras). Distr.- USA (Arizona) to Costa Rica. carioca LANE, 1958: 30 (female; Brazil). Distr.- Brazil (Rio de Janeiro). catarinensis SPINELLI & WIRTH, 1990: 17 (male, female; Brazil). Distr.- Brazil (Santa Catarina) cayoensis SPINELLI & WIRTH, 1991: 6 (male, female; Belize). Distr.- Belize. chilensis SPINELLI & RONDEROS, 2001: 752 (Male, female; Chile). Distri.- Chile, Argentina (Salta, Rio Negro). clavipennis SPINELLI & WIRTH, 1989b: 775 (female; Brazil). Distr.- Colombia, Ecuador, Venezuela, Brazil (Amazonas, Rondônia). dentifemur SPINELLI & WIRTH, 1991: 3 (male, female; Colombia). Distr.- Belize, Colombia, Trinidad, Brazil (Pará). filiducta SPINELLI & WIRTH, 1991: 3 (as filiductus; female; Colombia). Distr.- Honduras, Colombia. flinti SPINELLI & WIRTH, 1989a: 113 (male, female; Dominica). Distr.- Dominica. fluminensis LANE, 1948: 236 (female; Brazil). Distr.- Brazil (Brasilia, D.F., Rio de Janeiro), Argentina (Corrientes). fusca SPINELLI & WIRTH, 1991: 9 (female; Colombia). Distr.- Colombia. gibbera (COQUILLETT), 1905: 60 (Ceratopogon; female; Cuba). Distr.- USA (Arizona to Florida) to Panama, Cuba, Jamaica, Tobago. glabra (COQUILLETT), 1902: 85 (Ceratopogon; female; USA, Florida). Distr.- North America (Alaska to New Brunswick to California and Florida), Belize, El Salvador. globulosa SPINELLI & WIRTH, 1990: 18 (male, female; Puerto Rico). Distr.- Puerto Rico. goianensis LANE, 1961a: 44 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).

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grogani SPINELLI & WIRTH, 1990: 19 (male, female; Panama). Distr.- Mexico (Morelos), Panama, Colombia. hondurensis SPINELLI & WIRTH, 1990: 20 (male, female; Honduras). Distr.- Mexico (Veracruz, Chiapas), Honduras, El Salvador, Panama. jamaicensis SPINELLI & WIRTH, 1989a: 115 (male, female; Jamaica). Distr.- Jamaica, Haiti. jubata SPINELLI & WIRTH, 1990: 20 (male, female; Colombia). Distr.- Colombia. leei SPINELLI & WIRTH, 1990: 21 (male, female; Colombia). Distr.- Colombia. lenkoi LANE, 1958: 31 (male, female; Brazil). Distr.- Panama, Brazil (Amazonas, Pará, Brasilia, D.F., São Paulo, Rio de Janeiro). mathisi SPINELLI & WIRTH, 1989a: 117 (female; Ecuador). Distr.- Ecuador. mazaruni MACFIE, 1940c: 193 (female; Guyana). Distr.- USA (California, Florida), El Salvador, Venezuela, Guyana, Caribbean to Brazil (Amazonas, Rondônia). megatheca SPINELLI & WIRTH, 1990: 22 (female; Colombia). Distr.- Colombia, Brazil (Rondônia). mesotibialis SPINELLI & WIRTH, 1990: 23 (female; Belize). Distr.- Belize, Trinidad. nigritibialis SPINELLI & WIRTH, 1991: 5 (female; Belize). Distr.- Belize, Costa Rica. nobilis (WINNERTZ), 1852: 79 (Ceratopogon; Germany). Distr.- Palaearctic, North America (Alaska to Nova Scotia) to southern Argentina. acanthodes MACFIE, 1940c: 192. Guyana. atlantica WIRTH & WILLIAMS, 1957: 13. Bermuda. pseudogibbera SPINELLI & WIRTH, 1990: 23 (male, female; Panama). Distr.- Belize, Honduras, Panama. pseudovenustula SPINELLI & WIRTH, 1991: 8 (female; Brazil). Distr.- Brazil (Pará). pulchripes KIEFFER, 1917b: 330 (female; Paraguay). Distr.- Widespread, from Mexico (Tabasco) to northeastern Argentina. punctipennis (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- USA (California, Nevada, Arizona, Texas, Florida), Mexico (Yucatán, Tabasco, Quintana Roo), El Salvador, Honduras, Jamaica, Dominica, Brazil (Amazonas), Argentina (Buenos Aires province). raposoensis SPINELLI & WIRTH, 1991: 4 (male, female; Colombia). Distr.- Mexico (Yucatán), Colombia. roldani SPINELLI & WIRTH, 1981: 187 (larva, pupa, male, female; Argentina). Distr.- Eastern Argentina (Entre Rios, Buenos Aires). schmitzorum DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 54 (female; Brazil). Distr.Brazil (Rondônia). setigera SPINELLI & WIRTH, 1990: 24 (male, female; Colombia). Distr.- El Salvador, Colombia. snowi LANE, 1958: 34 (male, female; Guatemala). Distr.- Guatemala, Panama, Colombia, Trinidad, Brazil (Pará). subfusca MACFIE, 1939b: 218 (female; Brazil). Distr.- Brazil (Santa Catarina). venustula (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- Widespread, from Mexico (Veracruz, Morelos, Chiapas) to Argentina. concoloripes MACFIE, 1940d: 31. Guyana. woodruffi SPINELLI & WIRTH, 1989a: 120 (female; Jamaica). Distr.- Jamaica.

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Genus CLASTRIEROMYIA SPINELLI & GROGAN CLASTRIEROMYIA SPINELLI & GROGAN, 1985: 330. Type species: Clastrieromyia schnacki SPINELLI & GROGAN, by original designation. REFERENCE: SPINELLI & GROGAN, 1986 (key to females). dycei SPINELLI & GROGAN, 1986: 456 (pupa, male, female; Uruguay). Distr.- Uruguay, eastern and central Argentina. kremeri SPINELLI & GROGAN, 1985: 332 (female; Brazil). Distr.- Brazil (Amazonas). schnacki SPINELLI & GROGAN, 1985: 331 (female; Ecuador). Distr.- Ecuador, northern Argentina. uruguayensis SPINELLI & GROGAN, 1986: 458 (female; Uruguay). Distr.- Northeastern and central Argentina, Uruguay. Genus PACHYHELEA WIRTH PACHYHELEA W IRTH , 1959a: 50. Type species: Ceratopogon magnus C OQUILLETT (= Ceratopogon pachymerus WILLISTON), by original designation. albidiventris (KIEFFER), 1917b: 316 (Sphaeromias; female; Colombia). Distr.- Colombia. pachymera (WILLISTON), 1900: 224 (Ceratopogon; female; Mexico). Distr.- USA (Texas) to central Argentina. latifemoris (INGRAM & MACFIE), 1931: 231 (Johannsenomyia). Argentina. Genus PALPOMYIA MEIGEN PALPOMYIA MEIGEN, 1818: 82. Type species: Ceratopogon flavipes MEIGEN, by monotypy. Generic name first published in synonymy with Ceratopogon but available under ICZN Code Article 11(e). REFERENCES: INGRAM & MACFIE, 1931 (key males and females from Patagonia and southern Chile); LANE, 1960 (key to females). aculeata INGRAM & MACFIE, 1931: 215 (female; Chile). Distr.- Southern Chile. almeidai (LANE), 1946b: 219 (Sphaeromias; female; Brazil). Distr.- Brazil (São Paulo). barrettoi LANE, 1947b: 442 (female; Brazil). Distr.- Brazil (Rio de Janeiro). boliviensis KIEFFER, 1917b: 324 (female; Bolivia). Distr.- Bolivia. brasiliensis MACFIE, 1939b: 213 (female; Brazil). Distr.- Brazil (Minas Gerais, Santa Catarina). callangana KIEFFER, 1917b: 323 (female; Peru). Distr.- Peru. carioca LANE, 1960: 384 (Brazil; female). Distr.- Brazil (Rio de Janeiro). carrerai (LANE), 1948: 235 (Dicrobezzia, as carreirai; female; Brazil). Distr.- Brazil (São Paulo). castanea MACFIE, 1939b: 211 (female; Brazil). Distr.- Brazil (São Paulo, Santa Catarina). catarinensis LANE, 1960: 385 (female; Brazil). Distr.- Brazil (Santa Catarina).

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chilensis INGRAM & MACFIE, 1931: 218 (female; Chile). Distr.- Southern Chile, ? Brazil (São Paulo). columbiana KIEFFER, 1917b: 323 (female; Colombia). Distr.- Colombia. conifera MACFIE, 1939b: 214 (male; Brazil). Distr.- Brazil (Brasilia, D.F., Goiás, Santa Catarina). coroicoensis WIRTH, 1974: 54. New name for boliviensis. Distr.- Bolivia. boliviensis (KIEFFER), 1917b: 331 (Jenkinsia; female) (preoccupied by Palpomyia boliviensis KIEFFER, 1917b). Bolivia. crassicrus KIEFFER, 1917b: 321 (female; Paraguay). Distr.- Brazil (São Paulo), Paraguay. guarani LANE, 1946b: 221 (pupa, male, female; Brazil). Distr.- Brazil (São Paulo, Espírito Santo, Rio de Janeiro), Paraguay. guyana CLASTRIER, 1992: 117 (male, female; French Guiana). Distr.- French Guiana. hispida BORKENT, in BORKENT & WIRTH, 1997: 132. New name for multispinosa. Distr.- El Salvador to Colombia. multispinosa SPINELLI & GROGAN, 1989: 6 (male, female) (preoccupied by Palpomyia multispinosa (PIERCE), 1966). El Salvador. iberaensis SPINELLI & CAZORLA, 2006: 1068 (female, male genitalia; northeastern Argentina). Distr.- Argentina. inermicollis KIEFFER, 1917b: 322 (female; Paraguay). Distr.- Paraguay. insularis SPINELLI & GROGAN, 1989: 3 (male, female; Puerto Rico). Distr.- Jamaica, Dominica, Puerto Rico. johannseni (LANE), 1948: 233 (Dicrobezzia; female; Brazil). Distr.- Brazil (São Paulo). lacustris LANE, FORATTINI & RABELLO, 1955: 83 (pupa, male; Brazil). Distr.- Brazil (São Paulo). limnochares (MACFIE), 1940a: 76 (Sphaeromias; male; Brazil). Distr.- Northeastern Brazil. lutzi LANE, 1947b: 440 (female; Brazil). Distr.- Brazil (Rio de Janeiro). mellichroa MACFIE, 1939b: 212 (female; Brazil). Distr.- Brazil (Santa Catarina). microchela KIEFFER, 1917b: 321 (female; Colombia). Distr.- Colombia. nigroflava LANE, 1947b: 446 (female; Brazil). Distr.- Brazil (São Paulo). nigroscutellata LANE, 1947b: 444 (female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro). oliveirai LANE, 1947b: 443 (female; Brazil). Distr.- Brazil (Rondônia, Minas Gerais). pampana LANE, 1960: 386 (female; Brazil). Distr.- Brazil (Rio Grande do Sul). paraensis LANE, 1960: 387 (female; Brazil). Distr.- Brazil (Pará). patagonica INGRAM & MACFIE, 1931: 222 (female; Argentina). Distr.- Southwestern Argentina. paulistensis LANE, 1947b: 445 (female; Brazil). Distr.- Panama, Brazil (São Paulo). pseudolacustris DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 55 (male, female; Brazil). Distr.- Brazil (Rondônia). scabra (COQUILLETT), 1905: 62 (Ceratopogon; female; Mexico). Distr.- Mexico (Tabasco), Argentina (Buenos Aires province). spinifemur (LANE), 1948: 234 (Dicrobezzia; male, female genitalia; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro). spinosa LUTZ, 1914: 93 (female; Brazil). Distr.- Brazil (Rio de Janeiro). subaspera (COQUILLETT), 1901a: 606 (Ceratopogon; female; USA, New Mexico). Distr.- North America (Alberta to Ontario to California, Florida), Mexico (Chihuahua), Argentina. maculicrus INGRAM & MACFIE, 1931: 230. Argentina.

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subfuscula INGRAM & MACFIE, 1931: 216 (male, female; Chile). Distr.- Southern Chile, southwestern Argentina. tamioi LANE, 1960: 388 (female; Brazil). Distr.- Brazil (Rio de Janeiro). tanycornis BORKENT, in BORKENT & WIRTH, 1997: 134. New name for longicornis. Distr.- St. Vincent; Brazil (Brasilia, D.F.). longicornis (WILLISTON), 1896: 280 (Ceratopogon; female) (preoccupied by Ceratopogon longicornis WALTL, 1837). St. Vincent. tenuicrus KIEFFER, 1917b: 320 (female; Paraguay). Distr.- Paraguay. umbella MACFIE, 1939b: 208 (female; Brazil). Distr.- Brazil (Mato Grosso, Rio de Janeiro, Santa Catarina, Rio Grande do Sul). versicolor MACFIE, 1939b: 215 (female; Brazil). Distr.- Brazil (Rondônia, Santa Catarina). wirthi LANE, FORATTINI & RABELLO, 1955: 82 (pupa, female; Brazil). Distr.- Brazil (São Paulo). Genus PHAENOBEZZIA HAESELBARTH PHAENOBEZZIA HAESELBARTH, 1965: 297. Type species: Probezzia pistiae INGRAM & MACFIE, by original designation. REFERENCE: SPINELLI & WIRTH, 1986a (key to males and females). ateles (MACFIE), 1940a: 78 (Bezzia; male; Brazil). Distr.- Colombia, Ecuador, Brazil (Bahia). astyla SPINELLI & WIRTH, 1986a: 232. Colombia. maya SPINELLI & WIRTH, 1986a: 234 (male, female; Belize). Distr.- USA (Texas) to Brazil (Rondônia). TRIBE STENOXENINI COQUILLETT, 1899: 61 Genus PARYPHOCONUS ENDERLEIN PARYPHOCONUS ENDERLEIN , 1912: 57. Type species: Paryphoconus angustipennis ENDERLEIN, by original designation. REFERENCES: HUERTA, 1996 (key to females from Mexico); SPINELLI & WIRTH, 1984b (key to females). aemulus MACFIE, 1940b: 180 (female; Guyana). Distr.- Peru, Guyana, Brazil (Amazonas, Mato Grosso). amapaensis LANE, 1961b: 450 (female; Brazil). Distr.- Brazil (Amapá). angustipennis ENDERLEIN, 1912: 57 (female; Brazil). Distr.- Colombia, Peru, Venezuela, Brazil (Goiás, São Paulo, Santa Catarina), northeastern Argentina. anomalicornis KIEFFER, 1917b: 333 (female; Colombia and Paraguay). Distr.- Mexico San Luis Potosi) to Colombia, Venezuela, Brazil (Amazonas), Paraguay, northeastern Argentina. apicalis SPINELLI & WIRTH, 1984b: 889 (female; Ecuador). Distr.- Panama, Ecuador. barrettoi LANE, 1946a: 203 (female; Brazil). Distr.- Brazil (Goiás, São Paulo).

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batesi LANE, 1961c: 455 (female; Brazil). Distr.- Brazil (Brasilia, D.F.). brunneipennis SPINELLI & WIRTH, 1984b: 890 (male, female; Brazil). Distr.- Colombia, Brazil (Amazonas). ecuadorensis SPINELLI & WIRTH, 1984b: 892 (female; Ecuador). Distr.- Ecuador. enderleini LANE, 1956a: 301 (female; Brazil). Distr.- Brazil (Mato Grosso). fittkaui SPINELLI & WIRTH, 1984b: 892 (female; Brazil). Distr.- Brazil (Amazonas). flavidus (JOHANNSEN), 1943: 761 (Ceratobezzia; female; Guyana). Distr.- Guyana, Brazil (Amazonas, Mato Grosso). lanei WIRTH, 1959b: 236. Brazil. flinti SPINELLI & WIRTH, 1984b: 894 (female; Mexico). Distr.- Mexico (Tabasco, Chiapas), Belize. fusciradialis SPINELLI & WIRTH, 1984b: 895 (female; Brazil). Distr.- Brazil (Pará). fuscus LANE, 1946a: 206 (female; Brazil). Distr.- Brazil (Goiás). goianensis LANE, 1961c: 456 (female; Brazil). Distr.- Brazil (Brasilia, D.F.). grandis MACFIE, 1939c: 6 (female; Brazil). Distr.- Mexico (on train quarantined in Arizona), Belize, Venezuela, Brazil (Pará, Santa Catarina). guianae MACFIE, 1940b: 180 (female; Guyana). Distr.- Ecuador, Guyana, Brazil (Pará, Mato Grosso, São Paulo). kiefferi LANE, 1956a: 302 (female; Brazil). Distr.- Brazil (Goiás). latipennis SPINELLI & WIRTH, 1984b: 898 (male, female; Colombia). Distr.- Colombia, Venezuela. leei SPINELLI & WIRTH, 1984b: 898 (female; Colombia.). Distr.- Colombia. macfiei LANE, 1946a: 203 (female; Brazil). Distr.- Ecuador, Venezuela, Brazil (Pará, Mato Grosso, São Paulo). maya SPINELLI & WIRTH, 1984b: 899 (female; Belize). Distr.- Mexico (San Luis Potosi), Belize. mayeri WIRTH, 1959b: 236 (female; Brazil). Distr.- Brazil (Amazonas, Rondônia). misionensis SPINELLI, 1998: 52 (female; Argentina). Distr.- Northeastern Argentina. neotropicalis (LANE), 1948: 229 (Macropeza; female; Brazil). Distr.- Brazil (São Paulo). nigripes MACFIE, 1939c: 8 (female; Argentina). Distr.- Guyana, Brazil (Pará, Rondônia), northeastern Argentina. nubifer MACFIE, 1939c: 5 (female; Brazil). Distr.- Bolivia, Brazil (Amazonas, Rio de Janeiro, Santo Catarina), Paraguay, northeastern Argentina, Uruguay. oliveirai LANE, 1956a: 303 (female; Brazil). Distr.- Colombia, Brazil (Amazonas, Pará). paranaensis SPINELLI & WIRTH, 1984b: 902 (female; Argentina). Distr.- Northeastern Argentina. paulistensis LANE, 1961c: 457 (female; Brazil). Distr.- Brazil (São Paulo). sonorensis WIRTH & RATANAWORABHAN, 1972b: 1374 (female; Mexico). Distr.- USA (Oklahoma), Mexico (Sonora), Belize, Panama. steineri SPINELLI & WIRTH, 1984b: 905 (female; Peru). Distr.- Panama, Peru. subflavus MACFIE, 1940d: 23 (female; Guyana). Distr.- Colombia, Guyana, Brazil (Amazonas, Pará, Rondônia). travassosi LANE, 1956a: 304. Brazil. taragui SPINELLI, 1998: 53 (female; Argentina). Distr.- Northeastern Argentina. telmatophilus (MACFIE), 1940a: 77 (Bezzia; male; Brazil). Distr.- Northeastern Brazil. terminalis (COQUILLETT), 1904a: 90 (Ceratopogon; female; Nicaragua). Distr.- Belize to Peru and Ecuador, Brazil (Amazonas, Santa Catarina). unimaculatus MACFIE, 1940b: 179 (female; Guyana). Distr.- Guyana, Brazil (Pará).

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wirthi LANE, 1961c: 458 (female; Brazil). Distr.- Brazil (Brasilia, D.F.). wygodzinskyi LANE, 1946a: 208 (female; Brazil). Distr.- Brazil (Rio de Janeiro). Genus STENOXENUS COQUILLETT STENOXENUS COQUILLETT, 1899: 61. Type species: Stenoxenus johnsoni COQUILLETT, by monotypy. REFERENCE: WIRTH & RATANAWORABHAN, 1972b (key to females). aductus DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 56 (female; Brazil). Distr.- Brazil (Rondônia). arcuatus WIRTH & RATANAWORABHAN, 1972b: 1379 (female; Panama). Distr.- Panama. blantoni WIRTH & RATANAWORABHAN, 1972b: 1379 (female; Panama). Distr.- Panama. brasiliensis MACFIE, 1939b: 140 (female; Brazil). Distr.- Brazil (Amazonas, Santa Catarina), northeastern Argentina. carrerai LANE, 1956a: 305 (as carreirai; female; Brazil). Distr.- Brazil (Goiás). dimorphus KIEFFER, 1909: 47 (male, female; Peru). Distr.- Peru. excentricus LANE, 1961c: 453 (female; Brazil). Distr.- Brazil (Pará). fulvus JOHANNSEN, 1927: 70 (female; Peru). Distr.- Peru. johnsoni COQUILLETT, 1899: 61 (female; USA, New Jersey). Distr.- USA (Missouri, New Jersey, Texas) to Panama. lanei WIRTH & RATANAWORABHAN, 1972b: 1381 (female; Panama). Distr.- Panama. limpidus WIRTH & RATANAWORABHAN, 1972b: 1383 (female; Costa Rica). Distr.- Costa Rica. marginalis WIRTH & RATANAWORABHAN, 1972b: 1385 (female; Panama). Distr.- Panama. niger LANE, 1948: 230 (as nigrus; male, female; Brazil). Distr.- Brazil (Goiás). paraensis LANE, 1956a: 306 (female; Brazil). Distr.- Brazil (Pará). pseudofulvus SPINELLI, 1998: 54 (female; Argentina). Distr.- Northeastern Argentina, Uruguay. setiger MACFIE, 1939b: 138 (female; Brazil). Distr.- Jamaica, Dominican Republic, Brazil (Santa Catarina), northeastern Argentina. Unplaced genus of Forcipomyiinae DIDYMOPHLEPS WEYENBERGH , 1883: 108. Type species: Didymophleps hortorum WEYENBERGH, by monotypy. Unplaced species of Forcipomyiinae Didymophleps hortorum WEYENBERGH, 1883: 110. Argentina. Unplaced species of Ceratopogonidae Ceratopogon claripennis LYNCH ARRIBÁLZAGA, 1893: 229. Argentina.

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7 IDENTIFICATION BORKENT & WIRTH (1997) listed the living and extinct species of biting midges for the World and an updated version is available on the web (http://www.inhs.uiuc.edu/cee/FLYTREE/) CeratopogonidaeCatalog.pdf). BORKENT & SPINELLI (2000) provide a catalog of the ceratopogonids south of the USA which included those species from the Nearctic portion of Mexico. Additional information was given concerning which life stages were originally described and the distribution of each of the 1066 mentioned species. References to all pertinent taxonomic literature was provided for each genus, including available keys. Here, we provide an updated catalog of those species restricted to the Neotropical Region, as defined by GRIFFITHS (1980) (Table 4). The number of species in each genus is given in Table 5. WIRTH & GROGAN (1988) provided a revision of the tribe Ceratopogonini, the most diverse tribe at the generic level. This revision included a key to the genera and subgenera, and a diagnosis, distribution and list of included species for each genus. The best work for identifying Neotropical Culicoides is a wing atlas published by WIRTH et al. (1988), which contains wing photographs and a summary of the numerical characters of the majority of the species presently known for the region. This work may be used together with the revision of the genus for Panama by WIRTH & BLANTON (1959), and the keys for the Amazon Basin by WIRTH & BLANTON (1973), for the species occurring south of the Amazon Basin by SPINELLI & WIRTH (1986b), and for Argentina (SPINELLI et al., 2005b). Description of further species may be found in SPINELLI & BORKENT (2004a) for Costa Rica, BRICKLE & HAGAN (1999) for Belize, FELIPPE-BAUER & QUINTELAS (1994) and FELIPPE-BAUER et al. (2000a, b) for Brazil, FELIPPE-BAUER et al. (2003, 2005) for Peru and PERRUOLO (1990, 2001) for Venezuela. Adult ceratopogonids are easily collected with a variety of methods and are best preserved in 70% alcohol. These specimens can be identified to genus while in fluid but generally need to be placed on microscope slides to be identified to species. Specimens that are not slide mounted should be pinned, as old alcohol material is impossible to slide mount and study properly. For more information, see chapter ‘Capture and study of Ceratopogonidae’.

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Table 5. Numbers of named valid species in each genus of Neotropical Ceratopogonidae. Taxon Subfamily Leptoconopinae NOÈ Leptoconops SKUSE Subfamily Forcipomyiinae LENZ Atrichopogon KIEFFER Forcipomyia MEIGEN Subfamily Dasyheleinae LENZ Dasyhelea KIEFFER Subfamily Ceratopogoninae NEWMAN Tribe Culicoidini KIEFFER Culicoides LATREILLE Paradasyhelea MACFIE Tribe Ceratopogonini Allohelea KIEFFER Alluaudomyia KIEFFER Austrohelea WIRTH & GROGAN Baeodasymyia CLASTRIER & RACCURT Baeohelea WIRTH & BLANTON Bahiahelea WIRTH Borkenthelea SPINELLI & GROGAN Brachypogon KIEFFER Cacaohelea WIRTH & GROGAN Ceratoculicoides WIRTH & RATANAWORABHAN Diaphanobezzia INGRAM & MACFIE Downeshelea WIRTH & GROGAN Echinohelea MACFIE Fittkauhelea WIRTH & BLANTON Isthmohelea INGRAM & MACFIE Leptohelea WIRTH & BLANTON Macrurohelea INGRAM & MACFIE Monohelea KIEFFER Nannohelea GROGAN & WIRTH Notiohelea GROGAN & WIRTH Parabezzia MALLOCH Parastilobezzia WIRTH & BLANTON Rhynchohelea WIRTH & BLANTON Schizonyxhelea CLASTRIER Stilobezzia KIEFFER Tribe Heteromyiini WIRTH Clinohelea KIEFFER Heteromyia SAY Pellucidomyia MACFIE

Number of species 12 100 211 58

266 3 1 19 1 5 1 1 4 25 1 1 4 18 11 1 1 1 12 18 1 2 24 1 1 2 65 12 11 4

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Table 5. Continued. Taxon

Number of species

Physohelea GROGAN & WIRTH Tribe Sphaeromiini NEWMAN Austrosphaeromias SPINELLI Groganhelea SPINELLI & DIPPOLITO Johannsenomyia MALLOCH Lanehelea WIRTH & BLANTON Mallochohelea WIRTH Neobezzia WIRTH & RATANAWORABHAN Nilobezzia KIEFFER Sphaerohelea SPINELLI & FELIPPE-BAUER Tribe Palpomyiini ENDERLEIN Amerohelea GROGAN & WIRTH Bezzia KIEFFER Clastrieromyia SPINELLI & GROGAN Pachyhelea WIRTH Palpomyia MEIGEN Phaenobezzia HAESELBARTH Tribe Stenoxenini COQUILLETT Paryphoconus ENDERLEIN Stenoxenus COQUILLETT Unplaced genus Didymophleps WEYENBERGH Unplaced species Total number of species (not including unplaced taxa)

DIAGNOSIS

OF THE

2 2 1 2 2 6 8 3 1 11 46 4 2 46 2 40 16 1 2 1095

CERATOPOGONIDAE

The following distinguishes Ceratopogonidae from all other Diptera: small-medium sized biting midges with wing length = 0.4-7.0 mm. Ocelli absent. Antenna generally with 13 flagellomeres (some have fewer but always 6 or more present). Most males with plumose antennae. Most females with serrate mandibles. Anepisternum divided into anterior and posterior portions by anepisternal cleft. Postnotum lacking medial longitudinal groove. Wing with 1-2 radial veins reaching margin, with two median vein branches (posterior one may be weak) reaching wing margin. Wings overlapping each other over the abdomen in living individuals at rest (with exceptions in a few species of Stilobezzia). Foreleg shorter than hind leg. Tarsomere 1 of each leg equal to or longer than tarsomere 2 (with a few exceptions in Forcipomyia). Adults are similar to some Chironomidae but may be distinguished by the combination of a lack of a medial groove on the postnotum (otherwise only in Podonominae) and a lack of a m-cu crossvein (present in Podonominae).

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Pupae with well-developed, undivided respiratory organs with series of small spiracles. Third leg curled under wing sheath. Apex of the abdomen not curled under thorax. Two pointed anal processes. Larvae with well-developed head capsule, well-developed pharyngeal complex, and no open spiracles. Eggs are not diagnosable at this time (but see key below).

KEY

TO THE GENERA OF ADULT

CERATOPOGONIDAE

IN THE

NEOTROPICAL REGION

There are currently 1095 species of Ceratopogonidae recorded from the Neotropical Region (Table 5) but there are many more awaiting description, especially in the genera Forcipomyia, Atrichopogon, Dasyhelea, Culicoides and Stilobezzia. Species of the family may be found at all altitudes and in virtually every habitat where there is even a little moisture. The following key includes genera known from the Neotropics and includes males, for the first time, of all genera (males not known for Groganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, and Sphaerohelea). This key is intended to work for both alcohol and slide mounted material. We have indicated in bold type those character states that are best seen with a compound microscope (i.e. with slide mounted material), although with skill and patience many of these character states may also be observed in alcohol specimens. Some wing veins may be difficult to see in some taxa; for alcohol specimens, tilting the wing may help to see faint wing veins and with slide material, phase contrast microscopy provides the best resolution. When the two claws at the end of a leg are the same size, they are referred to as being equal; conversely when they differ in size they are called unequal. The costal ratio is the length of the costa from the wing base (measured from the arculus), divided by the total wing length. Terms for wing veins and cells follow the system of the Manual of Nearctic Diptera (MCALPINE et al., 1981), with modifications proposed by SZADZIEWSKI (1996) and are fully labeled here in Figs. 10C, D and 11I. As is widely used by other Dipterists, veins are in capital letters and those of cells in small letters. 1

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Eyes widely separated dorsally, lacking frontal suture (Fig. 6A); palpus with 4 segments (only one beyond elongate and/or swollen third) and with membranous gap between segment 3 and 4+5 (Fig. 9A); wing without crossvein r-m (Fig. 10A); female with radial cells fused into swollen unit (Fig. 10A); without palisade setae on first tarsomere of hind leg (Fig. 14B); male gonostylus with an apical, socketed peg (Fig. 19A); female cercus very elongate (Fig. 22E) ........................... ................................................................................................................... Leptoconops SKUSE Eyes narrowly or widely separated dorsally, with frontal suture (Figs. 6B-G); palpus with 2-5 segments but never with a membranous gap between the last two segments (Figs. 9B-E); wing with crossvein r-m (Figs. 10B-J); radial cells distinct or, if fused or absent (Figs. 11A, F), r-m clearly present; with (Fig. 14A) or without (Fig. 14B) palisade setae on first tarsomere of hind leg; male gonostylus without apical, socketed peg (Figs. 19B-F, 20A-F) although apex may be pointed (Fig. 19E); female cercus short (Figs. 22F, 23A-E) .............................................................. 2

104

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales Frontal suture

Palpus Lacinia Mandible Labrum Labium

A. Leptoconops B. Culicoides ‡

C. Culicoides †

Frontal suture

Clypeus Labium

D. Rhynchohelea

E. Echinohelea

F. Parabezzia

G. Fittkauhelea

Figure 6. Head capsules in anterior view. A: Female Leptoconops floridensis WIRTH (Nearctic Region). B: Female Culicoides yukonensis HOFFMAN, mouthparts spread laterally (Nearctic Region). C: Male Culicoides yukonensis, mouthparts spread laterally (Nearctic Region). D: Female Rhynchohelea monilicornis WIRTH & BLANTON (Nearctic and Neotropical Regions). E: Female Echinohelea sp. F: Female Parabezzia sp. G: Female Fittkauhelea amazonica WIRTH & BLANTON (Neotropical Region). (AC from MCALPINE et al., 1981; D-G courtesy of INBio, Costa Rica).

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Wing with (Figs. 10B, E, G) or without (Figs. 10D, J, 11A-C) numerous macrotrichia on membrane; hind first tarsomere with scattered setae or, at most, a row of strong bristles (but these not forming a tight row, especially basally on the tarsomere) (Fig. 14B); female with two small, equal claws on each leg (Figs. 14CE); with (Figs. 14C, D) or without (Figs. 14E, F) well-developed empodia .......... 3 Wing bare or with scattered macrotrichia on apical half of wing (a few species have some macrotrichia more basally on the wing but these are scattered) (Figs. 11E, I); hind first tarsomere with row of palisade setae with at least more basal setae with abutting bases, some groups with an additional pronounced basal stout spine (Figs. 14A, G-I); females with two small, equal claws on each leg or with 12 claws on one or more of the legs which, in some, are large (Figs. 14F-I, 15A-C, G, H, 16A-E); without well-developed empodia ......................................................... 7 Costa reaching well beyond middle of wing (Figs. 10C, D); both radial cells welldeveloped, with second radial cell elongate and moderately open (Figs. 10C, D); paratergite well-developed, with 1 strong and 0-5 more slender, shorter setae (Fig. 9F) ...................................................................................................... Atrichopogon KIEFFER Costa short or long; if long, then second radial cell either wide (Figs. 10F, G) or narrow (Figs. 10B, E); paratergite narrow (as in Fig. 9G) or well-developed, without setae or, if setae present, these number at least 10 and all are equal in size and length ............................................................................................................................ 4 Apical flagellomere with terminal nipple which is constricted basally (Figs. 7A, B); claws strongly curved (Figs. 14C, D); empodia vestigial or well-developed (Figs. 14C, D) ..................................................................................... Forcipomyia MEIGEN Apical flagellomere without terminal nipple (although it may be strongly tapered) (Fig. 7C); claws only slightly curved (Fig. 14E); empodia poorly developed ....... 5 Eyes broadly abutting medially; antennal flagellomeres without sensilla coeloconica; male with short setae on first flagellomere, less than 1/2 the length of those on following flagellomeres (Fig. 7C); male flagellomeres sculptured (Fig. 7C); without tibial spur on foreleg; wing membrane without distinct pattern of dark pigmentation (Fig. 10E) .......................... Dasyhelea KIEFFER Eyes contiguous to widely separated medially (Figs. 6B, C); some antennal flagellomeres with sensilla coeloconica; male with setae on first flagellomere about as long as those on following flagellomeres; male flagellomeres not sculptured; with tibial spur on foreleg; wing membrane with or without distinct patterns of dark and/or light pigmentation (Figs. 10F-H); CULICOIDINI .................................................................................................................. 6 Wing with 1-2 radial cells; costa extending beyond 0.50 of wing length (10F, G); females with or without toothed mandibles; wing membrane with or without distinct patterns of dark and/or light pigmentation (10F, G) ................................... ............................................................................................................... Culicoides LATREILLE Wing with both radial cells obliterated (Fig. 10H); costa not reaching beyond 0.43 of wing length; females with reduced, vestigial mandibles; wing membrane without distinct patterns of dark and/or light pigmentation (Fig. 10H) ................. .............................................................................................................. Paradasyhelea MACFIE

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A. Forcipomyia ‡

B. Forcipomyia ‡

C. Dasyhelea †

D. Rhynchohelea ‡ Figure 7. Antennae. A: Female Forcipomyia glauca MACFIE (Palaearctic, Nearctic and Neotropical Regions). B: Female Forcipomyia monilicornis (COQUILLETT) (Holarctic Region). C: Male Dasyhelea pseudoincisurata WAUGH and WIRTH (Nearctic Region). D: Female Rhynchohelea monilicornis WIRTH & BLANTON (Nearctic and Neotropical Regions) (all from MCALPINE et al., 1981).

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M forking beyond r-m (i.e. medial fork petiolate, in some, with base quite short (Fig. 11I), in some, base of M2 not present (Figs. 10I, J, 11A) ................................ 8 – M forking at or before r-m (i.e. medial fork sessile, not petiolate) (Figs. 11G, 12FJ, 13A-I); base of M2 always present ............................................................................ 52 8(7) Wing with reduced radial cells (Figs. 11A, B, D, F); costal ratio less than 0.65 (Figs. 11A, B, D, F); female claws more or less equal (the male of Rhynchohelea is unknown but likely with broadly separated eyes, antennal flagellomeres 10 and 11 fused and a katepisternal seta) ........................................................................................ 9 – Wing with at least one well-developed radial cell (Figs. 10J, 11E, I); costal ratio more than 0.50; female claws equal or unequal, some with only a single claw on one or more of the legs .................................................................................................. 15 9(8) Male antennal flagellomeres 10 and 11 fused or with less than 13 flagellomeres; wing with or without radial cells; katepisternum with 1 or more lateral setae (can be seen in alcohol specimens by holding specimen upright and looking along plane of thoracic pleura in outline) (Fig. 9G) .................................. 10 – Male antenna with flagellomeres 10 and 11 separate and with 13 flagellomeres present; wing without radial cell (cells completely fused) (Figs. 11A, B, D); katepisternum without setae (Fig. 9H) ............................................. 13 10(9) Female antenna with 12 flagellomeres (Fig. 7D); female labium stout, truncate, with apical hooklike structures (Fig. 6D) ........................................................................ ........................................................... Rhynchohelea WIRTH & BLANTON (male unknown) – Female antenna with 13 flagellomeres; female proboscis normal (Figs. 6B, C, E-G) .............................................................................................................................................. 11 11(10) Male flagellum with 7-8 flagellomeres (Fig. 8A); palpus with 3 segments (Fig. 9C) ............................................................................................. Nannohelea GROGAN & WIRTH – Male flagellum with at least 9 flagellomeres (almost always with 13 but with 10 and 11 fused); palpus with 5 segments ........................................................................ 12 12(11) Anepisternum lacking setae; male antenna with flagellomeres 2-6 fused (6-11 also fused) ......................................................................... Brachypogon KIEFFER (in part) – Anepisternum with at least 1 seta on posterior margin (Fig. 9G); male antenna with flagellomeres 2-5 articulated, not fused (flagellomeres 7-11 fused, sometimes including flagellomere 6) ........................ Ceratoculicoides WIRTH & RATANAWORABHAN 13(9) Palpus with 5 segments; wing length 0.7-0.8 mm; male with well-developed parameres .................................................................................... Schizonyxhelea CLASTRIER – Palpus with 2 segments (Fig. 9B); wing length < 0.6 mm; male without parameres .............................................................................................................................................. 14 14(13) Fore tibia with apical spur; macrotrichia confined to margin and apex of wing (in some, also a few on apices of M1 and M2); costal ratio > 0.40 (Fig. 11B); male antenna with 6 separate flagellomeres (Fig. 8C); male with abdominal tergites 1-3 pale and 4-8 brown ........................................................... Baeohelea WIRTH & BLANTON – Fore tibia without apical spur; macrotrichia long and abundant over entire wing; costal ratio < 0.36; male antenna with 12 flagellomeres but with 2-7, 8-9, and 10-11 fused (Fig. 8B); male with abdomen uniformly brown ............................ ................................................................................... Baeodasymyia CLASTRIER & RACCURT

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A. Nannohelea †

B. Baeodasymyia †

C. Baeohelea † Sensilla coeloconica

D. Borkenthelea †

Figure 8. Antennae. A: Male Nannohelea bourioni (CLASTRIER) (Palaearctic Region). B: Male Baeodasymyia modesta CLASTRIER and RACCURT (Neotropical Region). C: Male Baeohelea nana WIRTH & BLANTON (Neotropical Region). D: Male Borkenthelea harii (Neotropical Region). (A-B from WIRTH & GROGAN, 1988; C courtesy of INBio, Costa Rica).

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Fused 4+5

B. Baeohelea

C. Nannohelea

D. Leptohelea

A. Leptoconops

E. Parabezzia Paratergite

Anepisternum

Katepisternum

F. Atrichopogon

G. Ceratoculicoides

Forecoxa

Midcoxa

H. Parabezzia Figure 9. Female structures. A: Leptoconops bequaerti (KIEFFER) right palpus (Nearctic and Neotropical Regions). B: Baeohelea nana WIRTH & BLANTON right palpus (Neotropical Region). C: Nannohelea sp. right palpus. D: Leptohelea micronyx WIRTH & BLANTON right palpus (Neotropical Region). E: Parabezzia sp. right palpus. F: Atrichopogon sp. thorax, in lateral view. G: Ceratoculicoides sp. thorax, in lateral view. H: Parabezzia bystraki GROGAN & WIRTH thorax, in lateral view (Nearctic Region). (A, E courtesy of INBio, Costa Rica; D from WIRTH & GROGAN, 1988; F from SZADZIEWSKI et al., 1997).

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Wing with characteristic pattern of darkly mottled pigmentation, more or less scattered on membrane (Fig. 12C); thorax and at least hind femur and tibia with patterned pigmentation; male aedeagus with distinctive posteromedial loop (Fig. 21A) ....................................................................................................... Monohelea KIEFFER1 Wing without pigmentation or, if present, restricted to radial sector or with discrete dots, stripes or bands (Figs. 11E, H), or if with substantial pattern of pigmentation (Fig. 11H) then thorax and all femora and tibiae uniformly dark brown; other taxa with variably pigmented thorax and legs; aedeagus, if present, without posteromedial loop ........................................................................................... 16 Costa extending nearly to tip of wing; female wing with costa extending beyond apex of second radial cell (Figs. 12B, D, E) (extension also in male wing of Bahiahelea and Leptohelea) ................................................................................................. 17 Costa not extending beyond 0.85 of wing length (Figs. 11E, I); female wing with costa ending at or extending slightly past apex of second radial cell ................... 25 Halter saclike, without constriction below knob (Fig. 13J) ......................................... ...................................................................................... Diaphanobezzia INGRAM & MACFIE Halter with knob and distinct stem .............................................................................. 18 Male ..................................................................................................................................... 19 Female ................................................................................................................................. 21 Wing with costa not extending beyond apex of second radial cell .......................... ...................................................................................... Parastilobezzia WIRTH & BLANTON Wing with costa extending well beyond apex of second radial cell ..................... 20 With flagellomeres 10-13 more elongate than preceding flagellomeres; antennal plume well developed; parameres fused into large, single sphere (Fig. 19F) .......... .................................................................................................................... Bahiahelea WIRTH With flagellomeres 2-13 more or less equal in length; antennal plume short; parameres elongate, slender ........................................... Leptohelea WIRTH & BLANTON Palpus with three segments, with last segment small and short (Fig. 9D); claws small, equal, without basal inner tooth (Fig. 15D) ................. Leptohelea WIRTH & BLANTON Palpus with 4-5 segments, with last segment elongate (Fig. 9E); claws variable ....... 22 Clypeus not fused to head (as in Fig. 6B); wing with first radial cell small, second elongate (Fig. 12E); with single claw (with thick basal tooth) on each leg (Fig. 15E) ...................................................................................... Parastilobezzia WIRTH & BLANTON Clypeus laterally fused to head (Figs. 6F, G); wing with 1 radial cell (Fig. 12B); with 2 claws on each leg, although hind leg claws may be unequal ............................... 23 Eyes moderately separated, bare (Fig. 6F); claws without basal inner tooth .......... .............................................................................................................. Parabezzia MALLOCH Eyes broadly separated, bare or pubescent (Fig. 6G); claws with basal inner tooth .............................................................................................................................................. 24 Palpus with 4 segments (Fig. 6G); eyes pubescent; spermathecae without distinct necks .............................................................. Fittkauhelea WIRTH & BLANTON

M. fairchildi LANE & WIRTH does not key out well. Adults have banded wings, as in Downeshelea, but the male has an aedeagus with a distinctive posteromedial loop.

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A. Leptoconops r-m

B. Forcipomyia

c

First radial cell

R3

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Second radial cell r3 m1

M1 A

M2 CuP

CuA2

CuA1

C. Atrichopogon

E. Dasyhelea

G. Culicoides

I. Cacaohelea

anal

m2 cua1

D. Atrichopogon

F. Culicoides

H. Paradasyhelea

J. Brachypogon

Figure 10. Female wings. A: Leptoconops sp. B: Forcipomyia fairfaxensis WIRTH (Nearctic Region). C: Atrichopogon levis (COQUILLETT) (Nearctic Region). D: Atrichopogon sp. E: Dasyhelea pseudoincisurata WAUGH & WIRTH (Nearctic Region). F: Culicoides insignis LUTZ (Nearctic and Neotropical Regions). G: Culicoides copiosus ROOT & HOFFMAN (Nearctic Region). H: Paradasyhelea ingrami (Neotropical Region) I: Cacaohelea youngi WIRTH & GROGAN (Neotropical Region). J: Brachypogon stigmalis (COQUILLETT). (A-G, J, from MCALPINE et al., 1981; I from WIRTH & GROGAN, 1988).

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A. Rhynchohelea

B. Baeohelea

C. Leptohelea

D. Schizonyxhelea

F. Brachypogon E. Alluaudomyia

G. Echinohelea R1

R2

I. Stilobezzia

H. Allohelea R3

J. Borkenthelea

Figure 11. Female wings. A: Rhynchohelea monilicornis WIRTH & BLANTON (Nearctic and Neotropical Regions). B: Baeohelea nana WIRTH & BLANTON (Neotropical Region). C: Leptohelea micronyx WIRTH & BLANTON (Neotropical Region). D: Schizonyxhelea forattinii WIRTH & GROGAN (Neotropical Region). E: Alluaudomyia bella (COQUILLETT) (Nearctic Region). F: Brachypogon paraensis WIRTH & BLANTON (Neotropical Region). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Allohelea johannseni (WIRTH) (Nearctic Region). I: Stilobezzia lutea (MALLOCH) (Nearctic Region). J: Borkenthelea nothofagus SPINELLI & WIRTH (Neotropical Region). (A, E, G, I from MCALPINE et al., 1981; C, D, FH from WIRTH & GROGAN, 1988).

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A. Austrohelea

B. Parabezzia

C. Monohelea

D. Bahiahelea

E. Parastilobezzia

F. Downeshelea

G. Physohelea

H. Clinohelea

I. Paryphoconus

J. Stenoxenus †

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Figure 12. Wings. A: Female Austrohelea shannoni WIRTH & GROGAN (Neotropical Region). B: Female Parabezzia petiolata MALLOCH (Nearctic Region). C: Female Monohelea hieroglyphica KIEFFER (Neotropical Region). D: Female Bahiahelea brasiliensis WIRTH (Neotropical Region). E: Female Parastilobezzia leei WIRTH & BLANTON (Neotropical Region). F: Female Downeshelea sp. G: Female Physohelea oedidactyla (INGRAM & MACFIE) (Neotropical Region). H: Female Clinohelea bimaculata (LOEW) (Nearctic Region). I: Female Paryphoconus angustipennis ENDERLEIN (Neotropical Region). J: Male Stenoxenus coomani SÉGUY (Oriental Region) (A, E from WIRTH & GROGAN, 1988; B-C, H-J from MCALPINE et al., 1981).

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C. Bezzia

D. Phaenobezzia

E. Neobezzia

F. Groganhelea

G. Mallochohelea

H. Amerohelea

J. Diaphanobezzia I. Clastrieromyia Figure 13. Female wings. A: Stenoxenus coomani SÉGUY (Oriental Region). B: Palpomyia plebeja (LOEW) (Nearctic Region). C: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). D: Phaenobezzia pistiae (INGRAM & MACFIE) (Afrotropical Region). E: Neobezzia blantoni WIRTH & RATANAWORABHAN (Neotropical Region). F: Groganhelea rondoniensis SPINELLI & DIPPOLITO (Neotropical Region). G: Mallochohelea smithi (Lewis) (Nearctic Region). H: Amerohelea sp. (Neotropical Region) I: Clastrieromyia schnacki SPINELLI & GROGAN (Neotropical Region). J: Halter Diaphanobezzia araucaria SPINELLI (Neotropical Region) (A-D, G from MCALPINE et al., 1981; H from GROGAN & WIRTH, 1981; F from SPINELLI & DIPPOLITO, 1995; I from SPINELLI & GROGAN, 1986; E from WIRTH & RATANAWORABHAN, 1972a; J from SPINELLI, 1996).

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Palpus with 5 segments; eyes bare; spermathecae with distinct slender necks .................................................................................................................... Bahiahelea WIRTH Male ..................................................................................................................................... 26 Female ................................................................................................................................. 39 Hind leg with single long claw and small basal tooth (Fig. 14H) .............................. ................................................................................................................... Allohelea KIEFFER Hind leg with two equal or subequal claws ................................................................ 27 Katepisternum with 1 or more lateral setae (can be seen in alcohol specimens by holding specimen upright and looking along plane of thoracic pleura in outline) (Fig. 9G) ............................................................... Brachypogon KIEFFER (in part) Katepisternum without setae .................................................................................... 28 Wing with 1 radial cell (Fig. 11E) ................................................................................. 29 Wing with 2 radial cells ................................................................................................... 33 Fore coxa with long spine-like or slender, tapering setae (Fig. 9H) .......................... .............................................................................................................. Parabezzia MALLOCH Fore coxa with only short, simple setae (sometimes rubbed off in alcohol specimens) ......................................................................................................................... 30 Wing with pigmentation spots at least immediately basal to or over r-m and over apex of radial cell (Fig. 11E), some with further pigmentation ................................ ............................................................................................................ Alluaudomyia KIEFFER Wing without pigmentation spots ................................................................................ 31 Palpus with 5 segments; hind tarsomere 1 with at least 1 1/2 rows of palisade setae; aedeagus composed of 2 lateral sclerites (so that aedeagus appears to be split medially) (Fig. 20E) or with aedeagus absent but with a transverse bar ventral to the aedeagus (Fig. 20D) ..................... Stilobezzia KIEFFER (in part) Palpus with 4 segments (as in Fig. 9E); hind tarsomere 1 with only 1 row of palisade setae; aedeagus continuous medially (as in Fig. 21C) ......................... 32 Eyes without pubescence; genitalia with fused, very short, parameres ............ .................................................................. undescribed genus near Parabezzia MALLOCH Eyes pubescent; genitalia without parameres ....................................................... .......................................................................................... Fittkauhelea WIRTH & BLANTON Wing with patches of pigmentation, including discrete lighter patches on M2 and CuA2 (Fig. 12F); aedeagus continuous medially (Fig. 20F) ................................... ............................................................................................ Downeshelea WIRTH & GROGAN Wing with or without pattern of pigmentation but if pattern present, then without discrete patches on M2 and CuA2 (posterior portion of wing may be generally more darkly pigmented) (Figs. 11J, 12A) and with aedeagus divided medially (Figs. 20B, C) .................................................................................................. 34 Antenna with apical 5 flagellomeres more elongate than basal 8 flagellomeres (female-like antenna), without plume .......................... Cacaohelea WIRTH & GROGAN Antenna with apical 3 flagellomeres more elongate than basal 10 flagellomeres (male-like antenna), with or without plume ............................................................... 35 Radial cells separated by a short longitudinal vein (R2); wing without macrotrichia; sternite 9 elongate (Fig. 20C) ......................................... Isthmohelea INGRAM & MACFIE

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Both radial cells close to one another, separated by a short transverse vein (R2) (Figs. 11J, 12A); wing with or without macrotrichia; sternite 9 short or moderately elongate (Figs. 20D, E) .................................................................................................... 36 Flagellomeres 3-10 fused (Fig. 8D) ........................ Borkenthelea SPINELLI & GROGAN Antenna with flagellomeres separate, not fused ........................................................ 37 Length of tarsomeres 2-5 of hind leg about 1.4-1.7as long as length of tarsomere 1 .......................................................................................... Austrohelea WIRTH & GROGAN Length of tarsomeres 2-5 of hind leg about as long as tarsomere 1 ................... 38 Hind tarsomere 1 with at least 1 1/2 rows of palisade setae; wing with or without macrotrichia; aedeagus composed of 2 lateral sclerites (so that aedeagus appears to be split medially) (Fig. 20E) or with aedeagus absent but with a transverse bar ventral to the aedeagus (Fig. 20D); antennal flagellomeres without sensilla coeloconica ........................................................ Stilobezzia KIEFFER Hind tarsomere 1 with only 1 row of palisade setae (as in Fig. 14A); wing without macrotrichia; aedeagus undivided, somewhat triangular; some antennal flagellomeres with sensilla coeloconica ............. Macrurohelea INGRAM & MACFIE Katepisternum with 1 or more lateral setae (can be seen in alcohol specimens by holding specimen upright and looking along plane of thoracic pleura in outline) (Fig. 9G) ............................................................... Brachypogon KIEFFER (in part) Katepisternum without seta ...................................................................................... 40 Wing with 1 radial cell (Figs. 11C, E) .......................................................................... 41 Wing with 2 radial cells (first may be small) (Figs. 10I, 11H, I) ............................ 43 Palpus with 4 segments (as in Fig. 9E); claws on hind legs equal ............................. .................................................................. undescribed genus near Parabezzia MALLOCH Palpus with 5 segments; claws on hind legs a single talon or unequal (Figs. 14F, 15A-C) ................................................................................................................................ 42 Wing with pigmentation spots immediately basal to or over r-m and over apex of radial cell (Fig. 11E); hind tarsomere 1 with only one row of palisade setae ......... ............................................................................................................ Alluaudomyia KIEFFER Wing without pigmentation spots; hind tarsomere 1 with at least 1 1/2 rows of palisade setae ....................................................................... Stilobezzia KIEFFER (in part) Tenth segment of female elongated and bent anteroventrally (Fig. 22F) ............... ...........................................................................................Macrurohelea INGRAM & MACFIE Tenth segment of female not elongated, directed posteriorly ............................... 44 Wing with patches of pigmentation, either 2-3 dark bands (Fig. 11H) or at least with patches of pigmentation at r-m and just posterior to the apex of the second radial cell ............................................................................................................................ 45 Wing without patches of discrete pigmentation ....................................................... 47 Fore- and midlegs each with one claw (each with a smaller basal tooth) ................ ................................................................................................ Stilobezzia KIEFFER (in part) Fore- and midlegs each with 2 equal claws ................................................................ 46 Hind claw with basal tooth (Fig. 14H) ............................................. Allohelea KIEFFER Hind claw without basal tooth (Fig. 14I) .................. Downeshelea WIRTH & GROGAN Fore- and midlegs each with 1 claw (some with a smaller basal tooth) ............... 48

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Fore- and midlegs each with paired, equal, or unequal-sized claws (as in Fig. 14F) .............................................................................................................................................. 49 First radial cell small, with R3 thickened (Fig. 10I); second radial cell somewhat broad; medial vein reduced to faint trace on basal portion; single claw on each leg short, stout and sharp (Fig. 15F).. ................................ Cacaohelea WIRTH & GROGAN First radial cell small to well-developed, with R3 at most thickened at apex (Fig. 11I); second radial cell narrow; medial veins clearly visible but with base of M2 sometimes absent; claw on at least hind leg elongate (Figs. 15A-C) ........................ ................................................................................................ Stilobezzia KIEFFER (in part) Radial cells separated by a short longitudinal vein (R2); wing with a few macrotrichia restricted to extreme periphery at tip; antennal flagellomeres without sensilla coeloconica ..................................... Isthmohelea INGRAM & MACFIE Radial cells separated by a short transverse vein (R2); wing with or without macrotrichia on apical 1/4; some antennal flagellomeres with sensilla coeloconica (either only on flagellomere 1 or only on flagellomeres 8 and 12) ........................................................................................................................................ 50 Wing without macrotrichia; mandibles reduced, vestigial; eyes bare or pubescent; claws small, equal in all legs ..................... Notiohelea GROGAN & WIRTH Wing with macrotrichia on apical 1/4; mandibles toothed; eyes pubescent; only fore- and midlegs with small, equal claws, or claws unequal on all legs (as in Fig. 14F) ............................................................................................................................. 51 Radial cells subequal, small; R3 thickened; midleg claws unequal and somewhat similar to those of hind leg; sensilla coeloconica only on flagellomere 1 ........ ........................................................................................ Borkenthelea SPINELLI & GROGAN Second radial cell about twice as long as first radial cell; R3 not thickened; midleg with small, equal claws, much shorter than elongate single claw (with basal tooth) of hind leg; sensilla coeloconica only on flagellomeres 8 and 12 ..................... ............................................................................................. Austrohelea WIRTH & GROGAN Tarsomere 5 of foreleg swollen (Figs. 15G, 17A, C); HETEROMYIINI ........... 53 Tarsomere 5 of foreleg not swollen ............................................................................. 56 M barely sessile, forking at r-m (Fig. 12G) .................. Physohelea GROGAN & WIRTH M broadly sessile, forking before r-m (Fig. 12H) ...................................................... 54 Fore femur swollen, with numerous thick ventral spines (Fig. 17C) ........................ ........................................................................................................................ Heteromyia SAY Fore femur slender, with at most a few slender spines ........................................... 55 Fourth tarsomere of all legs cylindrical or slightly cordate and lacking strong spines; female hind leg strikingly elongate, with tarsomere 2 at least 3/4 the length of tarsomere 1 (Fig. 15H); hind claw very long, either single or with a small basal tooth, much longer than claws of fore and midlegs (Fig. 15H) ................................ .............................................................................................................. Pellucidomyia MACFIE Fourth tarsomere of foreleg cordate of mid and hind leg bifid and spinose (Fig. 17B); female hind leg not much longer than other legs, with tarsomere 2 less than twice the length of tarsomere 1; both hind claws about same length as those on fore- and midleg ................................................................................... Clinohelea KIEFFER

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H H

B. Other subfamilies

Palisade setae

A. Most Ceratopogoninae H

H

Empodium

C. Atrichopogon

D. Forcipomyia F

M

H

H

E. Dasyhelea H

G. Echinohelea

F. Alluaudomyia H

H. Allohelea

H

I. Downeshelea

Figure 14. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Ceratopogon MEIGEN first tarsomere. B: Forcipomyia sp. first tarsomere. C: Atrichopogon levis (COQUILLETT) (Nearctic Region). D: Forcipomyia fusicornis (COQUILLETT) (Nearctic and Neotropical Regions). E: Dasyhelea pseudoincisurata WAUGH & WIRTH (Nearctic Region). F: Alluaudomyia bella (COQUILLETT) (Nearctic Region). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Allohelea neotropica WIRTH (Neotropical Region). I: Downeshelea sp. (C-E, G from MCALPINE et al., 1981; F, H-I from WIRTH & GROGAN, 1988).

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

F

H

B. Stilobezzia

A. Stilobezzia

F

M

C. Stilobezzia M F

M

F

H

119

H

H

M F

D. Leptohelea H F

M

E. Parastilobezzia H

H

F

M

H. Pellucidomyia F. Cacaohelea

G. Clinohelea

Figure 15. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Stilobezzia antennalis (COQUILLETT) fifth tarsomere (Nearctic and Neotropical Regions). B: Stilobezzia antennalis (COQUILLETT) fifth tarsomere (Nearctic and Neotropical Regions). C: Stilobezzia lutea (MALLOCH) fifth tarsomeres (Nearctic Region). D: Leptohelea micronyx WIRTH & BLANTON fifth tarsomeres (Neotropical Region). E: Parastilobezzia leei WIRTH & BLANTON fifth tarsomeres (Neotropical Region). F: Cacaohelea youngi WIRTH & GROGAN (Neotropical Region). G: Clinohelea sp. H: Pellucidomyia ugandae MACFIE tarsi (Afrotropical Region). (A-B from MCALPINE et al., 1981; C-F from WIRTH & GROGAN, 1988; G from GROGAN & WIRTH, 1975; H from DE MEILLON & WIRTH, 1991).

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F

M

H

H F

M

A. Neobezzia H

C. Groganhelea H

B. Neobezzia H

M F

E. Phaenobezzia

D. Groganhelea Figure 16. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Neobezzia sp. fifth tarsomeres. B: Neobezzia sp. tarsi. C: Groganhelea rondoniensis SPINELLI & DIPPOLITO tarsi (Neotropical Region). D: Groganhelea rondoniensis SPINELLI & DIPPOLITO tarsi (Neotropical Region). E: Phaenobezzia sp. hind leg. (A-B from SPINELLI & WIRTH, 1993; C-D from SPINELLI et al., 1995; E from WIRTH & GROGAN, 1982).

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H

F

A. Clinohelea

B. Clinohelea

F

M

C. Heteromyia

D. Heteromyia H

E. Heteromyia

F

H

F. Stenoxenus

G. Johannsenomyia

M

H. Johannsenomyia

H

I. Johannsenomyia

Figure 17. Female fifth tarsomeres (F = foreleg, M = midleg, H = hind leg). A: Clinohelea bimaculata (LOEW) (Nearctic Region). B: Clinohelea bimaculata (LOEW) (Nearctic Region). C: Heteromyia fasciata SAY (Nearctic Region). D: Heteromyia fasciata SAY (Nearctic Region). E: Heteromyia fasciata SAY (Nearctic Region). F: Stenoxenus coomani SÉGUY (Oriental Region). G: Johannsenomyia argentata (LOEW) (Nearctic Region). H: Johannsenomyia argentata (LOEW) (Nearctic Region). I: Johannsenomyia argentata (LOEW) (Nearctic Region) (all from MCALPINE et al., 1981).

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56(52) Male genitalia large, with gonocoxites fused dorsally and ventrally (Fig. 21B); female genitalia with sternite 8 large and plate like (Fig. 23A) .................................. ................................................................................................................. Echinohelea MACFIE – Male genitalia small or large, if large then gonocoxites not fused; female genitalia with sternite 8 moderately developed .......................................................................... 57 57(56) Male (male of Groganhelea, Sphaerohelea are unknown) .............................................. 58 – Female ................................................................................................................................. 72 58(57) Gonostylus much reduced in size (Figs. 21D, 22B) or fused with gonocoxite (Fig. 22D) .................................................................................................................................... 59 – Gonostylus large, clearly present (Figs. 21F, 22A, C) (male Lanehelea are not known but likely key to at least this point) ................................................................. 61 59(58) Wing with 2 clearly defined radial cells (Fig. 13I); gonostylus well- defined and with pointed apex (Fig. 21D) .................................Clastrieromyia SPINELLI & GROGAN – Wing with 1 radial cell or with, at most, a faint indication of a division between first and second radial cell (Fig. 13D); gonostylus either small and apically rounded (Fig. 22B) or not clearly defined and fused with gonocoxite (Fig. 22D) .............................................................................................................................................. 60 60(59) Anepisternum with numerous elongate setae; gonocoxite short and gonostylus present but small Fig. 22B) ................................................. Phaenobezzia HAESELBARTH – Anepisternum without setae; gonocoxite and gonostylus fused and elongate (Fig. 22D) .......................................................................................................Nilobezzia KIEFFER 61(58) Genitalia with gonocoxite and tergite 9 elongate (Fig. 22C) .................................. 62 – Genitalia more stout, with gonocoxite and tergite 9 not unusually elongate (Figs. 21F, 22A) ............................................................................................................................ 64 62(61) Wing with 1 radial cell (as in Fig. 13D) ....... Neobezzia WIRTH & RATANAWORABHAN – Wing with 2 radial cells (Fig. 13G) ............................................................................... 63 63(62) Forefemur with ventral thick spines .................................. Austrosphaeromias SPINELLI – Forefemur without ventral thick spines ....................................... Mallochohelea WIRTH (Note. Some Mallochohelea species have males with non-elongate genitalia) 64(61) Tarsomere 5 of at least hind leg with stout, blunt spines (Fig. 17H); apex of tergite 9 with single row of thick, stout setae (Fig. 21C) .............. Johannsenomyia MALLOCH – Tarsomere 5 without thick spines, or if present, these more slender and sharply pointed (Fig. 16A); tergite 9 with scattered setae, never arranged in a single row of thick, stout setae ............................................................................................................... 65 65(64) With 1-2 stout setae on apex of the cercus (Fig. 21E); parameres fused, with fusion restricted to most anteromedial portion (Fig. 21E) .................................. ............................................................................................. Amerohelea GROGAN & WIRTH – With several to many setae on the cercus; parameres separate or with fusion various ............................................................................................................................... 66 66(65) Forefemur with stout spines (Fig. 18A) ...................................................................... 67 – Forefemur without stout spines .................................................................................... 68 67(66) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part) – Wing with 2 radial cells (Fig. 13B) ................................... Palpomyia MEIGEN (in part) 68(66) Palpus with 4 segments ............................................................... Stenoxenus COQUILLETT

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– Palpus with 5 segments ................................................................................................... 69 69(68) Thorax slender and dorsoventrally flattened; paratergite with numerous small setae; 1 radial cell (Fig. 12I) ..................................................... Paryphoconus ENDERLEIN – Thorax stouter and not dorsoventrally flattened; paratergite without setae; 1-2 radial cells .......................................................................................................................... 70 70(69) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part) – Wing with 2 radial cells (Fig. 13B) ............................................................................... 71 71(70) Hind femur swollen (Fig. 18C); abdomen white, contrasting with dark thorax ..... .....................................................................................................................Pachyhelea WIRTH – Hind femur not swollen (Fig. 18A); abdomen light to dark brown, similar to dark thorax ...................................................................................... Palpomyia MEIGEN (in part) 72(57) Wing with M2 strongly curving near base (Fig. 13A); palpus with 4 segments ...... .......................................................................................................... Stenoxenus COQUILLETT – Wing with M2 with not or only slightly curved near base (Figs. 13B-I); palpus with 5 segments ......................................................................................................................... 73 73(72) Tarsomere 5 armed ventrally with stout black blunt spines (Figs. 16A, B, 17G-I) (these spines somewhat sharp pointed in Austrosphaeromias); abdomen without tergal apodemes; sternite 8 with or without (Sphaerohelea only) posterolateral hair tufts, with long, medially overlapping, curved setae (best examined at 100X) (Fig. 23B); ....................................... SPHAEROMIINI..........74 – Tarsomere 5 unarmed, or if with some ventral spines, these are slender and with sharp tips (Fig. 16E); abdomen with (Figs. 23C-E) or without tergal apodemes (not visible in some teneral or young specimens); sternite 8 without hair tufts .............................................................................................................................................. 81 74(73) Femora without spines; claws equal on foreleg, unequal on mid- and hind legs (Figs. 17G-I) ............................................................................... Johannsenomyia MALLOCH – Femora with or without spines; claws equal on all legs or with only forelegs unequal ............................................................................................................................... 75 75(74) Claws gently curved distally, with slender basal tooth on internal side ............... 76 – Claws straight or flattened distally, with blunt or slender external basal tooth (Figs. 16A, B) ............................................................................................................................... 78 76(75) Forefemur swollen, with numerous thick ventral spines ............................................. .................................................................................................... Austrosphaeromias SPINELLI – Fore femur slender, unarmed ........................................................................................ 77 77(76) Costa extending to wing tip; tarsomere 5 with 2-3 pairs of ventral spines ............. ............................................................................. Sphaerohelea SPINELLI & FELIPPE-BAUER – Costa not extending beyond 0.85 of wing length; tarsomere 5 with 4-5 pairs of ventral spines ...................................................................... Lanehelea WIRTH & BLANTON 78(75) Wing with 1 radial cell; costa long, extending nearly to wing tip with costal ratio over 0.87; femora without ventral spines .................................................................... 79 – Wing with 1-2 radial cells; costa short, extending to less than 0.84 of wing length; femora with or without ventral spines ......................................................................... 80

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F

M

H

A. Palpomyia

H

F

B. Bezzia

C. Pachyhelea

Figure 18. Female legs (F = foreleg, M = midleg, H = hind leg). A: Palpomyia plebeja (LOEW) (Nearctic Region). B: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). C: Pachyhelea pachymera (WILLISTON) (Nearctic and Neotropical Regions) (all from MCALPINE et al., 1981).

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Parameres

A. Leptoconops

B. Forcipomyia

Parameres

Parameres

C. Dasyhelea

D. Culicoides

Parameres

Parameres

Aedeagus

E. Stenoxenus

F. Bahiahelea

Figure 19. Male genitalia, ventral view. A: Leptoconops torrens (TOWNSEND) (Nearctic Region). B: Forcipomyia acidicola (TOKUNAGA) (Holarctic Region). C: Dasyhelea pseudoincisurata WAUGH & WIRTH (Nearctic Region). D: Culicoides haematopotus MALLOCH (Nearctic and Neotropical Regions). E: Stenoxenus coomani SÉGUY (Oriental Region). F: Bahiahelea brasiliensis WIRTH (Neotropical Region) (AE from Manual of Nearctic Diptera; F from BORKENT, unpubl.).

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A. Leptohelea

B. Cacaohelea

C. Isthmohelea

D. Stilobezzia

Aedeagus

Parameres Gonocoxite

Gonostylus Cercus

E. Stilobezzia

F. Downeshelea

Figure 20. Male genitalia, ventral view. A: Leptohelea micronyx WIRTH & BLANTON (Neotropical Region). B: Cacaohelea sp. C: Isthmohelea disjuncta INGRAM & MACFIE (Neotropical region). D: Stilobezzia sp. E: Stilobezzia sp. (Neotropical Region). F: Downeshelea sp. (A, B, D-F courtesy of INBio, Costa Rica; C from INGRAM & MACFIE, 1931).

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Aedeagus

A. Monohelea

C. Johannsenomyia

B. Echinohelea

Gonostylus

D. Clastrieromyia

Paramer

E. Amerohelea

F. Paryphoconus

Figure 21. Male genitalia, ventral view. A: Monohelea sp. B: Echinohelea sp. C: Johannsenomyia argentata (LOEW) (Nearctic Region). D: Clastrieromyia dycei SPINELLI & GROGAN (Neotropical Region). E: Amerohelea frontispina (DOW & TURNER) (Nearctic and Neotropical Regions). F: Paryphoconus angustipennis ENDERLEIN (Neotropical Region). (A-E courtesy of INBio, Costa Rica; F from MCALPINE et al., 1981).

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79(78) External basal tooth of tarsal claws pointed (Fig. 16A); tarsus of hind leg not elongate (Fig. 16B); wing with anal lobe poorly developed ........................................ ..............................................................................Neobezzia WIRTH & RATANAWORABHAN – External basal tooth of tarsal claws blunt (Fig. 16C); tarsus of hind leg elongate (Fig. 16D); wing with anal lobe well developed (Fig. 13F) ......................................... ...................................................................................... Groganhelea SPINELLI & DIPPOLITO 80(78) Wing with 2 radial cells (Fig. 13G); wing grayish with veins darker brown, anterior veins prominent; forefemur without ventral thick spines ........ Mallochohelea WIRTH – Wing with 1 radial cell; wing whitish, anterior veins pale; forefemur with ventral thick spines ...........................................................................................Nilobezzia KIEFFER 81(73) Thorax slender and dorsoventrally flattened; paratergite with numerous small setae; 1 radial cell (Fig. 12I) ..................................................... Paryphoconus ENDERLEIN – Thorax stouter and not dorsoventrally flattened; paratergite without setae; wing with 1-2 radial cells .................................................................. PALPOMYIINI..........82 82(81) Abdomen with 1 pair of tergal apodemes arising near lateral margins of sternite 7; 1 spermatheca present, with no evidence of a reduced second one ........................ ............................................................................................. Amerohelea GROGAN & WIRTH – Abdomen with 1 or more pairs of tergal apodemes (Figs. 23C-E), if only 1 pair present, these arise from more medially on posterior margin of sternite 7; 1-2 spermathecae but if apparently only one, a small reduced second one is present .............................................................................................................................................. 83 83(82) Wing with 2 radial cells (Fig. 13B) ............................................................................... 84 – Wing with 1 radial cell (Fig. 13C) ................................................................................. 86 84(83) Forefemur without stout spines; hind femur greatly swollen (Fig. 18C); abdomen white, contrasting with dark thorax ....................................................Pachyhelea WIRTH – Forefemur with or without stout spines (Fig. 18A); hind femur not greatly swollen (Fig. 18A); abdomen light to dark brown, similar to dark thorax ......................... 85 85(84) Flagellum short, only 1.2-1.4 times longer than the breadth of the head; wing membrane unpigmented; anal lobe of wing well-developed (Fig. 13I); abdomen with single pair of apodemes arising from the anterior margin of tergite 7 and extending to about the anterior margin of tergite 6 (Fig. 23C) ................................. ......................................................................................Clastrieromyia SPINELLI & GROGAN – Flagellum elongate, at least 2.5 times longer than the breadth of the head; wing membrane pigmented; anal lobe of wing poorly developed (Fig. 13B); abdomen with 2 or more pairs of elongate apodemes arising from the anterior margin of at least tergites 6-7 and extending much further anteriorly into abdominal cavity (Fig. 23E) ............................................................................................... Palpomyia MEIGEN 86(83) Costa moderately short with costal ratio 0.67-0.75 (Fig. 13C); forefemur with (Fig. 18B) or without stout spines; tarsomere 5 with or without stout ventral setae ..... ....................................................................................................... Bezzia KIEFFER (in part) – Costa more elongate with costal ratio more than 0.80 (Fig. 13D); forefemur without stout spines; tarsomere 5 with slender ventral setae with sharp, bent tips (Fig. 16E) ................................................................................ Phaenobezzia HAESELBARTH

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

KEY

TO THE GENERA OF EGGS OF

CERATOPOGONIDAE IN

THE

129

NEOTROPICAL REGION

There are only a few species of Ceratopogonidae for which the eggs have been described (Table 1). The following key is therefore very tentative. There are no keys to the eggs of Diptera which would allow those of Ceratopogonidae to be distinguished from those of other families. Nevertheless, some Ceratopogonidae have eggs which are either different from those of all other Diptera (Dasyhelea, Stenoxenus and Paryphoconus) or are laid in a distinctive matrix (Alluaudomyia and most Sphaeromiini). 1 – 2(1) – 3(2) – 4(3)

– 5(4) –

KEY

Egg C-shaped (Fig. 24A) .................................................................................... Dasyhelea Egg with, at most, a slight curvature (Figs. 24B-F) .................................................... 2 Egg with distinctive subapical frill and terminal nipple-like constriction at anterior end (Fig. 24F) ........................................................................ Stenoxenus and Paryphoconus Egg without any well developed ornamentation at either end; small spicules may be present (Figs. 24A-E) ................................................................................................... 3 Egg oval (Fig. 24B) ............................ Leptoconops (in part), Forcipomyiinae (in part) Egg elongate, slender (Figs. 24C-E) ............................................................................... 4 Eggs arranged in an elongate ribbon, with eggs lying at an angle to the length of the ribbon (Fig. 24E) ........................................................................................................... .................. most Sphaeromiini (known for Johannsenomyia, Mallochohelea, Macropeza) Eggs laid separately or in group; if in a ribbon, eggs are laid side by side (Figs. 24C-D) .................................................................................................................................. 5 Eggs laid in ribbon (Fig. 24D) ..................................... Alluaudomyia (at least in part) Eggs laid singly or in a loose group (but not attached to one another longitudinally) (Fig. 24C) .......................................................... other Ceratopogonidae TO THE GENERA OF LARVAE OF

CERATOPOGONIDAE

IN THE

NEOTROPICAL REGION

The immatures of Ceratopogonidae are among the most poorly known of all nematocerous families (Table 2). Here we provide a tentative key to the larvae of some common genera. A world key to the genera of larvae and pupae is being prepared by the senior author. It is important to realize that only 11 of 51 known genera of Neotropical Ceratopogonidae have been described as larvae in this region (Table 2), although an additional 11 genera are known as larvae from elsewhere. 1

–

2(1)

Proleg present on prothorax (Figs. 25B, C); head capsule hypognathous to prognathous (Figs. 25B, C); with strong setae on head capsule and body (Figs. 25B, C) (Forcipomyiinae) ........................................................................................................... 2 Without proleg on thorax (Figs. 25A, D-F); head capsule somewhat hypognathous or prognathous (Figs. 25A, D-F); without strong setae on head capsule and body anterior to anal segment (Figs. 25A, D-F) .................................................................... 3 Body somewhat flattened dorsoventrally, with lateral tubercles (Fig. 25B) ............. ............................................................................................................................. Atrichopogon

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Parameres

Gonostylus

Parameres

B. Phaenobezzia

A. Bezzia

Parameres

C. Mallochohelea

D. Nilobezzia

Tenth segment

Cercus

E. Leptoconops

F. Macrurohelea

Figure 22. Male genitalia, ventral view. A: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). B: Phaenobezzia pistiae (INGRAM & MACFIE) (Afrotropical Region). C: Mallochohelea albibasis (MALLOCH) (Nearctic Region). D: Nilobezzia minor (WIRTH) (Nearctic Region). Female genitalia, ventral view. E: Leptoconops bequaerti (KIEFFER) (Nearctic and Neotropical Regions). F: Macrurohelea sp. (Neotropical Region). (A-D from MCALPINE et al., 1981; E courtesy of INBio, Costa Rica; F from SPINELLI & WIRTH, 1993).

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Sternite 8

Sternite 9 Sternite 10 Cercus

Sternite 8

A. Echinohelea

B. Probezzia

Tergal apodeme

Tergal apodeme

C. Clastrieromyia

D. Paryphoconus

E. Palpomyia

Figure 23. Female genitalia, ventral view. A: Echinohelea sp.(Neotropical Region). B: Probezzia concinna (MEIGEN) (Holarctic Region). C: Clastrieromyia dycei SPINELLI & GROGAN (Neotropical Region). D: Paryphoconus angustipennis ENDERLEIN (Neotropical Region). E: Palpomyia lineata (MEIGEN) (Holarctic Region). (A courtesy of INBio, Costa Rica; B, D-E from MCALPINE et al., 1981).

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A. Dasyhelea

B. Forcipomyia

C. Culicoides

D. Alluaudomyia

E. Probezzia

F. Paryphoconus

Figure 24. Eggs. A: Dasyhelea flavifrons (GUÉRIN) (Palearctic Region). B: Forcipomyia mcateei WIRTH (Nearctic Region). C: Culicoides pulicaris (LINNAEUS) (Palearctic Region). D: Alluaudomyia needhami THOMSEN (Nearctic Region). E: Probezzia concinna (GUÉRIN) (Palearctic Region). F: Paryphoconus angustipennis ENDERLEIN (Neotropical Region). (A from SZADZIEWSKI et al., 1997; B from CHAN & LEROUX, 1965 (as F. saundersi); C from GUTSEVICH & GLUKHOVA, 1970; D from THOMSEN, 1937; E from WESENBERG-LUND, 1914; F from MAYER, 1952).

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A. Leptoconops

Prothoracic proleg

B. Atrichopogon

Prothoracic proleg

Posterior proleg

Posterior proleg

C. Forcipomyia

D. Dasyhelea Posterior proleg

E. Culicoides

F. Bezzia Figure 25. Larval habitus. A: Leptoconops gallicus CLASTRIER (Palearctic Region), lateral view. B: Atrichopogon fusculus (COQUILLETT) (Holarctic and Neotropical Regions), lateral view. C: Forcipomyia sp., lateral view. D: Dasyhelea sp., lateral view. E: Culicoides sp., dorsal view. F: Bezzia sp., dorsal view (A from CLASTRIER, 1973; B-F from MCALPINE et al., 1981, B as A. polydactylus).

134

– 3(1)

– 4(3)

–

5(4) – 6(5) –

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Body cylindrical in cross-section, without lateral tubercles (Fig. 25C) ........ Forcipomyia Head capsule directed anteroventrally; end of abdomen with series of hooks, sometimes arranged as two prolegs, (often retracted and difficult to see in alcohol material), never with elongate setae (Fig. 25D) ............................................. Dasyhelea Head capsule prognathous; end of abdomen simple, without hooks, with or without elongate setae (Figs. 25A, E, F) ....................................................................... 4 Head capsule somewhat poorly developed pigmented, with well-developed, posteriorly directed apodemes extending into prothorax (Figs. 26A, B); abdomen with segments secondarily divided (Fig. 25A) ............................................ Leptoconops Head capsule well-developed, lacking apodemes posterior to margin of head capsule (Figs. 26C-F); abdomen with segments normal, not secondarily divided (Figs. 25E, F) (Ceratopogoninae) ................................................................................... 5 Head capsule with all setae simple (Figs. 26C, D) ................................. Culicoides Head capsule with one or more of setae s, u and posterior o plumose (Figs. 26E, F; seta u not shown, it is generally lateral on head capsule) ........................... 6 Posteroventral margin of head capsule with posteriorly projecting process (Fig. 26F) ................................................................................................. Sphaeromiini (in part) Posteroventral margin of head capsule a straight or somewhat curved line, without posteriorly projecting process (Fig. 26E) ......................................................... ................................................... Sphaeromiini (in part), Palpomyiini, Ceratopogonini

CLAVE PARA LOS GÉNEROS DE ADULTOS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL Existen hasta la fecha 1095 especies de Ceratopogonidae descriptas de la región Neotropical (Tabla 5), aunque muchas más aguardan para a ser descriptas, particularmente en los géneros Forcipomyia, Atrichopogon, Dasyhelea, Culicoides y Stilobezzia. Especies de esta familia pueden ser halladas en todas las altitudes y virtualmente en todo hábitat donde exista por lo menos algo de humedad. La siguiente clave incluye a los géneros conocidos de la región Neotropical y, por primera vez, a los machos de todos los géneros (no son conocidos los machos de Groganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, y Sphaerohelea) Esta clave está destinada a identificar tanto material conservado en alcohol como material montado en lámina. Hemos señalado en negrita los caracteres que se observan mejor mediante el uso de microscopio (por ej., en láminas), aunque con práctica y paciencia muchos de estos caracteres también pueden ser observados en material conservado en alcohol. En algunos grupos, algunas nervaduras de las alas puedes ser dificiles de observar; con ejemplares conservados en alcohol, es posible reconocer débiles trazos de las nervaduras al examinarlos de perfil a la vez que se inclina levemente el ala, mientras que con material montado en lámina, la mejor resolución es la que brinda el microscopio de contraste de fase. Cuando las dos uñas al final de la pata son del mismo tamaño, se las refiere como iguales; cuando son de diferente tamaño, como desiguales. La relación costal es el largo de la costa desde la base del ala (medida desde el arculus), dividido por el largo total del ala. La terminología adoptada para las nervaduras y celdas de las alas sigue aquella del “Manual of Nearctic Diptera” (MCALPINE et al., 1981), con las modificaciones propuestas

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Apodemes

A. Leptoconops

B. Leptoconops

C. Culicoides

D. Culicoides

Seta s

Seta o

Projecting process

E. Nilobezzia

F. Nilobezzia

Figure 26. Larval head capsules. A: Leptoconops gallicus CLASTRIER (Palearctic Region), dorsal view. B: Leptoconops gallicus CLASTRIER (Palearctic Region), ventral view. C: Culicoides reconditus CAMPBELL & PELHAM-CLINTON (Palearctic Region), dorsal view. D: Culicoides reconditus CAMPBELL & PELHAMCLINTON (Palearctic Region), ventral view. E: Nilobezzia formosa (LOEW) (Palearctic Region), dorsal view. F: Nilobezzia formosa (LOEW) (Palearctic Region), ventral view. (A-B from CLASTRIER, 1973; CF from GLUKHOVA, 1979).

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por SZADZIEWSKI (1996), y rotuladas conforme a las Figs. 10C, D y 11I. Al igual que otros dipterólogos, las nervaduras son escritas aquí en letras mayúsculas y las celdas en letras minúsculas. 1

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Ojos ampliamente separados, sin sutura frontal (Fig. 6A); palpo con 4 segmentos (sólo uno más allá del tercero, el cual es alargado o hinchado) y con intervalo membranoso entre los segmentos 3 y 4+5 (Fig. 9A); ala sin nervadura transversal r-m (Fig. 10A); hembra con las celdas radiales fusionadas en un estigma (Fig. 10A); sin hileras de setas empalizadas en el primer tarsómero de la pata posterior (Fig. 14B); gonostilo del macho con una púa alveolada apical (Fig. 19A); cerco de la hembra muy alargado (Fig. 22E) .................................. Leptoconops SKUSE Ojos estrecha o anchamente separados en el dorso, con sutura frontal (Figs. 6BG); palpo con 2-5 segmentos, pero nunca con intervalo membranoso entre los dos últimos segmentos (Figs. 9B-E); ala con nervadura transversal r-m (Figs. 10B-J); celdas radiales distintivas o, si están fusionadas o ausentes (Figs. 11A, F), la nervadura transversal r-m está claramente presente; con (Fig. 14A) o sin (Fig. 14B) hileras de setas empalizadas en el primer tarsómero de la pata posterior; gonostilo del macho sin púa alveolada apical (Figs. 19B-F, 20A-F) aunque el ápice puede ser puntiagudo (Fig. 19E); cerco de la hembra corto (Figs. 22F, 23A-E) ......................................................................................................................... 2 Ala con (Figs. 10B, E, G) o sin (Figs. 10D, J, 11A-C) numerosas macrotriquias en la membrana; primer tarsómero posterior con setas dispersas o, a lo sumo, con una hilera de pelos robustos (pero éstos no forman un hilera apretada, especialmente en la base del tarsómero) (Fig. 14B); hembra con dos uñas pequeñas, iguales, en cada pata (Figs. 14C-E); con (Figs. 14C, D) o sin (Figs. 14E, F) empodio bien desarrollado ......................................................................................... 3 Ala desnuda o con macrotriquias dispersas en la mitad distal del ala (unas pocas especies tienen algunas macrotriquias en la base del ala, pero éstas se hallan dispersas) (Figs. 11E, I); primer tarsómero posterior con hilera de setas con al menos más setas basales cuyas bases contactan, algunos grupos con una fuerte espina basal adicional (Figs. 14A, G-I); hembra con dos uñas pequeñas, iguales, en cada pata, o con 1-2 uñas en una o más patas, las cuales en algunas son largas (Figs. 14F-I, 15A-C, G, H, 16A-E); sin empodio bien desarrollado ....................... 7 Costa sobrepasando la mitad del largo del ala (Figs. 10C, D); ambas celdas radiales bien desarrolladas, con la segunda celda radial larga y abierta moderadamente (Figs. 10C, D); paratergito bien desarrollado, con una seta fuerte y 0-5 setas finas más cortas (Fig. 9F) ........................................................................ Atrichopogon KIEFFER Costa corta o larga; si es larga, entonces la seguna celda radial es ancha (Figs. 10F, G) o angosta (Figs. 10B, E); paratergito angosto (como en Fig. 9G) o bien desarrollado, sin setas o, si hay setas presentes, son al menos 10 e iguales en tamaño y longitud .............................................................................................................. 4 Flagelómero apical con pezón terminal, el cual es estrechado basalmente (Figs. 7A, B); uñas fuertemente curvas (Figs. 14C, D); empodio vestigial o bien desarrollado (Figs. 14C, D) ........................................................... Forcipomyia MEIGEN

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Flagelómero apical sin pezón terminal (aunque puede ser fuertemente adelgazado) (Fig. 7C); uñas sólo ligeramente curvas (Fig. 14E); empodio pobremente desarrollado .............................................................................................. 5 Ojos anchamente colindantes en la línea media; flagelómeros antenales sin sensilas celocónicas; macho con setas cortas en el primer flagelómero, menores a la mitad del largo de aquellas de los flagelómeros subsiguientes (7C); flagelómeros del macho esculturados (Fig. 7C); sin espolón en la pata anterior; membrana del ala sin diseño distintivo de pigmentación oscura (Fig. 10E) ......................................................................................................... Dasyhelea KIEFFER Ojos contiguos a anchamente separados en la línea media (Figs. 6B, C); algunos flagelómeros antenales con sensilas celocónicas; macho con setas del primer flagelómero más o menos del mismo largo que aquellas de los flagelómeros subsiguientes; flagelómeros del macho no esculturados; con espolón tibial en la pata anterior; membrana del ala con o sin diseños distintivos de pigmentación oscura (Figs. 10F-H); CULICOIDINI ....................... 6 Ala con 1-2 celdas radiales; costa sobrepasando la mitad del largo del ala (Figs. 10F, G); hembras con o sin mandíbulas dentadas; membrana alar con o sin diseños distintivos de pigmentación oscura y/o clara (Figs. 10F, G) ....................... ............................................................................................................... Culicoides LATREILLE Ala con ambas celdas radiales obliteradas (Fig. 10H); costa no alcanzándo más allá de 0,43 del largo del ala; hembras con mandíbulas reducidas, vestigiales; membran alar sin diseños distintivos de pigmentación oscura y/o clara (Fig. 10H) .............................................................................................................. Paradasyhelea MACFIE M bifurcándose más allá de r-m (v.g. bifurcación mediana peciolada, en algunos de base muy corta) (Fig. 11I), en algunos la base de M2 no está presente (Figs. 10I, J, 11A) ....................................................................................................................................... 8 M bifurcándose al nivel o anterior de r-m (v.g. bifurcación mediana sésil, no peciolada) (Figs. 11G, 12F-J, 13A-I); base de M2 siempre presente ...................... 52 Ala con celdas radiales reducidas (Figs. 11A, B, D, F); relación costal menor que 0,65 (Figs. 11A, B, D, F); uñas de la hembra más o menos iguales (el macho de Rhynchohelea es desconocido, pero probablemente con ojos anchamente separados, flagelómeros antenales 10 y 11 fusionados, y con una seta katepisternal) ............ 9 Ala con al menos una celda radial bien desarrollada (Figs. 10J, 11E, I); relación costal mayor que 0,50; uñas de la hembra iguales o desiguales, algunas con sólo una uña en una o más de las patas ............................................................................... 15 Macho con los flagelómeros antenales 10 y 11 fusionados o con menos de 13 flagelómeros; ala con o sin celdas radiales; katepisterno con 1 o más setas laterales (pueden ser observadas en ejemplares preservados en alcohol, tomándolos verticalmente y observándolos de perfil a lo largo del plano de la pleura torácica) (Fig. 9G) ................................................................................................ 10 Antena del macho con flagelómeros 10 y 11 separados y con 13 flagelómeros presentes; ala sin celda radial (celdas completamente fusionadas) (Figs. 11A, B, D); katepisterno sin setas (Fig. 9H) ................................................ 13

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Antena de la hembra con 12 flagelómeros (Fig. 7D); labio de la hembra robusto, truncado, con estructuras apicales en forma de gancho (Fig. 6D) ............................ ................................................. Rhynchohelea WIRTH & BLANTON (macho desconocido) Antena de la hembra con 13 flagelómeros; proboscis de la hembra normal (Figs. 6B, C, E-G) ........................................................................................................................ 11 Flagelo del macho con 7-8 flagelómeros (Fig. 8A); palpo con 3 segmentos (Fig. 9C) ...................................................................................... Nannohelea GROGAN & WIRTH Flagelo del macho con al menos 9 flagelómeros (casi siempre con 13, pero con 10 y 11 fusionados); palpo con 5 segmentos ................................................................... 12 Anepisterno sin setas; antena del macho con flagelómeros 2-6 fusionados (611 también fusionados) .............................................. Brachypogon KIEFFER (en parte) Anepisterno con al menos 1 seta en el margen posterior (Fig. 9G); antena del macho con flagelómeros 2-5 articulados, no fusionados (flagelómeros 7-11 fusionados, a veces incluso el flagelómero 6) ................................................................ ...................................................................... Ceratoculicoides WIRTH & RATANAWORABHAN Palpo con 5 segmentos; largo del ala 0,7-0,8 mm; macho con parámeros bien desarrollados .............................................................................. Schizonyxhelea CLASTRIER Palpo con 2 segmentos (Fig. 9B); largo del ala < 0,6 mm; macho sin parámeros .............................................................................................................................................. 14 Tibia anterior con espolón apical; macrotriquias confinadas a los márgenes distales del ala (en algunos, también unas pocas en el ápice de M1 y M2); relación costal > 0,40 (Fig. 11B); antena del macho con 6 flagelómeros separados (Fig. 8C); macho con tergitos abdominales 1-3 claros y 4-8 pardos ........................................... ............................................................................................... Baeohelea WIRTH & BLANTON Tibia anterior sin espolón apical; macrotriquias largas y abundantes sobre todo el ala; relación costal ratio < 0,36; antena del macho con 12 flagelómeros, pero con 2-7, 8-9, y 10-11 fusionados (Fig. 8B); macho con abdomen uniformemente pardo ........................................................................ Baeodasymyia CLASTRIER & RACCURT Ala con diseño característico de una pigmentación oscura moteada, más o menos dispersa sobre la membrana (Fig. 12C); tórax y al menos fémur y tibia posteriores con diseño de pigmentación; edeago del macho con lazo posteromediano característico (Fig. 21A) .................................................................... Monohelea KIEFFER1 Ala sin pigmentación o, si es pigmentada, pigmentación restricta al sector radial, o con puntos, rayas o bandas discretas (Figs. 11E, H), o si el patrón de pigmentación es substancial (Fig. 11H) entonces tórax y todos los fémures y tibias uniformemente pardos oscuros; otros taxa con tórax y patas pigmentados de manera variable; edeago, si está presente, sin rizo posteromediano……16 Costa extendiéndose hasta cerca del extremo del ala; costa del ala de la hembra extendiéndose más allá del ápice de la segunda celda radial (Figs. 12B, D, E) (extensión también presente en machos de Bahiahelea y Leptohelea) ...................... 17

M. fairchildi Lane y Wirth no sale bien en la clave. Los adultos tienen alas bandeadas como en Downeshelea, pero el macho posee un edeago con lazo posteromediano característico.

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Costa no extendiéndose más allá de 0,85 del largo del ala (Figs. 11E, I); costa del ala de la hembra finalizando en el ápice de la segunda celda radial o extendiéndose más allá ligeramente ......................................................................................................... 25 Halterio en forma de saco, sin constricción por debajo del capitelo (Fig. 13J) ..... ...................................................................................... Diaphanobezzia INGRAM & MACFIE Halterio con capitelo y pedicelo bien definidos ........................................................ 18 Macho ................................................................................................................................ .19 Hembra ............................................................................................................................... 21 Ala con costa no extendiéndose más allá del ápice de la segunda celda radial ...... ...................................................................................... Parastilobezzia WIRTH & BLANTON Ala con costa extendiéndose apreciablemente más allá del ápice de la segunda celda radial ......................................................................................................................... 20 Con flagelómeros 10-13 más alargados que los flagelómeros precedentes; penacho antenal bien desarrollado; parámeros fusionados en una esfera voluminosa (Fig. 19F) ............................................................................ Bahiahelea WIRTH Con flagelómeros 2-13 más o menos iguales en longitud; penacho antenal corto; parámeros alargados, delgados ...................................... Leptohelea WIRTH & BLANTON Palpo con tres segmentos, último segmento corto y pequeño (Fig. 9D); uñas pequeñas, iguales, sin diente basal interno (Fig. 15D) .................................................. .............................................................................................. Leptohelea WIRTH & BLANTON Palpo con 4-5 segmentos, último segmento alargado (Fig. 9E); uñas variables ..... .............................................................................................................................................. 22 Clípeo no fusionado a la cabeza (como en la Fig. 6B); ala con la primera celda radial pequeña, segunda alargada (Fig. 12E); con una uña (con diente basal grueso) en cada pata (Fig. 15E) ............................ Parastilobezzia WIRTH & BLANTON Clípeo fusionado lateralmente a la cabeza (Figs. 6F, G); ala con 1 celda radial (Fig. 12B); con 2 uñas en cada pata, a pesar de que las uñas de la pata posterior pueden ser desiguales ..................................................................................................................... 23 Ojos moderadamente separados, sin pubescencia (Fig. 6F); uñas sin diente basal interno ................................................................................................ Parabezzia MALLOCH Ojos anchamente separados, con o sin pubescencia (Fig. 6G); uñas con diente basal interno ...................................................................................................................... 24 Palpo con 4 segmentos; ojos pubescentes; espermatecas sin cuellos diferenciados ............................................................... Fittkauhelea WIRTH & BLANTON Palpo con 5 segmentos; ojos sin pubescencia; espermatecas con cuellos diferenciados delgados ...................................................................... Bahiahelea WIRTH Macho ................................................................................................................................. 26 Hembra ............................................................................................................................... 39 Pata posterior con una uña larga y diente basal pequeño (Fig. 14H) ....................... ................................................................................................................... Allohelea KIEFFER Pata posterior con dos uñas iguales o subiguales ...................................................... 27 Katepisterno con 1 o más setas laterales (pueden ser observados en ejemplares preservados en alcohol tomándolos verticalmente y observándolos de perfil a lo largo del plano de la pleura torácica) (Fig. 9G) ....... Brachypogon KIEFFER (en parte)

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– 31(30)

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34(33) – 35(34). –

36(35) – 37(36) –

Katepisterno sin setas .................................................................................................. 28 Ala con 1 celda radial (Fig. 11E) .................................................................................. 29 Ala con 2 celdas radiales ................................................................................................. 33 Coxa anterior con setas espinosas largas, o setas delgadas afinadas distalmente (Fig. 9H) ............................................................................................. Parabezzia MALLOCH Coxa anterior con sólo setas cortas, simples (a veces perdidas en ejemplares en alcohol) ............................................................................................................................... 30 Ala con manchas pigmentadas, al menos inmediatamente basal a, o sobre r-m y sobre el ápice de la celda radial (Fig. 11E), algunos con pigmentación adicional ............................................................................................................ Alluaudomyia KIEFFER Ala sin manchas pigmentadas ........................................................................................ 31 Palpo con 5 segmentos; tarsómero 1 de la pata posterior con al menos 1 1/2 hileras de setas empalizadas; edeago compuesto por 2 escleritos laterales (por lo que el edeago parece estar fisurado en la línea media) (Fig. 20E) o con edeago ausente pero con una barra transversa ventral al edeago (Fig. 20D) ..............................................................................................Stilobezzia KIEFFER (en parte) Palpo con 4 segmentos (como en Fig. 9E); tarsómero 1 de la pata posterior con sólo 1 hilera de setas empalizadas; edeago continuo en la línea media (como en Fig. 21C) ............................................................................................................................. 32 Ojos sin pubescencia; genitalia con parámeros fusionados, muy cortos ......... .................................................................. undescribed genus near Parabezzia MALLOCH Ojos pubescentes; genitalia sin parámeros ........ Fittkauhelea WIRTH & BLANTON Ala con áreas pigmentadas, incluyendo áreas poco notorias, discontínuas en M2 and CuA2 (Fig. 12F); edeago continuo en la línea media (Fig. 20F) ................... ............................................................................................ Downeshelea WIRTH & GROGAN Ala con o sin diseño de pigmentación, pero si está presente, entonces sin áreas poco notorias, discontínuas en M2 and CuA2 (porción posterior del ala puede ser más oscura) (Figs. 11J, 12A) y con edeago dividido en la línea media (Figs. 20B, C) .............................................................................................................................................. 34 Antena con los 5 flagelómeros apicales más alargados que los 8 flagelómeros basales (antena tipo-hembra), sin penacho ................. Cacaohelea WIRTH & GROGAN Antena con los 3 flagelómeros apicales más alargados que los 10 flagelómeros basales (antena tipo-macho), con o sin penacho ....................................................... 35 Celdas radiales separadas por una nervadura longitudinal corta (R2); ala sin macrotriquias; esternito 9 alargado (Fig. 20C) ........... Isthmohelea INGRAM & MACFIE Las 2 celdas radiales cercanas una de la otra, separadas por una nervadura transversal corta (R2) (Figs. 11J, 12A); ala con o sin macrotriquias; esternito 9 corto o moderadamente alargado (Figs. 20D, E) ...................................................... 36 Flagelómeros 3-10 fusionados (Fig. 8D) ............... Borkenthelea SPINELLI & GROGAN Antena con flagelómeros separados, no fusionados ................................................ 37 Largo combinado de los tarsómeros 2-5 de la pata posterior aproximadamente 1,4-1,7 veces mayor que el tarsómero 1 ..................... Austrohelea WIRTH & GROGAN Largo combinado de los tarsómeros 2-5 de la pata posterior aproximadamente del mismo tamaño que el tarsómero 1 ............................................................................... 38

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38(37) Tarsómero 1 de la pata posterior con al menos 1 1/2 hileras de setas empalizadas; ala con o sin macrotriquias; edeago compuesto por 2 escleritos laterales (por lo que el edeago parece estar fisurado en la línea media) (Fig. 20E) o con edeago ausente pero con una barra transversa ventral al edeago (Fig. 20D); flagelómeros antenales sin sensilas celocónicas .................. Stilobezzia KIEFFER – Tarsómero 1 de la pata posterior con sólo 1 hilera de setas empalizadas (como en Fig. 14A); ala sin macrotríquias; edeago no dividido, más o menos triangular; algunos flagelómeros antenales con sensilas celocónicas ................................. ........................................................................................... Macrurohelea INGRAM & MACFIE 39(25) Katepisterno con 1 o más setas laterales (pueden ser observados en ejemplares preservados en alcohol tomándolos verticalmente y observándolos de perfil a lo largo del plano de la pleura torácica) (Fig. 9G) ......... Brachypogon KIEFFER (en parte) – Katepisterno sin setas .................................................................................................. 40 40(39) Ala con 1 celda radial (Figs. 11C, E) ............................................................................ 41 – Ala con 2 celdas radiales (la primera puede ser pequeña) (Figs. 10I, 11H, I) ........ 43 41(40) Palpo con 4 segmentos (como en Fig. 9E); uñas de la pata posterior iguales ........ ...................................................... género no descripto cercano a Parabezzia MALLOCH – Palpo con 5 segmentos; uñas de la pata posterior desiguales o representadas por un talón (Figs. 14F, 15A-C) ............................................................................................ 42 42(41) Ala con manchas pigmentadas inmediatamente basal a, o sobre r-m y sobre el ápice de la celda radial (Fig. 11E); tarsómero 1 de la pata posterior con sólo 1 hilera de setas empalizadas ........................................................... Alluaudomyia KIEFFER – Ala sin manchas pigmentadas; tarsómero 1 de la pata posterior con al menos 1 1/2 hilera de setas empalizadas .............................................Stilobezzia KIEFFER (en parte) 43(40) Décimo segmento de la hembra alargado y doblado anteroventralmente (Fig. 22F) ...........................................................................................Macrurohelea INGRAM & MACFIE – Décimo segmento de la hembra no alargado, dirigido posteriormente ............... 44 44(43) Alas con áreas pigmentadas, con 2-3 bandas oscuras (Fig. 11H), o al menos con áreas pigmentadas en r-m e inmediatamente posterior al ápice de la segunda celda radial .................................................................................................................................... 45 – Ala sin áreas pigmentadas discretas ............................................................................. 47 45(43) Patas anterior y media, cada una con una uña (cada una con un diente basal más pequeño) ............................................................................Stilobezzia KIEFFER (en parte) – Patas anterior y media, cada una con 2 uñas iguales ................................................ 46 46(45) Uña de la pata posterior con diente basal (Fig. 14H) .................... Allohelea KIEFFER – Uña de la pata posterior sin diente basal (Fig. 14I) ..... Downeshelea WIRTH & GROGAN 47(44) Patas anterior y media, cada una con 1 uña (algunos con diente basal más pequeño) ............................................................................................................................ 48 – Patas anterior y media, cada una con uñas pares, iguales, o desiguales (como en Fig. 14F) ............................................................................................................................. 49 48(47) Primera celda radial pequeña, R3 engrosada (Fig. 10I); segunda celda radial más o menos ancha; nervadura mediana reducida a un trazo poco perceptible en la base; una uña corta, robusta, puntiaguda en cada pata (Fig. 15F) ....................................... .............................................................................................. Cacaohelea WIRTH & GROGAN

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Primera celda radial pequeña a bien desarrollada, R3 a lo sumo engrosada en el ápice (Fig. 11I); segunda celda radial angosta; nervadura mediana claramente visible, pero algunas veces la base de M2 está ausente; uña, de al menos la pata posterior, alargada (Figs. 15A-C) ..................................Stilobezzia KIEFFER (en parte) Celdas radiales separadas por una nervadura longitudinal corta (R2); ala con macrotriquias escasas, restrictas al ápice extremo; flagelómeros antenales sin sensilas celocónicas ...................................................... Isthmohelea INGRAM & MACFIE Celdas radiales separadas por una nervadura transversal corta (R2); ala con o sin macrotriquias en el 1/4 apical; algunos flagelómeros antenales con sensilas celocónicas (sólo en el flagelómero 1 o sólo en flagelómeros 8 y 12) ........ 50 Ala sin macrotriquias; mandíbulas reducidas, vestigiales; ojos con o sin pubescencia; uñas pequeñas, iguales, en todas las patas ........................................... ............................................................................................... Notiohelea GROGAN & WIRTH Ala con macrotriquias en el 1/4 apical; mandíbulas dentadas; ojos pubescentes; sólo patas anterior y media con uñas pequeñas, iguales, o uñas desiguales en todas las patas (como en Fig. 14F) ...................................................... 51 Celdas radiales subiguales, pequeñas; R3 engrosada; uñas de la pata media desiguales y más o menos similares a las de la pata posterior; sólo flagelómero 1 con sensilas celocónicas ....................................... Borkenthelea SPINELLI & GROGAN Segunda celda radial aproximadamente el doble de largo que primera celda radial; R3 no engrosada; pata media con uñas pequeñas, iguales, más cortas que la uña única, alargada (con diente basal) de la pata posterior; sólo flagelómeros 8 y 12 con sensilas celocónicas ............................................ Austrohelea WIRTH & GROGAN Tarsómero 5 de la pata anterior hinchado (Figs. 15G, 17A, C); ................................ ................................................................................................. HETEROMYIINI..........53 Tarsómero 5 de la pata anterior no hinchado ............................................................ 56 M sésil, bifurcándose a nivel de r-m (Fig. 12G) ......... Physohelea GROGAN & WIRTH M anchamente sésil, bifurcándose anteriormente a r-m (Fig. 12H) ...................... 54 Fémur anterior hinchado, con numerosas espinas gruesas ventrales (Fig. 17C) .... ........................................................................................................................ Heteromyia SAY Fémur anterior delgado, a lo sumo con unas pocas espinas delgadas .................. 55 Tarsómero 4 de todas las patas cilíndrico o ligeramente cordiforme, sin espinas fuertes; pata posterior de la hembra llamativamente alargada, con tarsómero 2 al memos 3/4 del largo del tarsómero 1 (Fig. 15H); uña posterior muy larga, única o con diente basal pequeño, más larga que las uñas de las patas anterior y media (Fig. 15H) .................................................................................................... Pellucidomyia MACFIE Tarsómero 4 de la pata anterior cordiforme, tarsómeros 4 de las patas media y posterior bífido y espinoso (Fig. 17B); pata posterior de la hembra no mucho más larga que las otras patas, con tarsómero 2 menor que el doble del largo del tarsómero 1; ambas uñas posteriores aproximadamente del mismo largo que aquellas de las patas anterior y media .............................................. Clinohelea KIEFFER Genitalia masculina grande, con gonocoxitos fusionados dorsalmente y ventralmente (Fig. 21B); genitalia femenina con esternito 8 alargado, laminar (Fig. 23A) ........................................................................................................ Echinohelea MACFIE

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57(56) – 58(57) – 59(58) –

60(59) – 61(58) – 62(61) – 63(62) – 64(61)

–

65(64)

– 66(65) – 67(66) – 68(66) –

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Genitalia masculina pequeña o grande, si es grande entonces los gonocoxitos no están fusionados; genitalia femenina con esternito 8 moderadamente desarrollado .............................................................................................................................................. 57 Macho (macho de Groganhelea, Sphaerohelea desconocidos) ...................................... 58 Hembra ............................................................................................................................... 72 Gonostilo muy reducido en tamaño (Figs. 21D, 22B) o fusionado con el gonocoxito (Fig. 22D) ..................................................................................................... 59 Gonostilo grande, claramente presente (Figs. 21F, 22A, C) (macho de Lanehelea desconocido, pero probablemente se ubicaría en este sitio de la clave) .............. 61 Ala con 2 celdas radiales claramente definidas (Fig. 13I); gonostilo bien definido, con ápice puntiagudo (Fig. 21D) ..........................Clastrieromyia SPINELLI & GROGAN Ala con 1 celda radial o con, al menos, un indicio poco perceptible de división entre primera y segunda celda radial (Fig. 13D); gonostilo pequeño y redondeado apicalmente (Fig. 22B) o no claramente definido y fusionado con el gonocoxito (Fig. 22D) ........................................................................................................................... 60 Anepisterno con numerosas setas alargadas; gonocoxito corto y gonostilo presente pero pequeño (Fig. 22B) ..................................... Phaenobezzia HAESELBARTH Anepisterno sin setas; gonocoxito y gonostilo fusionados y alargados (Fig. 22D) .................................................................................................................Nilobezzia KIEFFER Genitalia con gonocoxito y tergito 9 alargados (Fig. 22C) ..................................... 62 Genitalia más robusta, con gonocoxito y tergito 9 no inusualmente alargados (Figs. 21F, 22A) ................................................................................................................. 64 Ala con 1 celda radial (como en Fig. 13D) ...... Neobezzia WIRTH & RATANAWORABHAN Ala con 2 celdas radiales (Fig. 13G) ............................................................................ 63 Fémur anterior con espinas guesas ventrales ................... Austrosphaeromias SPINELLI Fémur anterior sin espinas gruesas ventrales .............................. Mallochohelea WIRTH (Nota. Los machos de algunas especies de Mallochohelea no tienen genitalia alargada) Tarsómere 5 de al menos la pata posterior con espinas robustas, romas (Fig. 17H); ápice del tergito 9 con una hilera de setas gruesas, robustas (Fig. 21C) .................. ....................................................................................................... Johannsenomyia MALLOCH Tarsómero 5 sin espinas gruesas, o si están presentes, son más delgadas y finamente puntiagudas (Fig. 16A); tergito 9 con setas dispersas, nunca dispuestas en una hilera de setas gruesas, robustas ...................................................................... 65 Con 1-2 setas robustas en el ápice de los cercos (Fig. 21E); parámeros fusionados, fusión restricta a la porción más anteromedia (Fig. 21E) ............ ............................................................................................. Amerohelea GROGAN & WIRTH Con algunas a muchas setas en los cercos; parámeros separados o con diferente grado de fusión ........................................................................................... 66 Fémur anterior con espinas robustas (Fig. 18A) ....................................................... 67 Fémur anterior sin espinas robustas ............................................................................ 68 Ala con 1 celda radial (Fig. 13C) ........................................ Bezzia KIEFFER (en parte) Ala con 2 celdas radiales (Fig. 13B) .............................. Palpomyia MEIGEN (en parte) Palpo con 4 segmentos ............................................................... Stenoxenus COQUILLETT Palpo con 5 segmentos ................................................................................................... 69

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69(68) Tórax delgado y aplanado dorsoventralmente; paratergito con numerosas setas pequeñas; 1 celda radial (Fig. 12I) .......................................... Paryphoconus ENDERLEIN – Tórax más robusto y no aplanado dorsoventralmente; paratergito sin setas; 1-2 celdas radiales .................................................................................................................... 70 70(69) Ala con 1 celda radial (Fig. 13C) ........................................ Bezzia KIEFFER (en parte) – Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 71 71(70) Fémur posterior hinchado (Fig. 18C); abdomen blanco, contrastando con el tórax oscuro ........................................................................................................Pachyhelea WIRTH – Fémur posterior no hinchado (Fig. 18A); abdomen claro a pardo oscuro, similar al tórax oscuro ........................................................................ Palpomyia MEIGEN (en parte) 72(57) Ala con M2 fuertemente curvada cerca de la base (Fig. 13A); palpo con 4 segmentos ...................................................................................... Stenoxenus COQUILLETT – Ala con M2 no curvada cerca de la base, o sólo ligeramente (Figs. 13B-I); palpo con 5 segmentos ............................................................................................................... 73 73(72) Tarsómero 5 armado ventralmente con espinas robustas, negras, romas (Figs. 16A, B, 17G-I) (estas espinas algo puntiagudas en Austrosphaeromias); abdomen sin apodemas tergales; esternito 8 con o sin (sólo Sphaerohelea) mechones de pelos posterolaterales, con setas largas, curvadas, superpuestas en la línea media (mejor visión a 100X) (Fig. 23B);...................... SPHAEROMIINI..........74 – Tarsómero 5 inerme, o si existen algunas espinas ventrales, son delgadas y puntiagudas (Fig. 16E); abdomen con (Figs. 23C-E) o sin apodemas tergales (no visibles en algunos ejemplares jóvenes o pobremente esclerotizados); esternito 8 sin mechones de pelos ................................................................................................ 81 74(73) Fémures sin espinas; uñas iguales en la pata anterior, desiguales en las patas media y posterior (Figs. 17G-I) .......................................................... Johannsenomyia MALLOCH – Fémures con o sin espinas; uñas iguales en todas las patas o sólo desiguales en la pata anterior ...................................................................................................................... 75 75(74) Uñas suavemente curvas distalmente, con diente basal delgado en el margen interno ................................................................................................................................ 76 – Uñas rectas o aplanadas distalmente, con diente basal delgado o romo en el margen externo (Figs. 16A, B) ...................................................................................... 78 76(75) Fémur anterior hinchado, con numerosas espinas ventrales gruesas ........................ .................................................................................................... Austrosphaeromias SPINELLI – Fémur anterior delgado, inerme ................................................................................... 77 77(76) Costa extendiéndose hasta el extremo distal del ala; tarsómero 5 con 2-3 pares de espinas ventrales .............................................. Sphaerohelea SPINELLI & FELIPPE-BAUER – Costa no extendiéndose más allá de 0,85 del largo del ala; tarsómero 5 con 4-5 pares de espinas ventrales ............................................... Lanehelea WIRTH & BLANTON 78(75) Ala con 1 celda radial; costa larga, extendiéndose casi hasta el extremo distal del ala; relación costal mayor que 0,87; fémures sin espinas ventrales ....................... 79 – Ala con 1-2 celdas radiales; costa corta, no llegando a 0,84 del largo del ala; fémures con o sin espinas ventrales ............................................................................. 80

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79(78) Diente basal externo de las uñas tarsales puntiagudo (Fig. 16A); tarso de la pata posterior no alargado (Fig. 16B); ala con lóbulo anal pobremente desarrollado ... ..............................................................................Neobezzia WIRTH & RATANAWORABHAN – Diente basal externo de las uñas tarsales romo (Fig. 16C); tarso de la pata posterior alargado (Fig. 16D); ala con lóbulo anal bien desarrollado (Fig. 13F) .... ...................................................................................... Groganhelea SPINELLI & DIPPOLITO 80(78) Ala con 2 celdas radiales (Fig. 13G); ala grisácea, con nervaduras anteriores pardas oscuras; fémur anterior sin espinas gruesas ventrales ............... Mallochohelea WIRTH – Ala con 1 celda radial; ala blancuzca, nervaduras anteriores claras; fémur anterior con espinas gruesas ventrales ...........................................................Nilobezzia KIEFFER 81(73) Tórax delgado y aplanado dorsoventralmente; paratergito con numerosas setas pequeñas; 1 celda radial (Fig. 12I) .......................................... Paryphoconus ENDERLEIN – Tórax más robusto y no aplanado dorsoventralmente; paratergito sin setas; ala con 1-2 celdas radiales; ................................................................... PALPOMYIINI..........82 82(81) Abdomen con 1 par de apodemas tergales que surgen cerca de los márgenes laterales del esternito 7; 1 espermateca presente, sin trazos ni evidencias de una segunda reducida ............................................................. Amerohelea GROGAN & WIRTH – Abdomen con 1 o más pares de apodemas tergales (Figs. 23C-E), si sólo está presente 1 par, estos apodemas surgen más cerca de la línea media en el margen posterior del esternito 7; 1-2 espermatecas, pero si existe sólo 1, una segunda reducida está presente ..................................................................................................... 83 83(82) Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 84 – Ala con 1 celda radial (Fig. 13C) .................................................................................. 86 84(83) Fémur anterior sin espinas robustas; fémur posterior muy hinchado (Fig. 18C); abdomen blanco, contrastando con el tórax oscuro ........................Pachyhelea WIRTH – Fémur anterior con o sin espinas robustas (Fig. 18A); fémur posterior no muy hinchado (Fig. 18A); abdomen claro a pardo oscuro, similar al tórax oscuro ........... 85 85(84) Flagelo corto, sólo 1,2-1,4 veces más largo que el ancho de la cabeza; membrana alar sin pigmentación; ala con lóbulo anal bien desarrollado (Fig. 13I); abdomen con un par de apodemas que surgen del margen anterior del tergito 7 y se extienden hasta el margen anterior del tergito 6 (Fig. 23C) ........................................ ......................................................................................Clastrieromyia SPINELLI & GROGAN – Flagelo alargado, a menos 2,5 veces más largo que el ancho de la cabeza; membrana alar pigmentada; ala con lóbulo anal pobremente desarrollado (Fig. 13B); abdomen con 2 o más pares de apodemas alargados que surgen del margen anterior de al menos los tergitos 6-7 y se extienden anteriormente dentro de cavidad abdominal (Fig. 23E) ............................................................ Palpomyia MEIGEN 86(83) Costa moderadamente corta, relación costal 0,67-0,75 (Fig. 13C); fémur anterior con (Fig. 18B) o sin espinas robustas; tarsómero 5 con o sin setas ventrales robustas .................................................................................... Bezzia KIEFFER (en parte) – Costa más larga, relación costal ratio mayor que 0,80 (Fig. 13D); fémur anterior sin espinas robustas; tarsómero 5 con setas ventrales delgadas con extremos afinados y doblados (Fig. 16E) ........................................... Phaenobezzia HAESELBARTH

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CLAVE PARA LOS HUEVOS DE GÉNEROS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL Solamente han sido descriptos los huevos de pocas especies de ceratopogónidos (Table 1). La clave siguiente es, por lo tanto, muy tentativa. No se hallan disponibles claves para huevos de Diptera en las cuales los huevos de Ceratopogonidae pudan ser distinguidos de aquellos pertenecientes a otras familias. Sin embargo, algunos Ceratopogonidae poseen huevos que son diferentes de todos los demás (Dasyhelea, Stenoxenus y Paryphoconus) o las oviposiciones tienen lugar en una matriz característica (Alluaudomyia y la mayoría de los Sphaeromiini). 1 – 2(1) – 3(2) – 4(3)

– 5(4) –

Huevo en forma de C (Fig. 24A) ...................................................................... Dasyhelea Huevo, a lo sumo, con curvatura ligera (Figs. 24B-F) ............................................... 2 Huevo con collar característico subapical y constricción terminal a manera de pezón en el extremo anterior (Fig. 24F) .............................. Stenoxenus y Paryphoconus Huevo sin ornamentación en ninguno de los extremos, pequeñas espículas pueden estar presentes (Figs. 24A-E) ............................................................................ 3 Huevo oval (Fig. 24B) .................. Leptoconops (en parte), Forcipomyiinae (en parte) Huevo alargado, delgado (Figs. 24C-E) ........................................................................ 4 Huevos ordenados en una banda alargada, dispuestos en ángulo con respecto al eje longitudinal de la banda (Fig. 24E) ............................................................................ ...... mayoría de Sphaeromiini (conocidos para Johannsenomyia, Mallochohelea, Macropeza) Huevos colocados separadamente o en grupos; si están colocados en banda, los huevos se hallan lado a lado (Figs. 24C, D) ................................................................. 5 Huevos colocados en banda (Fig. 24D) ............... Alluaudomyia (al menos en parte) Huevos colocados separadamente o en grupos aislados (aunque no fijados uno otro longitudinalmente) (Fig. 24C) ......................................... otros Ceratopogonidae

CLAVE PARA LAS LARVAS DE GÉNEROS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL Los inmaduros de Ceratopogonidae se hallan entre los más pobremente conocidos entre todas las familias de nematóceros (Table 2). Se brinda aquí una clave tentativa para algunos géneros comunes. El primer autor de esta contribución está preparando una clave, a nivel genérico, para larvas y pupas de géneros de todo el Mundo. Es importante destacar que sólo 11 de los 51 géneros conocidos de Ceratopogonidae Neotropicales han sido descriptos como larvas sobre la base de material de esta región (Table 2), mientra que 11 otros géneros se conocen como larvas a partir de material de otras regiones. 1

–

Pseudópodo presente en el protórax (Figs. 25B, C); cápsula cefálica hipognata a prognata (Figs. 25B, C); con setas fuertes en cápsula cefálica y cuerpo (Figs. 25B, C) (Forcipomyiinae) ........................................................................................................... 2 Sin pseudópodo en el tórax (Figs. 25A, D-F); cápsula cefálica algo hipognata o prognata (Figs. 25A, D-F); sin setas fuertes en cápsula cefálica y cuerpo anteriormente al segmento anal (Figs. 25A, D-F) ....................................................... 3

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– 4(3)

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Cuerpo algo aplanado dorsoventralmente, con tubérculos laterales (Fig. 25B) ..... ............................................................................................................................. Atrichopogon Cuerpo cilíndrico en sección transversal, sin tubérculos laterales (Fig. 25C) ......... ............................................................................................................................... Forcipomyia Cápsula cefálica dirigida anteroventralmente; finalización del abdomen con serie de ganchos, a veces dispuestos como dos pseudópodos (a menudo retraídos y dificil de observar en ejemplares preservados en alcohol), nunca con setas alargadas (Fig. 25D) ............................................................................................. Dasyhelea Cápsula cefálica prognata; finalización del abdomen simple, sin ganchos, con o sin setas alargadas (Figs. 25A, E-F) ...................................................................................... 4 Cápsula cefálica con pigmentación pobremente desarrollada, con apodemas bien desarrollados dirigidos posteriormente y que se extienden en el protórax (Figs. 26A, B); abdomen con segmentos divididos secundariamente (Fig. 25A) .............. ............................................................................................................................... Leptoconops Cápsula cefálica bien desarrollada, sin apodemas posteriores al margen de cápsula cefálica (Figs. 26C-F); abdomen con segmentos normales, no divididos secundariamente (Figs. 25E, F) (Ceratopogoninae) .................................................... 5 Cápsula cefálica con todas las setas simples (Figs. 26C, D) ............... Culicoides Cápsula cefálica con una o más setas s, u y la posterior o plumosa (Figs. 26E, F, seta u no visible, generalmente lateral sobre la cápsula cefálica) .............. 6 Margen posteroventral de la cápsula cefálica con proceso que se proyecta posteriormente (Fig. 26F) ........................................................ Sphaeromiini (en parte) Margen posteroventral de la cápsula cefálica como una línea recta o algo curvada, sin proceso que se proyecta posteriormente (Fig. 26E) .............................................. ................................................. Sphaeromiini (en parte), Palpomyiini, Ceratopogonini

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8 SYNOPSIS OF THE CERATOPOGONIDAE OF THE NEOTROPICAL REGION References providing available keys to the species of the various genera are also given in the catalog in Table 4. Allohelea KIEFFER. Only a single species occurs in the Neotropical Region, A. neotropica WIRTH from Central America and Jamaica, but six species are known from the Nearctic (WIRTH, 1991a). Males are the only members in the family with massive, enlarged hind legs and a single hind claw, as in the predaceous female. Alluaudomyia KIEFFER. Nineteen species occur in the Neotropical Region. Of these species, seventeen were reviewed by SPINELLI & WIRTH (1984c). These small, brightly patterned predaceous midges have larvae that inhabit ponds, lakes and streams where they are unique in swimming on the surface film. Amerohelea GROGAN & WIRTH. Eleven species have been recorded from the Neotropical Region. Of these species, ten were reviewed by GROGAN & WIRTH (1981). Females differ from other members of the tribe Palpomyiini in having only a single spermatheca and a single pair of internal abdominal tergal apodemes that arise near the lateral margins of segment 7. Atrichopogon KIEFFER. One hundred species are known from the Neotropical Region with 18 recently described from Costa Rica (BORKENT & PICADO, 2004) and five from Patagonia (SPINELLI et al., 2006). Six new species from Peruvian Amazonia are being presently described (SPINELLI & MARINO, in prep.). The genus is clearly very diverse in the Neotropical Region and we suspect that hundreds of species remain to be described. Adults are among the most commonly encountered diurnal biting midges and can be found on a variety of flowers (mostly small, white or green) or resting on vegetation. The adult males of at least some species form swarms. Larvae are aquatic to terrestrial and have distinctive elongate dorsal setae (a feature shared with many Forcipomyia). The feeding habits of the adults are poorly known but some suck blood from blister beetles (WIRTH 1956a, b; BORKENT & ROCHA-FILHO, 2006), while others feed on dead insects, are pollinovorous or are non-feeding (autogenous). The larvae and pupae have important taxonomic features. EWEN & SAUNDERS (1958) reared nine species from several localities in the Neotropics. Austrohelea WIRTH & GROGAN. Only a single species occurs in the Neotropical Region, A. shannoni WIRTH & BLANTON from the temperate Nothofagus forests of Argentina and Chile. The genus exhibits a transantarctic distribution, with six other species known

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from Australia and New Zealand (WIRTH & GROGAN, 1988). At least three unnamed species have been collected from Patagonia. Austrosphaeromias SPINELLI. The genus is known only from two species from southern Argentina and Chile (SPINELLI, 1997a). It is the only genus of the tribe Sphaeromiini in the Neotropical Region inhabiting temperate Nothofagus forests. Baeodasymyia CLASTRIER & RACCURT. Five species of these minute, highly reduced biting midges are known in the Neotropical Region (BORKENT & CRAIG, 1999). Adults walk with a distinctive rapid, stuttering gait (best seen with a hand lens). The immatures are restricted to the headwaters of springs and small seeps. Baeohelea WIRTH & BLANTON. Only a single species (B. nana WIRTH & BLANTON) of this minute, highly reduced predaceous genus is known in the Neotropical Region, distributed from Costa Rica to Ecuador. Baeohelea and its sister group Baeodasymyia are the only two genera in the family to have their palpi reduced to two segments. Bezzia KIEFFER. Forty-six species occur in the Neotropical Region. Most species are shining dark black, but many have striking banded leg patterns. This is the most common genus in the tribe Palpomyiini in the region and the second most common predaceous genus after Stilobezzia. The larva of at least one species in Costa Rica lives in bromeliads. Others likely occur in larger water bodies such as rivers and lakes. Borkenthelea SPINELLI & GROGAN. The genus is known from four species from the temperate Nothofagus forests of southern Argentina and Chile (SPINELLI & GROGAN, 2001). Males are unique in the tribe Ceratopogonini in having flagellomeres 3-10 fused. Brachypogon KIEFFER. Twenty-five species of these small, black midges are known from the Neotropical Region. Four new species from the Peruvian Amazonia are being described (SPINELLI & MARINO, in prep.). One undescribed species is common in some rocky coastal areas in Costa Rica and an additional 11 new species are known from Costa Rica. This suggests that numerous further species remain to be discovered in this region. Cacaohelea WIRTH & GROGAN. Only a single described species of these tiny midges is known, and the original specimens came from a cacao plantation in Costa Rica (WIRTH & GROGAN, 1988). Further material, including that of a second undescribed species, has been collected in Costa Rica in areas devoid of cacao. Ceratoculicoides WIRTH & RATANAWORABHAN. Within the region, only one species is known from Mexico, with a second, unnamed species from Costa Rica known from two female adults (HUERTA & BORKENT, 2005). All remaining species are restricted to the Holarctic Region (WIRTH & GROGAN, 1988). Clastrieromyia SPINELLI & GROGAN. The genus is known from four Neotropical species (SPINELLI & GROGAN, 1985, 1986). Females differ from other members of the tribe Palpomyiini in having only a single pair of internal abdominal tergal apodemes arising from the anterior margin of tergite 7 and extending to about the anterior margin of tergite 6. Clinohelea KIEFFER. Fourteen species of this predaceous genus occur in the Neotropical Region (SPINELLI & DURET, 1993). The elegant adults have long, slender legs and large dark maculations on their wings. Culicoides LATREILLE. Two hundred and sixty-six species of this vertebrate-biting genus have been recorded in the region. Of these, only 16 are serious pests of humans or domestic animals. The most serious coastal species biting humans in Central America and

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northern South America is C. furens, attacking on beaches and in homes near mangrove swamps. They may occur in such huge numbers that they provide the important ecological service of restricting humans from living in some areas. Various pest species are discussed above in the section on Bionomics. The adult males of at least some species form swarms. Larvae of species of Culicoides are semiaquatic (wet mud, mosses, rotting vegetation) to aquatic and feed on microorganisms and small invertebrates. There are clearly a significant number of species remaining to be discovered, especially from higher elevations in the Neotropical Region. Dasyhelea KIEFFER. Fifty-eight species of this common, mainly diurnal genus are recorded from the Neotropics, but many more remain undescribed. At least eleven new species are known from Patagonia (DÍAZ & SPINELLI, in prep.). Adults have reduced, nonbiting mouthparts but are common on many flowers (particularly those that are small and white) where they feed on nectar. Adults may be common in some coastal habitats, particularly where there are seeps, pools and, for a few species, in flat rocky areas. The adult males of at least some species form swarms. The larvae are aquatic grazers on algae and detritus and are generally present in natural small aquatic habitats (e.g. tree holes, rock pools) or artificial ones (e.g. cemetery flower-holding containers). This very diverse and common genus is in urgent need of revision. Diaphanobezzia INGRAM & MACFIE. The genus is known from four species from Argentinean Patagonia. The peculiar halter, without a constriction below the knob, is unique within the family. Downeshelea WIRTH & GROGAN. Eighteen species occur in the Neotropical Region (LANE & WIRTH, 1964). They have distinctive large blotches on their wings and may be reasonably common in some Malaise trap samples. Echinohelea MACFIE. Eleven species of this genus are known in the Neotropical Region (WIRTH, 1994b). The distinctive adults are patterned and bright yellow to golden brown in color, have numerous spines on their legs and both sexes have long proboscis and elongate antennal flagella. The species are strikingly similar to one another and therefore somewhat difficult to identify. Specimens are occasionally common in Malaise trap samples. Fittkauhelea WIRTH & BLANTON. The genus is known from a single species, F. amazonica WIRTH & BLANTON from Brazilian Amazonia. Females are similar to females of Parabezzia except for the presence of tarsal claws with an basal inner tooth. Forcipomyia MEIGEN. Two hundred and ten species of this common genus occur in the Neotropical Region. Adults of some species are ectoparasites of large insects (as described in the section on Bionomics above) but those of the subgenus Lasiohelea feed on vertebrate blood. Although adult females are common and many have biting mouthparts, the hosts for most of the species remain unknown. The adult males of at least some species form swarms. Most larvae have elongate dorsal setae (also present in Atrichopogon), are aquatic to terrestrial, and are found in a diverse array of habitats such as in mosses, under the bark of rotting trees, inside decaying fruits, and in aquatic epiphytes. Many species remain to be described. Groganhelea SPINELLI & DIPPOLITO. This genus of the tribe Sphaeromiini is known from a single species, G. rondoniensis SPINELLI & DIPPOLITO, from Brazilian Amazonia and Uruguay. Males are unknown.

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Heteromyia SAY. Eleven species occur in the Neotropical Region (DURET & LANE, 1955; WIRTH & GROGAN, 1977). The distinctive, large adult females have greatly swollen fore femora with dense ventral spines and extremely elongated hind legs, each with a single, very long claw which they project upward when at rest. Occasionally adults may be abundant. Isthmohelea INGRAM & MACFIE. The genus is known from a single species, I. disjuncta INGRAM & MACFIE, from its type-locality, in the Valdivian rain forests of southern Chile. The male has a strikingly elongate abdominal segment 9. Johannsenomyia MALLOCH. Only two species are known from the Neotropical Region. The larvae inhabit rivers and streams. Lanehelea WIRTH & BLANTON. This genus of the tribe Sphaeromiini is known from two species from Colombia. Males are unknown. Leptoconops SKUSE. Twelve species of this vertebrate blood sucking genus occur in the Neotropical Region. Females, especially those of L. bequaerti, L. chilensis FORATTINI and L. petrocchiae SHANNON & DEL PONTE, are often serious daytime biting pests of humans. Adults of both sexes have the unusual habit of burrowing into sand to rest and for females to lay their eggs. The larvae are pink and primarily inhabit wet, protected sands on beaches of oceans and lakes. Leptohelea WIRTH & BLANTON. The genus is known from a single female specimen, L. micronyx WIRTH & BLANTON (WIRTH & BLANTON, 1970a), from Colombia and one male from Costa Rica. The maxillary palpus of adults is 3-segmented. Males are included here, in a key, for the first time. Macrurohelea INGRAM & MACFIE. Twelve species occur in the Neotropical Region, particularly in Patagonia. SPINELLI & GROGAN (1990) provided a key to males and females of eleven of these species. The genus exhibits a transantarctic distribution, being known from 3 other species from Australia (GROGAN & WIRTH, 1985). Females are unique in the family in having the tenth segment elongated and bent anteroventrally. Mallochohelea WIRTH. Six species are known from the Neotropical Region. None are common. Monohelea KIEFFER. Eighteen species occur in the Neotropical Region, the majority of them included in a key to males and females by LANE & WIRTH (1964). Adults have wings with characteristic black and gray maculations, brightly banded legs and females have single large hind claws. Nannohelea GROGAN & WIRTH. Only one named species of this minute predaceous genus, N. clastrieri GROGAN & WIRTH, occurs in the Neotropical Region, from Colombia (GROGAN & WIRTH, 1980a, 1990). Another undescribed species is known from Costa Rica, based on one male. Otherwise, three species are recorded from the Old World. Neobezzia WIRTH & RATANAWORABHAN. Eight of these rare species occur in the Neotropical Region. WIRTH & RATANAWORABHAN (1972a) provided a key to females of seven of these species. Nilobezzia KIEFFER. Three species occur in the Neotropical Region. LANE (1961b) provided a key to males and females. Adults may be locally common, especially around lowland permanent ponds. Notiohelea GROGAN & WIRTH. The genus is known from two species from the temperate Nothofagus forests of southern Argentina and Chile (GROGAN & WIRTH, 1979b; SPINELLI & GROGAN, 1990). Males are unknown.

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Pachyhelea WIRTH. Two species occur in the Neotropical Region, the wide-ranging P. pachymera (WILLISTON) and P. albidiventris (KIEFFER) from Colombia (GROGAN & WIRTH, 1980c). It is similar to Palpomyia. Palpomyia MEIGEN. Forty six species occur in the Neotropical. The adult females feed on small Diptera or mayflies (GROGAN & WIRTH, 1979c). Parabezzia MALLOCH. Twenty-four species occur in the Neotropical region (SPINELLI & GROGAN, 1987). Paradasyhelea MACFIE. Three species are restricted to southern Chile and Argentina (SPINELLI & GROGAN, 2003). Seven further species are known from the Australasian region, and a single species occurs in the Nearctic region. Parastilobezzia WIRTH & BLANTON. This rare genus is known from a single species, P. leei WIRTH & BLANTON, from Costa Rica and Colombia (WIRTH & GROGAN, 1988). Paryphoconus ENDERLEIN. Forty species occur in the Neotropical Region. SPINELLI & WIRTH (1984b) provided a key to females of thirty eight of these species. The adults are medium-sized to large (female wing lengths 1.5-7 mm) and are generally rare. The few abundant collections from the Neotropical Region are from near rivers. Pellucidomyia MACFIE. Four species occur in the Neotropical Region (WIRTH & RATANAWORABHAN, 1971b). Adults are rare and females resemble those of Heteromyia in having greatly elongated hind legs and a single elongate claw, but they lack the swollen, spinose forefemur of that genus. Phaenobezzia HAESELBARTH. Three species are broadly distributed in the Neotropical Region (SPINELLI & WIRTH, 1986a). Physohelea GROGAN & WIRTH. The genus is known only from two species from southern Argentina and Chile (GROGAN & WIRTH, 1979a). It is the only genus of the tribe Heteromyiini in the Neotropical Region inhabiting temperate Nothofagus forests. Rhynchohelea WIRTH & BLANTON. A single species of this monotypic genus, R. monilicornis WIRTH & BLANTON, is known from Florida and Costa Rica (WIRTH & GROGAN, 1988). Within the Neotropical Region, this minute relative of Brachypogon is known from only one female specimen from Costa Rica (BORKENT, 1992). Schizonyxhelea CLASTRIER. Two species of these small, reduced relatives of Stilobezzia are known from the Neotropical Region (BORKENT, 2000b). The larvae of one species were collected from a moderately-sized spring in Costa Rica. Sphaerohelea SPINELLI & FELIPPE-BAUER. A single species of this monotypic genus, S. biestroi SPINELLI & FELIPPE-BAUER, is known from northeastern Argentina (SPINELLI & FELIPPE-BAUER, 1990b). Males are unknown. Stenoxenus COQUILLETT. Sixteen species occur in the Neotropical Region). WIRTH & RATANAWORABHAN (1972b) provided a key to females of fourteen of these species. This genus is closely related to Paryphoconus, the only two genera in the tribe Stenoxenini, and the eggs of both genera have a terminal nipple-like constriction with a fringed subapical collar. Female Stenoxenus have a strikingly different M2 wing vein (Fig. 13A) compared to their males and other members of the family. Adults may be common near rivers. Stilobezzia KIEFFER. Sixty-four species of these common, often brightly colored and patterned predaceous midges are recorded from the Neotropical Region (LANE & FORATTINI, 1961). However, the genus is clearly diverse in the region. For example, we have

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examined 87 species from Costa Rica alone and there are undoubtedly many more yet to be discovered in that small country. At least twelve new species in the subgenus Acanthohelea, from Nothofagus forest in Patagonia, are being described (CARZOLA & SPINELLI, in prep.). Undescribed genus near Parabezzia MALLOCH. A single species is known as male and female adults from Costa Rica. It will be fully described in a future publication.

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9 REFERENCES AITKEN, T.H.G., WIRTH, W.W, WILLIAMS, R.W, DAVIES, J.B & E.S. TIKASINGH (1975): A review of the bloodsucking midges of Trinidad and Tobago, West Indies (Diptera: Ceratopogonidae). - J. Entomol. (B) 44: 101-144. ATCHLEY, W.R. & W.W. WIRTH (1979): A review of the Culicoides haematopotus group in North America (Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 52: 524-545. BARBOSA, F.A.S. (1943): Descrição de “Culicoides recifensis” n. sp. e do macho de “Culicoides reticulatus” LUTZ (Diptera, Chironomidae). - Rev. Bras. Biol. 3: 261-264. BARBOSA, F.A.S. (1947): Culicoides (Diptera: Heleidae) da região neotropical. - Ann. Soc. Biol. Pernambuco 7: 3-30 (Nov.). BARBOSA, F.A.S. (1951): A change of specific name in the genus Culicoides (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 53: 163. BARBOSA, F.A.S. (1952): Novos subsidios para o conhecimento dos Culicoides neotrópicos (Diptera: Heleidae). - Thesis, Univ. Recife, Recife: Imprensa Industrial: 21 pp. BARBOSA, F.A.S. (1953): Novos subsidios para o conhecimento dos Culicoides neotrópicos (Diptera: Heleidae). - Pub. Avulsas Inst. Aggeu Magalhães 2: 11-41. BARRETTO, M.P. (1944): Sobre o género Culicoides LATREILLE, 1809, com a descrição de trés novas espécies (Diptera, Ceratopogonidae). - Ann. Fac. Med. Univ. São Paulo 20: 89-105, pls. 1-4. BECK, E.C. (1951): A new Culicoides from Florida (Diptera, Ceratopogonidae). - Fla. Entomol. 34: 135-136. BECK, E.C. (1956): A new species of Culicoides from Florida with additional distribution data for the genus (Diptera: Heleidae). - Fla. Entomol. 39: 133-138. BLANTON, F.S. & W.W. WIRTH (1979): The sand flies (Culicoides) of Florida (Diptera: Ceratopogonidae). -Arthropods of Florida and neighboring land areas 10: xv + 204 pp. BORKENT, A. (1991): The Ceratopogonidae (Diptera) of the Galápagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. - Entomol. Scand. 22: 97-122. BORKENT, A. (1992): A new key to some genera of Ceratopogonini in the Holarctic (Diptera: Ceratopogonidae). - Entomol. Scand. 22: 433-436. BORKENT, A. (1996): Biting midges (Ceratopogonidae: Diptera) feeding on a leatherback turtle in Costa Rica. - Brenesia 43-44: 25-30 (1995). BORKENT, A. (2000a): Biting midges (Ceratopogonidae: Diptera) from Lower Cretaceous Lebanese amber with a discussion of the diversity and patterns found in other ambers. - In: GRIMALDI, D. (ed.): Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey: 355-452. Backhuys Publ., Leiden: viii + 498 pp.

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BORKENT, A. (2000b): The larva and pupa of Schizonyxhelea forattinii WIRTH and GROGAN (Diptera: Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus. - Proc. Entomol. Soc. Washington 102: 862-868. BORKENT, A. (2004): 10. Ceratopogonidae. - In: MARQUARDT, W.C. (ed.): Biology of disease vectors: 113-126. 2nd edition. Elsevier Academic Press, Amsterdam: xxiii + 785 pp. BORKENT, A. & D.A. CRAIG (1999): A revision of the Neotropical genus Baeodasymyia CLASTRIER and RACCURT (Diptera: Ceratopogonidae) with a discussion of their phylogenetic relationships. Am. Mus. Novitates 3274: 1-26. BORKENT, A. & D.A. CRAIG (2001): Submerged Stilobezzia rabelloi LANE (Diptera: Ceratopogonidae) pupae obtain oxygen from the aquatic fern Salvinia minima BAKER. - Proc. Entomol. Soc. Washington 103: 655-665. BORKENT, A. & D.A. CRAIG (2004): Austroconops WIRTH and LEE, a Lower Cretaceous genus of biting midges yet living in Western Australia: a new species, first description of the immatures and discussion of their biology and phylogeny (Diptera: Ceratopogonidae). - Am. Mus. Novitates 3449: 1-67. BORKENT, A. & L. FORSTER (1986): Review of the Dasyhelea fasciigera species group (Diptera: Ceratopogonidae) with a revision of the Nearctic species. - Can. J. Zool. 64: 1280-1287. BORKENT, A. & A. PICADO (2004): Distinctive new species of Atrichopogon KIEFFER (Diptera: Ceratopogonidae) from Costa Rica. - Zootaxa 637: 1-37. BORKENT, A. & L.C. ROCHA-FILHO (2006): First record of female adult Atrichopogon KIEFFER (Diptera: Ceratopogonidae) biting in the Neotropical Region. - Proc. Entomol. Soc. Washington 108: 998-1001. BORKENT, A. & G.R. SPINELLI (2000): Catalog of New World biting midges south of the United States (Diptera: Ceratopogonidae). - Contrib. Entomol. Internat. 4: 1-107. BORKENT, A. & W.W. WIRTH (1997): World species of biting midges (Diptera: Ceratopogonidae). Bull. Am. Mus. Nat. Hist. 233: 1-257. BRÈTHES, J. (1912): Descripción de un nuevo género y especie nueva de Chironomidae (Dipt). - An. Mus. Nac. Buenos Aires 15: 451-453. BRÈTHES, J. (1914): Descripción de seis Cécidomyidae (Dipt.) de Buenos Aires. - An. Mus. Nac. Buenos Aires 26: 151-156. BRICKLE, D.S. & D.V. HAGAN (1999): The Culicoides (Diptera: Ceratopogonidae) of Belize, Central America. - Insecta Mundi 13: 39-44. BROWN, B.V. (2005): Malaise trap catches and the crisis in Neotropical Dipterology. - Am. Entomologist 51: 180-183. BROWNE, J.E. (1980): New species of biting midges of the genus Culicoides from Colombia and the first description of the male of C. florenciae (Diptera: Ceratopogonidae). - J. Med. Entomol. 17: 533-544. BYSTRAK, P.G. & W.W. WIRTH (1978): The North American species of Forcipomyia, subgenus Euprojoannisia (Diptera: Ceratopogonidae). - U.S. Dept. Agric., Tech. Bull. 1591: 1-51. CARTER, H.F. (1921): A revision of the genus Leptoconops SKUSE. - Bull. Entomol. Res. 12: 1-28 (June). CAVALIERI, F. (1961a): Notas sobre Ceratopogonidae (Diptera, Nematocera) I. Forcipomyia (Forcipomyia) wygodzinskyi sp. n. para Tierra del Fuego. - Revta. Soc. Entomol. Argent. 23: 17-19. CAVALIERI, F. (1961b): Notas sobre Ceratopogonidae (Diptera, Nematocera) II. Sobre dos nuevas especies de Forcipomyia para Tierra del Fuego, Forcipomyia (F.) delpontei n. sp. y F. (F.) piroskyi. Acta Zool. Lilloana 18: 169-175 (1962).

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CAVALIERI, F. (1962): Notas sobre Ceratopogonidae (Dipt. Nematocera) III. Sobre un nuevo díptero hematófago para Argentina, Lasiohelea saltensis n. sp. y notas sobre las especies neotropicales de Lasiohelea. - Acta Zool. Lilloana 18: 359-365. CAVALIERI, F. (1966): Notas sobre Ceratopogonidae (Diptera, Nematocera) V. Descripción de una nueva especie de Venezuela: Culicoides (Oecacta) birabeni sp. n. - Physis 26: 59-63. CAVALIERI, F. & I.F. CHIOSSONE (1966): Datos sobre la familia de dípteros hematófagos Leptoconopidae en la Argentina, con la descripción de Leptoconops (Leptoconops) casali sp. n. y la redescripción de L. (L.) petrocchiae SHANNON y DEL PONTE, 1927. - Physis 26: 43-57. CAVALIERI, F. & I.F. CHIOSSONE (1972): Notas sobre el género Atrichopogon KIEFFER, 1906, de la Argentina (Diptera, Ceratopogonidae). I. Consideraciones generales y descripción de Atrichpogon delpontei sp. nov. - Revta. Soc. Entomol. Argent. 34: 119-129. CAVALIERI, F. & I.F. CHIOSSONE (1973): Notas sobre el género Atrichopogon KIEFFER, 1906, de la Argentina (Diptera, Ceratopogonidae). II. Descripción de Atrichpogon casali sp. nov. - Physis 32: 151-159. CAZORLA, C.G. & P.I. MARINO (2004): The pupa of Stilobezzia punctulata LANE from Peruvian Amazonia (Diptera: Ceratopogonidae). - Amazoniana 18: 75-80. CAZORLA, C.G., DÍAZ, F. & M.M. RONDEROS (2006): Redescription of pupa and adult of Stilobezzia fiebrigi KIEFFER 1917 (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 132: 111-119. CAZORLA, C.G., SPINELLI, G.R. & F. DÍAZ (2005): Two new species of the subgenus Stilobezzia (Stilobezzia) KIEFFER from Peruvian Amazonia (Diptera: Ceratopogonidae). - Amazoniana 18: 289-297. CHAN, K.L. & E.J. LEROUX (1965): Description of Forcipomyia (Neoforcipomyia) sp. n. and redescription of Forcipomyia (Neoforcipomyia) eques (JOHANNSEN) (Diptera: Ceratopogonidae), with an account of the digestive and reproductive systems. - Phytoprotection 46: 74-104. C HAN , K L. & E.J. L E R OUX (1967): Ecological studies on three pond midges (Diptera: Ceratopogonidae) in Quebec. - Ann. Soc. Entomol. Québec 12: 14-68. CHAN, K L. & E.J. LEROUX (1971): A new subgenus and species of midge (Diptera: Ceratopogonidae) from Singapore. - Can. Entomol. 103: 271-276. CHAUDHURI, P.K., DAS GUPTA, S.K. & D.K. CHAUDHURI (1972): Biting midges of the genus Alluaudomyia KIEFFER from India (Diptera: Ceratopogonidae). - Oriental Insects 6: 83-110. CHAVEZ, R., COLAND, E. & I. PHILLIPS (1992): Fiebre de oropouche en Iquitos. Reporte preliminar de 5 casos. - Rev. Farmacol. Terap. 2: 12-14. CLASTRIER, J. (1968) : Deux Ceratopogonides nouveaux de la Guyane Française. - Arch. Inst. Pasteur Guyane Franc. Territ.- L’Inini 21 : 85-92. CLASTRIER, J. (1971) : Deux nouveaux Culicoides (Diptera, Ceratopogonidae) de la Guyane Française. - Ann. Parasitol. 46 : 285-294. CLASTRIER, J. (1972) : Forcipomyia (F.) squamithorax n. sp. de la Guyane Française. - Bull. Soc. Entomol. France 77 : 170-176. CLASTRIER, J. (1973) : Le genre Leptoconops, sous-genre Holoconops dans le midi de la France (Dipt. Ceratopogonidae). - Ann. Soc. Entomol. France 9: 895-920. CLASTRIER, J. (1976): Alluaudomyia prima n. sp. De la Guyane Française (Diptera, Ceratopogonidae). Nouv. Rev. Entomol. 6: 205-207. CLASTRIER, J. (1979): Un nouvel Atrichopogon de la Guyane Française (Diptera: Ceratopogonidae). Rev. Franc. Entomol. 1: 30-32. CLASTRIER, J. (1984): Schizonyxhelea guyana n. g., n. sp. de la Guyane Française (Diptera, Ceratopogonidae). - Rev. Franc. Entomol. 6 : 1-4.

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CLASTRIER, J. (1987) : Atrichopogon ornatipennis n. sp. de la Guyane Française. - Entomologiste 43 : 271-274. CLASTRIER, J. (1991) : Description de cinq nouveaux Stilobezzia apparentés à S. Insolita DAS GUPTA & W IRTH , originaires des régions afrotropicale, paléarctique et néotropicale (Dipt. Ceratopogonidae). - Bull. Soc. Entomol. France 95: 297-310 (1990). CLASTRIER, J. (1992): Description de Palpomyia guyana n. sp. de la Guyane Française (Diptera, Ceratopogonidae). - Rev. Franc. Entomol. 14: 117-121. CLASTRIER, J. & J.-C. DELÉCOLLE (1994): Description de Forcipomyia (Phytohelea) musae n. sp. de la Guyane française (Diptera, Ceratopogonidae). - Rev. Franc. Entomol. 16: 51-56. CLASTRIER, J. & J.-C. DELÉCOLLE (1997): Description de Forcipomyia (Trichohelea) roubaudi n. sp., ectoparasite d’un hétéroptère réduvide capturé dans la canopée de la foret guyanaise (Diptera, Ceratopogonidae). - Bull. Soc. Entomol. France 102: 379-383. CLASTRIER, J. & J. LEGRAND (1990): Forcipomyia (Pterobosca) incubans (MACFIE) et F. (Trichohelea) macheti n. sp. Parasites des ailes de libellules en Guyane Française (Diptera, Ceratopogonidae; Odonata). - Rev. Franc. Entomol. 12: 167-170. CLASTRIER, J. & C. RACCURT (1979): Quatre nouveaux Parabezzia de la Republique d’Haiti (Diptera, Ceratopogonidae). - Nouv. Rev. Entomol. 9: 165-175. CLASTRIER, J. & C. RACCURT (1979b): Baeodasymyia modesta n. g., n. sp, de la République d’Haiti (Diptera, Ceratopogonidae). - Ann. Parasitol. 54: 99-104. CLASTRIER, J. & W.W. WIRTH (1978): The Leptoconops kerteszi complex in North America (Diptera: Ceratopogonidae). - U.S. Dept. Agric. Tech. Bull. 1573: 1-58. CLASTRIER, J. & W.W. WIRTH (1995) : Révision des Forcipomyia du sous-genre Microhelea de la région Néotropicale, parasites de phasmes (Diptera : Ceratopogonidae). - Ann. Soc. Entomol. France 31: 97-150. COQUILLETT, D.W. (1899): A new dipterous family related to the Chironomidae. - Entomol. News 10: 60-61. COQUILLETT, D.W. (1901a): New Diptera in the U.S. National Museum. - Proc. U.S. Nat. Mus. 23: 593-618. COQUILLETT, D.W. (1901b): Papers from the Hopkins Stanford Galapagos Expedition, 1898-1899. II. Entomological Results (2): Dipteria. - Proc. Wash. Acad. Sci. 3: 371-379. COQUILLETT, D.W. (1902): New Diptera from North America. - Proc. U.S. Nat. Mus. 25: 83-126. COQUILLETT, D.W. (1904a): New Diptera from Central America. - Proc. Entomol. Soc. Washington 6: 90-98. COQUILLETT, D.W. (1904b): A new Ceratopogon from Brazil. - J. New York Entomol. Soc. 12: 35-36. COQUILLETT, D.W. (1905): New nematocerous Diptera from North America. - J. New York Entomol. Soc. 13: 56-69. COQUILLETT, D.W. (1910): The type-species of the North American genera of Diptera. - Proc. U.S. Nat. Mus. 37: 499-647. CORDERO, E.H. (1929): Contribución al estudio de los Dipteros del Uruguay, I. Lophomyidium uruguayense n. Gen., n. Sp. Nueva Ceratopogonina hematofaga. - An. Mus. Hist. Nat. Montevideo 3: 93-108. COSTA LIMA, A. DA (1928): Ceratopogonineos ectoparasitos de phasmideos. - Mem. Inst. Oswaldo Cruz, Suppl. 3: 84-85, 2 pls. COSTA L IMA, A. DA (1937ª): Chave das espécies de Culicoides da região neotropical (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 32: 411-422. COSTA LIMA, A. DA (1937b): Primeira espécie americana do gênero Pterobosca (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 32: 615-616, pls. 1-2.

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PHILIPPI, R.A. (1865): Aufzählung der chilenischen Dipteren. - Verh. Kaiser.-König. Zool.-Botan. Gesell. Wien 15: 595-782. PIERCE, W.D. (1966): Fossil arthropods of California. 29. Silicified Miocene pupae of ceratopogonid flies. - Bull. South. Calif. Acad. Sci. 65: 81-98. PINHEIRO, F.P., TRAVASSOS DA ROSA, A.P.A. & F.C. VASCONCELOS (1998): An overview of oropouche fever epidemics in Brazil and neighbour countries. - In: TRAVASSOS DA ROSA, A.P.A. et al. (eds.): An overview of arbovirology in Brazil and neighbouring countries. Instituto Evandro Chagas, Belém: 296 pp. POEY, F. (1853): Memorias sobre la historia natural de la Isla de Cuba, acompañadas de sumarios latinos y extractos en francés. - Vol. 1 (pt. 4): 201-280, Havanna. RAFINESQUE, C.S. (1815): Analyse de la nature ou tableau de l’univers et des corps organisés. Le nature est mon guide, et LINNÉUS mon maitre.- Palmero, privately published: 224 pp. REMM, H. (1971): On the fauna of Ceratopogonidae of South Primorye (Ussuri Land) [in Russian, English summary]. - In: Living Nature of the Far East: 182-220, Akademiia nauk Estonskoi SSR: 240 pp. REMM, H. (1975): On the classification of the biting midges (Diptera, Ceratopogonidae) (in Russian). - Parazitol. 9: 393-397. ROBACK, S.S. (1957): A new Palpomyia from Peru (Diptera: Heleidae). - Not. Nat. Acad. Nat. Sci. Philadelphia 297: 1-2. RODRÍGUEZ, M.C. DE & W.W. WIRTH (1986): A new species of man-biting Culicoides from the high Andes of Colombia (Diptera: Ceratopogonidae). - Fla. Entomol. 69: 311-314. RONDEROS, M.M. (1988): Estudio sistemático de los géneros hematófagos de Ceratopogonidae (Diptera: Nematocera) de la República Argentina. - Ph.D.-thesis, Univ. Nac. La Plata, Argentina: 135 pp. + 49 pls. RONDEROS, M.M. (1990a): Dos especies nuevas del género Culicoides para la Argentina y Uruguay (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 48: 115-120. R ONDEROS , M.M. (1990b): A new species of Lepteconops [sic] from Argentina (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 34: 423-426. RONDEROS, M.M. & G.R. SPINELLI (1989): The pupa of the biting midge Culicoides venezuelensis ORTÍZ and MISRA 1950 (Diptera: Ceratopogonidae). - Biota 5: 77-81. R ONDEROS , M.M. & G.R. S PINELLI (1992): A key to the Neotropical Leptoconops (Diptera: Ceratopogonidae), with the description of a new species from Argentina. - Revta. Soc. Entomol. Argent. 51: 41-45. RONDEROS, M.M. & G.R. SPINELLI (1993): Dos especies nuevas de Leptoconops neotropicales, y descripcion del macho de Leptoconops (Leptoconops) chilensis (Diptera: Ceratopogonidae). - Gayana Zool. 57: 305-308. RONDEROS, M.M. & G.R. SPINELLI (1994): Dos especies nuevas de Culicoides de la Argentina (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 53: 47-50. RONDEROS, M.M. & G.R. SPINELLI (1995a): Two new neotropical species of Culicoides from the Paraná River Basin of Argentina and Paraguay (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 121: 59-63. RONDEROS, M.M. & G.R. SPINELLI (1995b): Redescripción de Culicoides lahillei y descripción de Culicoides ichesi, n. sp. de la Argentina, Paraguay y Uruguay (Diptera: Ceratopogonidae). - Neotrópica 41: 77-81. RONDEROS, M.M. & G.R. SPINELLI (1997): Culicoides patagoniensis n. sp. del sur de Argentina y Chile (Diptera: Ceratopogonidae). - Gayana Zool. 61: 33-39.

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RONDEROS, M.M. & G.R. SPINELLI (1998): Las especies de Culicoides en el área de influencia de la represa de Yacyretá (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 57: 79-83. RONDEROS, M.M. & G.R. SPINELLI (1999): On the subgenus Forcipomyia (Lasiohelea) in the Neotropical Region (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 125: 151-161(June). RONDEROS, M.M. & G.R. SPINELLI (2000): The larva and pupa of Culicoides bambusicola LUTZ observed with SEM, and additional notes on the adult (Diptera Ceratopogonidae). - Trans. Am. Entomol. Soc. 126: 133-144. RONDEROS, M.M., DÍAZ, F. & G.R. SPINELLI (2004a): A new species of Dasyhelea KIEFFER from Argentina described as adult and pupa (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 103: 193-200. RONDEROS, M.M., SPINELLI, G.R. & F. DÍAZ (2004b): Description of larva and redescription of pupa and adult of Palpomyia guarani (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 63: 45-54. RONDEROS, M.M., SPINELLI, G.R. & F. DÍAZ (2006): Observations on the immatures of Dasyhelea necrophila SPINELLI & RODRÍGUEZ in laboratory (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 132: 291-297. RONDEROS, M.M., SPINELLI, G.R., HUERTA, H. & F. DÍAZ (2003): Immature stages of two Neotropical species of Dasyhelea KIEFFER, 1911 (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 129: 295-308. RONDEROS, M.M., SPINELLI, G.R. & P. SARMIENTO (2000): Preparation and mounting of biting midges of the genus Culicoides LATREILLE (Diptera: Ceratopogonidae) to be observed with a scanning electron microscope. - Trans. Am. Entomol. Soc. 126: 125-132. ROOT, F.M. & W.A. HOFFMAN (1937): The North American species of Culicoides. - Am. J. Hyg. 25: 150-176, pls. 1-8. SÁENZ, M.R. & E.C. GREINER (1994): Culicoides aspirated from cattle in Costa Rica, Honduras, Panama and Puerto Rico, and their role as potential vectors of bluetongue viruses. - Med. Vet. Entomol. 8: 15-19. SAUNDERS, L.G. (1925): On the life history, morphology and systematic position of Apelma KIEFF. and Thyridomyia n. g. (Diptera, Nemat. Ceratopogoninae). - Parasitol. 17: 252-277. SAUNDERS, L.G. (1957): Revision of the genus Forcipomyia based on characters of all stages (Diptera, Ceratopogonidae). - Can. J. Zool. 34: 657-705 (1956). SAUNDERS, L.G. (1959): Methods for studying Forcipomyia midges, with special reference to cacaopollinating species (Diptera, Ceratopogonidae). - Can. J. Zool. 37: 33-51. SAUNDERS, L.G. (1964): New species of Forcipomyia in the Lasiohelea complex described in all stages (Diptera, Ceratopogonidae). - Can. J. Zool. 42: 463-482. SAUNDERS, L.G. & G.F. BOWMAN (1956): Cacao pollination in Costa Rica. - Banco de Costa Rica, San Jose: 13 pp. SAY, T. (1825): American entomology, or descriptions of the insects of North America. - Vol. 2. Philadelphia: 121 pp., pls. 19-36. SÉGUY, E. (1941): Quelques Cératopogonides vulnérants parasites des insectes. - Rev. Franc. Entomol. 8: 82-88. SHANNON, R.C. & E. DEL PONTE (1927): Cuatro notas sobre especies nuevas de Dipteros, Nematoceros, hematofagos o no, de la Republica Argentina. - Rev. Inst. Bacterial. Dept. Nac. Hig. 4: 724-736. SKUSE, F.A.A. (1889): Diptera of Australia. Part VI. - The Chironomidae. - Proc. Linn. Soc. N. S. W. 4: 215-311, pls. 11-14.

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SORIA, S. & P.G. BYSTRAK (1975): A new species of Forcipomyia (Diptera, Ceratopogonidae) described in all stages, with an account of its role as a cacao pollinator. - Rev. Theobroma (Brasil) 5: 3-11. SPINELLI, G.R. (1982): Cuatro especies nuevas del género Artichopogon [sic] KIEFER (Diptera: Ceratopogonidae) en la República Argentina. - Revta. Soc. Entomol. Argent. 41: 201-210. SPINELLI, G.R. (1983a): Notas sobre Ceratopogonidae de la República Argentina (Diptera, Nematocera) II. Nuevos aportes al conocimiento del género Forcipomyia MEIGEN. - Neotrópica 29: 121-129. SPINELLI, G.R. (1983b): Notas sobre Ceratopogonidae (Diptera, Nematocera) de la República Argentina I. Una nueva especie del género Alluaudomyia KIEFFER, redescripción de Dasyhelea penthesileae MACFIE, y nuevas citas para el género Stilobezzia KIEFFER. - Limnobios 2: 403-411. SPINELLI, G.R. (1983c): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina. III. Nuevos aportes al conocimiento de la tribu Palpomyiini. - Limnobios 2:17-24. SPINELLI, G.R. (1984): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina. IV. Descripción de adulto y pupa de Neobezzia termophila sp. nov. - Neotrópica 30: 197-200. SPINELLI, G.R. (1987): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina. VI. Las hembras de Paradasyhelea brevipalpis y de Macrurohelea paracaudata. - Limnobios 2: 667-670. SPINELLI, G.R. (1988): Two new species of Neotropical Alluaudomyia KIEFER, 1913 (Insecta, Diptera, Certopogonidae [sic]). - Iheringia, Ser. Zooll. 68: 129-136. SPINELLI, G.R. (1989): Amerohelea similis n. sp., from Argentina and Uruguay (Diptera: Ceratopogonidae). - Rev. Asoc. Cs. Nat. Litoral 20: 25-28. SPINELLI, G.R. (1990): The genus Brachypogon in Argentina, with a key to the Neotropical species (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 34: 743-755. SPINELLI, G.R. (1994): The male of Physohelea turgidipes (Diptera: Ceratopogonidae). - Neotrópica 40: 73-74. SPINELLI, G.R. (1996): A new species in the genus Diaphanobezzia from subantarctic Argentina (Diptera: Ceratopogonidae). - Neotrópica 42: 77-79. SPINELLI, G.R. (1997a): A new genus of the tribe Sphaeromiini from subantarctic Argentina and Chile related to Mackerrasomyia DEBENHAM (Diptera: Ceratopogonidae). - Mem. Entomol. Soc. Washington 18: 224-229. SPINELLI, G.R. (1997b): Further notes on Neotropical Ceratopogonini (Diptera: Ceratopogonidae). Revta. Soc. Entomol. Argent. 56:123-124. SPINELLI, G.R. (1998): Three new species and new records of Neotropical Stenoxenini (Diptera: Ceratopogonidae). - Neotrópica 44: 51-55. SPINELLI, G.R. & A. BORKENT (2004a): New species of Central American Culicoides LARTREILLE (Diptera: Ceratopogonidae) with a synopsis of species from Costa Rica. - Proc. Entomol. Soc. Washington 106: 361-395. SPINELLI, G.R. & A. BORKENT (2004b): A new species and first record of the subgenus Forcipomyia (Schizoforcipomyia) CHAN and LEROUX from the Neotropical Region (Diptera: Ceratopogonidae). - Zootaxa 572: 1-8. SPINELLI, G.R. & C.G. CAZORLA (2004): A new species of Brachypogon (Isohelea) from arid zones of Argentina (Diptera: Ceratopogonidae). - Zootaxa 570: 1-6. SPINELLI, G.R. & C.G. CAZORLA (2006): A new predaceous midge of the Palpomyia tibialis group from northeastern Argentina (Diptera: Ceratopogonidae). - Rev. Biol. Trop. 54: 1067-1070. SPINELLI, G.R. & A. DIPPOLITO (1995): Two new neotropical species of the Forcipomyia (F.) argenteola species group (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 54: 155-158.

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SPINELLI, G.R. & J.P. DURET (1993): Las especies neotropicales de Clinohelea (Diptera: Ceratopogonidae). - Graellsia 49: 39-50. SPINELLI, G.R. & M.L. FELIPPE-BAUER (1990a): Two new Neotropical species of predaceous midges of the tribe Sphaeromiini (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 85: 87-90. SPINELLI, G.R. & M.L. FELIPPE-BAUER (1990b): Sphaerohelea, a new Neotropical predaceous midge genus of the tribe Sphaeromiini (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 85: 195-198. SPINELLI, G.R. & W.L. GROGAN (1984): Three new species of Macrurohelea from Argentina with a key to the Neotropical species (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 86: 961-967. SPINELLI, G.R. & W.L. GROGAN (1985): Clastrieromyia, a new Neotropical genus of predaceous midges related to Palpomyia and Bezzia (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 87: 329-334. SPINELLI, G.R. & W.L. GROGAN (1986): Two new species of the predaceous midge genus Clastrieromyia from Uruguay with a new record of C. schnacki for Argentina (Diptera: Ceratopogonidae). Proc. Entomol. Soc. Washington 88: 455-460. SPINELLI, G.R. & W.L. GROGAN (1987): A revision of the Neotropical species of Parabezzia (Diptera: Ceratopogonidae). - Biol. Acuat. 11: 1-45. SPINELLI, G.R. & W.L. GROGAN (1989): Dos especies nuevas de Palpomyia, grupo distincta de la región Neotropical (Diptera: Ceratopogonidae). - Neotrópica 35: 3-8. SPINELLI, G.R. & W.L. GROGAN (1990): New species of predaceous midges of the tribe Ceratopogonini from subantarctic Argentina (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 92: 127-134. SPINELLI, G.R. & W.L. GROGAN (1993): Borkenthelea, a new predaceous midge genus from subantarctic Argentina (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 95: 321-326. SPINELLI, G.R. & W.L. GROGAN (1994): Two new Neotropical species of the subgenus Isohelea of Brachypogon, with the description of the female of B. (I.) misionensis and a key to the Neotropical species (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 53: 1-8. SPINELLI, G.R. & W.L. GROGAN (1998): A revision of the Neotropical predaceous midges of Brachypogon (Brachypogon) KIEFFER (Diptera: Ceratopogonidae). - Insecta Mundi 12: 59-79. SPINELLI, G.R. & W.L. GROGAN (1999): A new species of Macrurohelea INGRAM and MACFIE, and new records of biting midges of the tribes Culicoidini and Ceratopogonini (Diptera: Ceratopogonidae) from Tierra del Fuego and the Magallanes. - Proc. Entomol. Soc. Washington 101: 708-713. SPINELLI, G.R. & W.L. GROGAN (2001): A revision of the Patagonian predaceous midge genus Borkenthelea SPINELLI and GROGAN (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 103: 147-156. SPINELLI, G.R. & W.L. GROGAN (2003): A revision of the Neotropical biting midges of the genus Paradasyhelea INGRAM and MACFIE (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 105: 568-577. SPINELLI, G.R. & P.I. MARINO (1997): Two new species of the subgenus Thyridomyia of Forcipomyia from Argentina and new records of F. (Synthyridomyia) sanctaeclarae (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 123: 187-190. SPINELLI, G.R. & M.E. MARTINEZ (1992): The genus Culicoides in Uruguay (Diptera: Ceratopogonidae). - Insecta Mundi 5: 175-179 (1991).

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SPINELLI, G.R. & E.A. RODRÍGUEZ (1999): A new species of Dasyhelea from Argentina described as adult, pupae and fourth instar larvae (Diptera: Ceratopogonidae). - Neotrópica 45: 59-62. SPINELLI, G.R. & M.M. RONDEROS (1987): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina. V. Nuevos aportes al conocimiento del género Dasyhelea KIEFFER. Neotrópica 33: 11-17. SPINELLI, G.R. & M.M. RONDEROS (1991): Los polvorines del género Culicoides en area de influencia de la represa de Salto Grande (Diptera: Ceratopogonidae). - Neotrópica 37: 83-94. SPINELLI, G.R. & M.M. RONDEROS (2001): First record of the genus Bezzia in Chile, with a description of a new species of the venustula group (Diptera: Ceratopogonidae). - Rev. Chil. Hist. Nat. 74: 751-754. SPINELLI, G.R. & W.W. WIRTH (1981): A new species of predaceous midge of the genus Bezzia KIEFFER from Argentina (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 40: 187-192. SPINELLI, G.R. & W.W. WIRTH (1984a): Ocho especies nuevas del género Culicoides LATREILLE de la región Neotropical. Primera descripción del macho de C. flinti WIRTH, y de la hembra de C. lenti TAVARES y LUNA DIAS (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 43: 171-185. SPINELLI, G.R. & W.W. WIRTH (1984b): A review of the Neotropical predaceous midge genus Paryphoconus (Diptera: Ceratopogonidae). - Proc. Biol. Soc. Washington 97: 882-908. SPINELLI, G.R. & W.W. WIRTH (1984c): The Neotropical predaceous midges of the genus Alluaudomyia (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 86: 673-702. SPINELLI, G.R. & W.W. WIRTH (1984d): Las especies Neotropicales del género Dasyhelea, grupo cincta (Diptera: Ceratopogonidae). - Limnobios 2: 586-608. S PINELLI , G.R. & W.W. W IRTH (1986a): The Neotropical species of Phaenobezzia (Diptera: Ceratopogonidae). - Fla. Entomol. 69: 231-236. SPINELLI, G.R. & W.W. WIRTH (1986b): Clave para la identificación de las especies del género Culicoides LATREILLE presentes al sur de la cuenca Amazónica. Nuevas citas y notas sinonímicas (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 44: 49-73. SPINELLI, G.R. & W.W. WIRTH (1989a): Las especies Neotropicales del género Bezzia (Diptera: Ceratopogonidae). II. Los grupos nobilis y punctipennis. - Revta. Soc. Entomol. Argent. 45: 109-129. SPINELLI, G.R. & W.W. WIRTH (1989b): The Neotropical predaceous midges of the genus Bezzia (Diptera: Ceratopogonidae) Part I. The glabra and brevicornis groups. - Limnobios 2: 762-778. SPINELLI, G.R. & W.W. WIRTH (1990): Neotropical predaceous midges of the genus Bezzia (Diptera: Ceratopogonidae) Part III. The gibbera group of species. - Insecta Mundi 4: 11-32. SPINELLI, G.R. & W.W. WIRTH (1991): The Neotropical predaceous midges of the genus Bezzia (Diptera: Ceratopogonidae) Part IV. The dentifemur and venustula groups. - Insecta Mundi 5: 1-17. SPINELLI, G.R. & W.W. WIRTH (1993): Los Ceratopogonidae de la Argentina (Insect: Diptera). - In: CASTELLANOS, Z. (ed.). Fauna de agua dulce de la República Argentina. Volumen 38. Diptera Fascículo 3. Ceratopogonidae: 124 pp. SPINELLI, G.R., DIPPOLITO, A. & W.W. WIRTH (1995): A report on a collection of Ceratopogonidae (Diptera) from Rondonia, Brazil. 2. Tribes Heteromuiini [sic] and Sphaeromiini. - Insecta Mundi 9: 165-169. SPINELLI, G.R., GREINER, E.C. & W.W. WIRTH (1993): The Neotropical bloodsucking midges of the Culicoides guttatus group of the subgenus Hoffmania (Diptera: Ceratopogonidae). - Contrib. Am. Entomol. Inst. 27: 1-91. SPINELLI, G.R., MARINO, P.R. & P. POSADAS (2006): The Patagonian species of the genus Atrichopogon KIEFFER, with a biogeographic analysis based on Forcipomyiinae (Diptera: Ceratopogonidae). - Ins. Syst. and Evol. 37: 301-324.

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SPINELLI, G.R., MARINO, P.I. & M.M. RONDEROS (2005a): The fourth instar larva and pupa of the Neotropical biting midge Forcipomyia (Forcipomyia) rioplatensis MARINO and SPINELLI (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 107: 108-114. SPINELLI, G.R., RONDEROS, M.M. & E.G. BALSEIRO (1989): Análisis de la actividad diaria de dípteros nematocéros en Punta Lara (Pdo. de Ensenada, Prov. de Buenos Aires). II. Ceratopogonidae, con la descripción de una especie nueva en el género Atrichopogon. - Limnobios 2: 733-737. SPINELLI, G.R., RONDEROS, M.M., DÍAZ, F. & P.I. MARINO (2005b): The bloodsucking biting midges of Argentina (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 100: 137-150. SPINELLI, G.R., SCHAEFER, E.F. & A.I. KEHR (2002): First record of biting midges (Diptera: Ceratopogonidae) attacking frogs in the Neotropical Region. - Proc. Entomol. Soc. Washington 104: 527-528. STAEGER, R.C. (1839): Systematisk fortegnelse over de i Danmark hidtil fundne Diptera. - Naturhist. Tids. 2: 549-600. SZADZIEWSKI, R. (1996): Biting midges from Lower Cretaceous amber of Lebanon and Upper Cretaceous Siberian amber of Taimyr (Diptera, Ceratopogonidae). - Studia Dipterol. 3: 23-86. SZADZIEWSKI, R., KRZYWINSKI, J. & W. GILKA (1997): Diptera Ceratopogonidae, biting midges. In: NILSSON, A.N. (ed.). Aquatic insects of North Europe - A taxonomic handbook. Vol. 2: 243-263. TAVARES, O. & A.P. LUNA DIAS (1980): Quatro novas espécies do gênero Culicoides LATREILLE, 1809, do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 40: 393-399. TAVARES, O. & R.A. RUIZ (1980): Sobre duas espécies novas do gênero Culicoides LATREILLE, 1809, do Brasil (Diptera, Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 75: 27-32. TAVARES, O. & M.A. DE SOUZA (1978): Sobre três espécies novas do gênero Culicoides LATREILLE, 1809 do Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 38: 619-624. TAVARES, O. & M.A. DE SOUZA (1979): Sobre algumas espécies do gênero Culicoides LATREILLE, 1809 do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 39: 611-613. TAVARES, O. & M.A. DE SOUZA (1980): Duas espécies novas do gênero Monohelea KIEFFER, 1917, do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 40: 95-100. TAVARES, O. & A.J. SILVA PEREIRA (1978): Duas espécies novas do gênero Monohelea KIEFFER, 1917, do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 38: 157-160. THIENEMANN, A. (1928): Chironomiden - Metamorphosen I. - Arch. Hydrobiol. 19: 585-623. THOMSEN, L.C. (1937): Aquatic Diptera Part V. Ceratopogonidae. - Cornell Univ. Agric. Exp. Sta. Mem. 210: 57-80, pls. 10-18. TOKUNAGA, M. (1961): Notes on biting midges III. - Pub. Ent. Lab., Univ. Osaka Pref. 6: 115-122. TOWNSEND, C.H.T. (1893): An interesting blood-sucking gnat of the family Chironomidae. - Psyche 6: 369-371. UTMAR, J.A. & W.W. WIRTH (1976): A revision of the new world species of Forcipomyia, subgenus Caloforcipomyia (Diptera: Ceratopogonidae). - Fla. Entomol. 59: 109-133. VARGAS, L. (1953a): Beltranmyia n. subgen. de Culicoides (Insecta: Heleidae). - Rev. Inst. Sal. Enf. Trop. 13: 33-36. VARGAS, L. (1953b): Culicoides wirthomyia n. sp. y Culicoides stigmalis WIRTH, 1952 (Insecta, Diptera). Rev. Inst. Sal. Enf. Trop. 13: 227-233. VARGAS, L. (1954): Dos nuevas especies de Culicoides Mexicanos (Diptera, Heleidae). - Rev. Inst. Sal. Enf. Trop. 14: 25-32. VARGAS, L. (1960): The subgenera of Culicoides of the Americas (Diptera, Ceratopogonidae). - Rev. Biol. Trop. 8: 35-47.

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VARGAS, L. & W.W. WIRTH (1955): Culicoides blantoni n. sp. (Diptera, Heleidae). - Rev. Inst. Sal. Enf. Trop. 15: 33-35. VILLALOBOS, C. DE & M. RONDEROS (2003): Dasyhelea necrophila SPINELLI et RODRÍGUEZ, 1999 (Diptera: Ceratopogonidae) a new potential paratenic host of Paragordius varius (LEIDY, 1851) (Gordiida, Nematomorpha). - Acta Parasitol. 48: 218-221. VITALE, G.C., WIRTH, W.W. & T.H.G. AITKEN (1981): New species and records of Culicoides reared from arboreal habitats in Panama, with a synopsis of the debilipalpis group (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 83: 140-159. WALKER, F. (1856): Vol. 1. Diptera. - In: Insecta Saundersiana: or characters of undescribed insects in the collection of William Wilson SAUNDERS: 415-474, Esq., F.R.S., F.L.S. & C. Van Voorst, London: 474 pp. WALTL, J. (1837): Neue Gattungen von Mücken bei Passau. - Isis (Oken’s) 1837: 279-283. WARMKE, H.E. (1951): Studies on pollination of Hevea brasiliensis in Puerto Rico. - Science 113: 646-648. WARMKE, H.E. (1952): Studies on natural pollination of Hevea brasiliensis in Brazil. - Science 116: 474-475. WATTS, D., PHILLIPS, M.I., CALLAHAN, D.J., GRIEBENOW, W., HYAMS, K.C. & C.G. HAYESL (1997): Oropouche virus transmission in the Amazon river basin of Peru. - Am. J. Trop. Med. Hyg. 56: 148-152. WAUGH, W.T. & W.W. WIRTH (1976): A revision of the genus Dasyhelea KIEFFER of the eastern United States north of Florida (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 69: 219-247. WESENBERG-LUND, C. (1914): Bidrag til nogle myggeslaegters, saerlig Mochlonyx og Corethra biologi. Mindeskrift i Anledning of Hundredaaret for Japetus Steenstrups Fodsel, no. 34: 25 pp. WESTWOOD, J.O. (1840): Order XIII. Diptera Aristotle (Antliata FABRICIUS. Halteriptera Clairv.). An introduction to the modern classification of insects; founded on the natural habits and corresponding organisation of the different families: 125-154. - Synopsis of the genera of British insects, London: 158 pp. WEYENBERGH, H. (1883): Die Gattung Didymophleps m. - Stett. Entomol. Zeit. 1883: 108-111. WILLIAMS, F.X. (1944): Biological studies in Hawaiian water-loving insects. Part III. Diptera or flies. D. Culicidae, Chironomidae and Ceratopogonidae. - Proc. Hawaiian Entomol. Soc. 12: 149-180. WILLIAMS, R.W. (1956): The biting midges of the genus Culicoides found in the Bermuda Islands (Diptera, Heleidae) I. A description of C. bermudensis n. sp. with a key to the local fauna. - J. Parasitol. 42: 297-300. WILLISTON, S.W. (1896): On the Diptera of St. Vincent (West Indies). - Trans. Entomol. Soc. London 1896: 253-446, pls. 8-14. WILLISTON, S.W. (1900): Supplement (Part). - In: GODMAN, F.D. & O. SALVIN (eds.): Biologia CentraliAmericana. Zoologia-Insects-Diptera: 217-248, Vol. 1. London: 378 pp., 6 pls. WINDER, J.A. (1977a): Field observations on Ceratopogonidae and other Diptera: Nematocera associated with cocoa flowers in Brazil. - Bull. Entomol. Res. 67: 57-63. WINDER, J.A. (1977b): Biology, activity, and behavior of cocoa pollinators (abstract). - Mosquito News 37: 289. WINNERTZ, J. (1852): Beitrag zur Kenntniss der Gattung Ceratopogon MEIGEN. - Linn. Entomol. 6: 1-80. WIRTH, W.W. (1951a): A new biting midge of the genus Leptoconops from Florida, with new records of other American species (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 53: 281-284. WIRTH, W.W. (1951b): New species and records of Virginia Heleidae (Diptera). - Proc. Entomol. Soc. Washington 53: 313-326. WIRTH, W.W. (1952a): The Heleidae of California. - Univ. Calif. Pub. Entomol. 9: 95-266.

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WIRTH, W.W. (1952b): Los insectos de las Islas Juan Fernández. 7. Heleidae and Tendipedidae (Diptera). - Rev. Chil. Entomol. 2: 87-104. WIRTH, W.W. (1952c): Two new species of anthropophilic Culicoides from Guatemala (Diptera: Heleidae). - J. Parasitol. 38: 245-247. WIRTH, W.W. (1952d): The immature stages of two species of Florida salt marsh sand flies (Diptera, Heleidae). - Fla. Entomol. 35: 91-100. WIRTH, W.W. (1953a): Biting midges of the heleid genus Stilobezzia in North America. - Proc. U.S. Nat. Mus. 103: 57-85. WIRTH, W.W. (1953b): American biting midges of the heleid genus Monohelea. - Proc. U.S. Nat. Mus. 103: 135-154. WIRTH, W.W. (1955): Three new species of Culicoides from Texas (Diptera, Heleidae). - J. Washington Acad. Sci. 45: 355-359. WIRTH, W.W. (1956a): New species and records of biting midges ectoparasitic on insects (Diptera, Heleidae). - Ann. Entomol. Soc. Am. 49: 356-364. WIRTH, W.W. (1956b): The biting midges ectoparasitic on blister beetles (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 15-23. WIRTH, W.W. (1956c): The heleid midges involved in the pollination of rubber trees in America (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 241-250. WIRTH, W.W. (1959a): Pachyhelea, a new genus of American Ceratopogonidae related to Palpomyia (Diptera). - Bull. Brooklyn Entomol. Soc. 54: 50-52. WIRTH, W.W. (1959b): New species and records of Heleidae from Brazil (Diptera). - Deut. Entomol. Zeitsch. 6: 234-237. WIRTH, W.W. (1960): The genus Pellucidomyia MACFIE (Diptera, Ceratopogonidae). - Bull. Brooklyn Entomol. Soc. 55: 1-3. WIRTH, W.W. (1962): A reclassification of the Palpomyia-Bezzia-Macropeza groups, and a revision of the North American Sphaeromiini (Diptera, Ceratopogonidae). - Ann. Entomol. Soc. Am. 55: 272-287. WIRTH, W.W. (1965a): A new Johannsenomyia from Brazil (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 67: 4. WIRTH, W.W. (1965b): A revision of the genus Parabezzia MALLOCH (Diptera, Ceratopogonidae). Proc. Entomol. Soc. Washington 67: 215-230. WIRTH, W.W. (1965c): Two new species of Macrurohelea from Chile (Diptera, Ceratopogonidae). Pan-Pac. Entomol. 41: 46-50. WIRTH, W.W. (1966): A new Jamaican blood-sucking midge from lacewings (Diptera, Ceratopogonidae). - Proc. Entomol. Soc. Washington 68: 29-32. WIRTH, W.W. (1969): New species and records of Galapagos Diptera. - Proc. Calif. Acad. Sci. 36: 571-594. WIRTH, W.W. (1970): The neotropical Forcipomyia midges of the subgenus Thyridomyia SAUNDERS (Diptera, Ceratopogonidae). - Studia Entomol. 13: 429-440. WIRTH, W.W. (1971): A review of the “stick-ticks”, neotropical biting midges of the Forcipomyia subgenus Microhelea parasitic on walking stick insects (Diptera: Ceratopogonidae). - Entomol. News 82: 229-245. WIRTH, W.W. (1972): The neotropical Forcipomyia (Microhelea) species related to the caterpillar parasite F. fuliginosa (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 65: 564-577. WIRTH, W.W. (1974): Family Ceratopogonidae. - In: A catalog of the diptera of the Americas south of the United States, Fasc. 14: 89 pp.

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WIRTH, W.W. (1976): Forcipomyia pictoni MACFIE and descriptions of two new related species from Florida (Diptera: Ceratopogonidae). - Fla. Entomol. 59: 77-84. WIRTH, W.W. (1977): A review of the pathogens and parasites of the biting midges (Diptera: Ceratopogonidae). - J. Washington Acad. Sci. 67:60-75. WIRTH, W.W. (1978): New species and records of intertidal biting midges of the genus Dasyhelea KIEFFER from the Gulf of California (Diptera: Ceratopogonidae). - Pacific Insects 18: 191-198. WIRTH, W.W. (1982a): The cacao-pollinating midges of the Forcipomyia argenteola group (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 84: 568-585. WIRTH, W.W. (1982b): New species of neotropical Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol. 65: 248-253. WIRTH, W.W. (1983a): The North American predaceous midges of the Bezzia bicolor group (Diptera: Ceratopogonidae). - Fla. Entomol. 66: 292-310. WIRTH, W.W. (1983b): A review of the American predaceous midges of the Bezzia nobilis group (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 85:670-685. WIRTH, W.W. (1991a): The predaceous midge genus Allohelea KIEFFER in the western hemisphere (Diptera: Ceratopogonidae). - Fla. Entomol. 74: 491-505. WIRTH, W.W. (1991b): New and little-known species of Forcipomyia (Diptera: Ceratopogonidae) associated with cocoa pollination in Brazil. - Proc. Entomol. Soc. Washington 93: 163-175. WIRTH, W.W. (1991c): Forcipomyia bicolor and related species of the subgenus Lepidohelea in Brazil (Diptera: Ceratopogonidae). - Fla. Entomol. 74: 506-517. WIRTH, W.W. (1992): A new genus of Ceratopogonini (Diptera: Ceratopogonidae) from Brazil. Proc. Entomol. Soc. Washington 94: 276-281. WIRTH, W.W. (1994a): The subgenus Atrichopogon (Lophomyidium) with a revision of the Nearctic species (Diptera: Ceratopogonidae). - Insecta Mundi 8: 17-36. WIRTH, W.W. (1994b): The western hemisphere species of the predaceous midge genus Echinohelea, with descriptions of six new species (Diptera: Ceratopogonidae). - Insecta Mundi 8: 227-242. WIRTH, W.W. & W.R. ATCHLEY (1973): A review of the North American Leptoconops (Diptera: Ceratopogonidae). - Grad. Stud. Texas Tech Univ. 5: 1-57. WIRTH, W.W. & P. BARRETO (1978): New species of Culicoides biting midges (Diptera: Ceratopogonidae) from Colombia. - J. Med. Entomol. 14: 553-564. WIRTH, W.W. & F.S. BLANTON (1953a): Studies in Panama Culicoides (Diptera: Heleidae): I, Descriptions of six new species. - J. Washington Acad. Sci. 43: 69-77. WIRTH, W.W. & F.S. BLANTON (1953b): Studies in Panama Culicoides (Diptera, Heleidae). II. Descriptions of six additional new species. - J. Parasitol. 39: 229-236. WIRTH, W.W. & F.S. BLANTON (1953c): Studies in Panama Culicoides (Diptera, Heleidae). III. A new species related to phlebotomus (WILLISTON). - Entomol. News 64: 113-120. WIRTH, W.W. & F.S. BLANTON (1955a): Studies in Panama Culicoides (Diptera, Heleidae) IV. Descriptions of three new species. - Bull. Brooklyn Entomol. Soc. 50: 100-106 (October). W IRTH , W.W. & F.S. B LANTON (1955b): Studies in Panama Culicoides (Diptera, Heleidae) V. Descriptions of three new species of the subgenus Oecacta POEY. - Bull. Brooklyn Entomol. Soc. 50: 121-127. WIRTH, W.W. & F.S. BLANTON (1956a): A new species of salt-marsh sand fly from Florida, the Bahamas, Panama and Ecuador: its distribution and taxonomic differentiation from Culicoides furens (POEY) (Diptera, Heleidae). - Fla. Entomol. 39: 157-162.

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WIRTH, W.W. & F.S. BLANTON. (1956b): Studies in Panama Culicoides VII. The species of the pulicaris and cova-garciai group (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 211-227. WIRTH, W.W. & F.S. BLANTON (1956c): Studies in Panama Culicoides (Diptera, Heleidae). IX. Two new species related to leoni BARBOSA and reevesi WIRTH. - Bull. Brooklyn Entomol. Soc. 51: 45-52. WIRTH, W.W. & F.S. BLANTON (1959): Biting midges of the genus Culicoides from Panama (Diptera: Heleidae). - Proc. U.S. Nat. Mus. 109: 237-482. WIRTH, W.W. & F.S. BLANTON (1967): The North American Culicoides of the guttipennis group (Diptera: Ceratopogonidae). - Fla. Entomol. 50: 207-232. WIRTH, W.W. & F.S. BLANTON (1968a): A revision of the neotropical biting midges of the hylas group of Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol. 51: 201-215. WIRTH, W.W. & F.S. BLANTON (1968b): A new Culicoides from Guyana (Diptera: Ceratopogonidae). Fla. Entomol. 51: 251-252. WIRTH, W.W. & F.S. BLANTON (1970a): New genera of Neotropical Ceratopogonidae (Diptera). - Fla. Entomol. 53: 7-14. W IRTH , W.W. & F.S. B LANTON (1970b): New species of Neotropical Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol. 53: 39-45. WIRTH, W.W. & F.S. BLANTON (1970c): Notes on Brachypogon KIEFFER (Diptera, Ceratopogonidae), a new species, and two new Neotropical genera of the tribe Ceratopogonini. - Fla. Entomol. 53: 93-104. WIRTH, W.W. & F.S. BLANTON (1970d): A review of the Culicoides nigrigenus group, with two new species (Diptera: Ceratopogonidae). - Entomol. News 81: 141-151. WIRTH, W.W. & F.S. BLANTON (1971): New Neotropical sandflies of the Culicoides debilipalpis group (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 34-43. WIRTH, W.W. & F.S. BLANTON (1972a): Lanehelea, a new neotropical genus of Sphaeromiini (Diptera: Ceratopogonidae). - Studia Entomol. 15: 433-438. WIRTH, W.W. & F.S. BLANTON (1972b): A new Patagonian biting midge of the the genus Monohelea, Isthmohelea (Diptera: Ceratopogonidae). - Fla. Entomol. 55: 173-176. WIRTH, W.W. & F.S. BLANTON (1973): A review of the maruins or biting midges of the genus Culicoides (Diptera: Ceratopogonidae) in the Amazon Basin. - Amazoniana 4: 405-470. WIRTH, W.W. & F.S. BLANTON (1974a): A new Florida sand fly closely related to Culicoides haematopotus MALLOCH (Diptera: Ceratopogonidae). - Fla. Entomol. 57: 23-26. WIRTH, W.W. & F.S. BLANTON (1974b): The West Indian sandflies of the genus Culicoides (Diptera: Ceratopogonidae). - Agri. Res. Serv., U.S. Dept. Agric., Tech. Bull. 1474: iv + 98 pp. W IRTH , W.W. & F.S. B LANTON (1978): Two new species of neotropical Culicoides (Diptera: Ceratopogonidae). - Pan-Pac. Entomol. 54: 236-240. W IRTH , W.W. & J.L. C ASTNER (1990): New neotropical species of “stick-tick” (Diptera: Ceratopogonidae) from katydids. - Fla. Entomol. 73: 157-160. WIRTH, W.W. & M.I. DOW (1971): Studies on the genus Forcipomyia III. Blantonia, a new subgenus in the Trichohelea complex (Diptera: Ceratopogonidae). - Fla. Entomol. 54: 289-295. WIRTH, W.W. & M.I. DOW (1972): Studies on the genus Forcipomyia 4. Rhynchoforcipomyia, a new neotropical subgenus in the Trichohelea complex (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 65: 862-872. WIRTH, W.W. & A.L. DYCE (1985): The current taxonomic status of the Culicoides vectors of bluetongue viruses. - In: BARBER, T.L. & M.M. JOCHIM (eds.): 151-162. Bluetongue and related orboviruses. Alan R. Liss, New York: 746 pp.

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WIRTH, W.W. & M.L. FELIPPE-BAUER (1989): The neotropical biting midges related to Culicoides paraensis (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 84:551-565. WIRTH, W.W. & W.L. GROGAN (1977): Taxonomic notes on the genus Heteromyia SAY, and a new species from Nicaragua (Diptera: Ceratopogonidae). - Fla. Entomol. 60: 177-185. WIRTH, W.W. & W.L. GROGAN (1981): Natural History of Plummer’s Island, Maryland XXV. Biting midges (Diptera: Ceratopogonidae). 3. The species of the tribe Stilobezziini. - Bull. Biol. Soc. Washington 5: 1-102. WIRTH, W.W. & W.L. GROGAN (1982): The predaceous midges of the genus Phaenobezzia in North America (Diptera: Ceratopogonidae). - Mem. Entomol. Soc. Washington 10:179-192. WIRTH, W.W. & W.L. GROGAN (1983): The Nearctic species of the Bezzia bivittata group (Diptera: Ceratopogonidae). - Proc. Biol. Soc. Washington 96: 489-523. WIRTH, W.W. & W.L. GROGAN (1988): The predaceous midges of the world (Diptera: Ceratopogonidae; Tribe Ceratopogonini). - Flora and Fauna Handbook 4. E.J. Brill, Leiden: xv + 160 pp. WIRTH, W.W. & A.A. HUBERT (1960): Ceratopogonidae (Diptera) reared from cacti, with a review of the copiosus group of Culicoides. - Ann. Entomol. Soc. Am. 53: 639-658. WIRTH, W.W. & V.H. LEE (1967): New species of Culicoides from high altitudes in the Colombian Andes (Diptera: Ceratopogonidae). - Proc. U.S. Nat. Mus. 124: 1-22. WIRTH, W.W. & D.H. MESSERSMITH (1971): Studies on the genus Forcipomyia. 1. The North American parasitic midges of the subgenus Trichohelea (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 64: 15-26. WIRTH, W.W. & B.A. MULLENS (1992): Culicoides boydi (Diptera: Ceratopogonidae): a potential vector of hemorrhagic disease viruses to desert bighorn sheep in southern California. - J. Med. Entomol. 29: 1006-1010. WIRTH, W.W. & N.C. RATANAWORABHAN (1971a): Ceratoculicoides, a new genus related to Ceratopogon MEIGEN (Diptera, Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 170-177. WIRTH, W.W. & N.C. RATANAWORABHAN (1971b): Notes on Neotropical Pellucidomyia (Diptera, Ceratopogonidae). - Ann. Entomol. Soc. Am. 64: 446-448. WIRTH, W.W. & N.C. RATANAWORABHAN (1972a): Neobezzia, a new neotropical biting midge genus of the tribe Sphaeromiini (Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 45: 476-490. WIRTH, W.W. & N.C. RATANAWORABHAN (1972b): A revision of the tribe Stenoxenini (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 65: 1368-1388. WIRTH, W.W. & N.C. RATANAWORABHAN (1978): Studies on the genus Forcipomyia. V. Key to subgenera and description of a new subgenus related to Euprojoannisia BRÈTHES (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 80: 493-507. WIRTH, W.W. & S.J. SORIA (1975): A new Neotropical Forcipomyia midge closely related to F. (F.) genualis (LOEW) (Diptera: Ceratopogonidae). - Rev. Theobroma 5: 19-27. WIRTH, W.W. & S.J. SORIA (1980): Studies on the genus Forcipomyia VI. The Neotropical species of the subgenus Warmkea (Diptera: Ceratopogonidae). - Rev. Theobroma 9: 137-161. WIRTH, W.W. & S.J. SORIA (1981): Two Culicoides biting midges reared from inflorescences of Calathea in Brazil and Colombia, and a key to the species of the discrepans groups (Diptera: Ceratopogonidae). - Rev. Theobroma 11: 107-117. WIRTH, W.W. & G.R. SPINELLI (1992a): Immature stages of Forcipomyia seminole WIRTH and a related new neotropical species (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 349-356.

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WIRTH, W.W. & G.R. SPINELLI (1992b): American predaceous midges of the subgenus Eukraiohelea of Stilobezzia (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 342-349. WIRTH, W.W. & G.R. SPINELLI (1993a): The American species of the annulatipes group of the subgenus Lepidohelea, genus Forcipomyia (Diptera: Ceratopogonidae). - Insecta Mundi 6: 109-125 (1992). WIRTH, W.W. & G.R. SPINELLI (1993b): The North American species of the Forcipomyia (Lepidohelea) bicolor subgroup (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 95: 611-634. WIRTH, W.W. & W.T. WAUGH (1976): Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of Cocoa. - Studia Entomol. 19: 223-236. WIRTH, W.W. & R.W. WILLIAMS (1957): The biting midges of the Bermuda Islands, with descriptions of five new species. - Proc. Entomol. Soc. Washington 59: 5-14. WIRTH, W.W., DYCE, A.L. & G.R. SPINELLI (1988): An atlas of wing photographs, with a summary of the numerical characters of the Neotropical species of Culicoides (Diptera: Ceratopogonidae). Contrib. Am. Entomol. Inst. 25: 1-72. YOUNG, A.M. (1982): Effects of shade cover and availability of midge breeding sites on pollinating midge populations and fruit set in two cocoa farms. - J. Appl. Ecol. 19: 47-63. YOUNG, A.M. (1983): Seasonal differences in abundance and distribution of cocoa-pollinating midges in relation to flowering and fruit set between shaded and sunny habitats of the La Lola cocoa farm in Costa Rica. - J. Appl. Ecol. 20: 801-831. YOUNG, A.M. (1986a): Cocoa Pollination. - Cocoa Grower’s Bull. 37: 5-23. YOUNG, A.M. (1986b): Notes on the distribution and abundance of midges (Diptera: Ceratopogonidae and Cecidomyiidae) in some Central American cacao plantations. - Brenesia 24: 273-286.

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10 SUBJECT INDEX abercrombyi MACFIE (Forcipomyia) 52 acanthodes MACFIE (Bezzia) 31, 94 Acanthohelea KIEFFER 40, 85, 153 acidicola (TOKUNAGA) (Forcipomyia) 57, 125 acotylus LUTZ (Culicoides) 71 aculeata INGRAM & MACFIE (Palpomyia) 95 adamsoni MACFIE (Atrichopogon) 44 aductus DIPPOLITO & SPINELLI (Stenoxenus) 99 aegealitis SPINELLI & WIRTH (Dasyhelea) 59 aemulus MACFIE (Paryphoconus) 97 aenipes (MACFIE) (Mallochohelea) 91 aeria SAUNDERS (Forcipomyia) 26, 59 aeronautica MACFIE (Forcipomyia) 58 affinis FELIPPE-BAUER & SPINELLI (Monohelea) 83 aguirrei TAVARES & SOUZA (Monohelea) 83 aitkeni DE MEILLON & WIRTH (Forcipomyia) 55 aitkeni SPINELLI & WIRTH (Bezzia) 93 aitkeni WIRTH & BLANTON (Culicoides) 67 aitkeni WIRTH (Echinohelea) 81 alahialinus BARBOSA (Culicoides) 70 alambiculorum MACFIE (Culicoides) 65 albibasis (MALLOCH) (Mallochohelea) 91, 130 albicoxa LANE & FORATTINI (Stilobezzia) 86 albidiventris (KIEFFER) (Pachyhelea) 95, 152 albinensis INGRAM & MACFIE (Atrichopogon) 44 albitarsis WIRTH & RATANAWORABHAN (Neobezzia) 91 albocincta (Stilobezzia) 86 albopennis LANE (Clinohelea) 88 albopicta INGRAM & MACFIE (Dasyhelea) 59 albuquerquei LANE (Bezzia) 93 albuquerquei WIRTH & BLANTON (Culicoides) 70 alexanderi WIRTH (Parabezzia) 30, 84 *

Figures in bold

*

aliciae HUERTA & BORKENT (Ceratoculicoides) 80 alleni CLASTRIER & WIRTH (Forcipomyia) 54 Allohelea KIEFFER 32, 76, 101, 115-116, 139, 141, 148 Alluaudomyia KIEFFER 32, 76, 101, 115-116, 129, 140-141, 146, 148 almeidai (LANE) (Palpomyia) 95 almirantei WIRTH & BLANTON (Culicoides) 74 altivolans MACFIE (Atrichopogon) 44 alvarezi ORTÍZ (Culicoides) 75 amapaensis LANE (Paryphoconus) 97 amazonica CLASTRIER (Stilobezzia) 85 amazonica SPINELLI & WIRTH (Alluaudomyia) 76 amazonica WIRTH & BLANTON (Fittkauhelea) 81, 104, 150 amazonica WIRTH (Forcipomyia) 54 amazonius MACFIE (Culicoides) 69 ameliae BROWNE (Culicoides) 62 americana KIEFFER (Stilobezzia) 86 Amerohelea GROGAN & WIRTH 32, 41, 92, 102, 114, 122, 128, 143, 145, 148 amnicola (MACFIE) (Neobezzia) 30, 91 amnigena (MACFIE) (Stilobezzia) 86 Amossovia GLUKHOVA 62 ancora (COQUILLETT) (Dasyhelea) 59 andensis INGRAM & MACFIE (Dasyhelea) 59 andicola WIRTH & LEE (Culicoides) 63 andinus WIRTH & LEE (Culicoides) 73 angustipennis ENDERLEIN (Paryphoconus) 31, 97, 113, 127, 131-132 Anilomyia VARGAS 40, 62 anitae HUERTA & IBAÑEZ-BERNAL (Forcipomyia) 52

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annettae SPINELLI & BORKENT (Culicoides) 27, 67 annulatipes MACFIE (Forcipomyia) 52 annuliductus WIRTH (Culicoides) 27, 66 anomalicornis KIEFFER (Paryphoconus) 97 antefurcatus WIRTH & BLANTON (Culicoides) 71 antennalis (COQUILLETT) (Stilobezzia) 30, 86, 119 antequerae (LYNCH ARRIBÁLZAGA) (Heteromyia) 89 antiguensis SAUNDERS (Forcipomyia) 26, 56 antioquiae CLASTRIER & WIRTH (Forcipomyia) 54 antunesi FORATTINI (Culicoides) 73 Apelma KIEFFER 22-27, 56, 58 apicalis SPINELLI & WIRTH (Paryphoconus) 97 apricans (INGRAM & MACFIE) (Austrosphaeromias) 90 apunctipennis SPINELLI & GROGAN (Brachypogon) 78 aragaoi TAVARES & LUNA DIAS (Culicoides) 66 araucana SPINELLI & WIRTH (Bezzia) 93 araucaria SPINELLI (Diaphanobezzia) 80, 114 archboldi DE MEILLON & WIRTH (Forcipomyia) 55 archboldi WIRTH & BLANTON (Culicoides) 72 archboldi WIRTH (Atrichopogon) 47 archibaldoi TAVARES & SOUZA (Monohelea) 83 arcuatus WIRTH & RATANAWORABHAN (Stenoxenus) 99 arenosa CLASTRIER & RACCURT (Parabezzia) 84 argentata (LOEW) (Johannsenomyia) 90, 121, 127 argenteola MACFIE (Forcipomyia) 25, 49 argentina LANE & DURET (Clinohelea) 88 argentinensis KIEFFER (Dasyhelea) 59 aridus SPINELLI & MARINO (Atrichopogon) 44 arubae FOX & HOFFMAN (Culicoides) 17, 27, 75 assimilis INGRAM & MACFIE (Atrichopogon) 46 asuturus BORKENT & PICADO (Atrichopogon) 44 ateles (MACFIE) (Phaenobezzia) 97 atelis WIRTH (Culicoides) 74 atlantica WIRTH & WILLIAMS (Bezzia) 94 atlantis WIRTH & WILLIAMS (Dasyhelea) 27, 59 Atrichopogon KIEFFER 15, 18, 20, 22, 24, 32, 44, 54, 101, 103, 105, 109, 111, 129, 134, 136, 147-148, 150 atrichopogon LANE & FORATTINI (Stilobezzia) 85 atripalpis WIRTH & BLANTON (Culicoides) 71 attenuata SAUNDERS (Forcipomyia) 26, 52 aureus ORTÍZ (Culicoides) 74 auricoma KIEFFER (Atrichopogon) 44 australis WIRTH (Dasyhelea) 59

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Austrohelea WIRTH & GROGAN 32, 40, 77, 101, 116-117, 140, 142, 148 austroparaensis SPINELLI (Culicoides) 66 Austrosphaeromias SPINELLI 40, 90, 102, 122-123, 143-144, 149 Avaritia FOX 63 avilaensis ORTÍZ & MIRSA (Culicoides) 70 azureus WIRTH & BLANTON (Culicoides) 70 bachmanni SPINELLI (Culicoides) 66 Baeodasymyia CLASTRIER & RACCURT 32, 40-41, 77, 101, 107, 138, 149 Baeohelea WIRTH & BLANTON 40-41, 77, 101, 107, 138, 149 bahamensis (JOHNSON) (Dasyhelea) 27, 59 Bahiahelea WIRTH 40, 78, 101, 110, 115, 138-139 bahiensis BARBOSA (Culicoides) 64 bahiensis WIRTH & SPINELLI (Forcipomyia) 52 bajensis WIRTH (Dasyhelea) 59 balboa (LANE & WIRTH) (Downeshelea) 80 balsapambensis ORTÍZ & LEÓN (Culicoides) 72 balseiroi SPINELLI & GROGAN (Parabezzia) 30, 84 balseiroi SPINELLI (Atrichopogon) 44 bambusicola LUTZ (Culicoides) 27, 64 banksi (GERRY) (Nilobezzia) 92 barbatus BORKENT & PICADO (Atrichopogon) 44 barbosai WIRTH & BLANTON (Culicoides) 17, 27, 70 barrettoi LANE & DURET (Clinohelea) 88 barrettoi LANE (Palpomyia) 95 barrettoi LANE (Paryphoconus) 97 barthi TAVARES & SOUZA (Culicoides) 70 basifemoralis WIRTH & SPINELLI (Forcipomyia) 52 bassoi RONDEROS & SPINELLI (Leptoconops) 43 batesi LANE (Paryphoconus) 98 batesi WIRTH & BLANTON (Culicoides) 67 baueri WIRTH (Forcipomyia) 58 bayano WIRTH (Culicoides) 27, 66 beaveri WIRTH & BARRETO (Culicoides) 71 beccus BORKENT & PICADO (Atrichopogon) 45 beckae WIRTH (Stilobezzia) 86 beebei FOX (Culicoides) 62 bejaranoi DURET & LANE (Heteromyia) 89 belemensis CLASTRIER & WIRTH (Forcipomyia) 54 belemensis WIRTH & BLANTON (Culicoides) 71 bella (COQUILLETT) (Alluaudomyia) 29, 76, 112, 118

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Beltranmyia VARGAS 63 benarrochi ORTÍZ & MIRSA (Culicoides) 73 bequaerti (KIEFFER) (Leptoconops) 17, 25, 33, 43, 109, 130, 151 bermudae WIRTH & WILLIAMS (Dasyhelea) 60 bermudensis WILLIAMS (Culicoides) 27, 63 Bezzia KIEFFER 32, 86, 91-93, 97-98, 102, 122123, 128, 133, 143-145, 149 bicellii (LANE) (Mallochohelea) 91 bicinctipes INGRAM & MACFIE (Stilobezzia) 85 bicolor LANE (Stilobezzia) 30, 86 bicolor LUTZ (Forcipomyia) 52, 59 bicolor SAUNDERS (Forcipomyia) 26, 58-59 bicornis FELIPPE-BAUER & QUINTELAS (Downeshelea) 80 bicuspis BORKENT & PICADO (Atrichopogon) 45 bidentata FELIPPE-BAUER & SPINELLI (Monohelea) 83 biestroi SPINELLI & FELIPPE-BAUER (Sphaerohelea) 92, 152 biestroi SPINELLI & RONDEROS (Culicoides) 67 biestroi SPINELLI (Alluaudomyia) 29, 76 bifida WIRTH & SPINELLI (Forcipomyia) 52 bifidus EWEN (Atrichopogon) 25, 45 bifidus SPINELLI & GROGAN (Brachypogon) 78 bimaculata (LOEW) (Clinohelea) 113, 121 bimaculata LANE & FORATTINI (Stilobezzia) 86 bimaculatus FLOCH & ABONNENC (Culicoides) 68 bimaculatus SPINELLI & GROGAN (Brachypogon) 78 birabeni CAVALIERI (Culicoides) 70 bispinosa KIEFFER (Stilobezzia) 86 bivittata (COQUILLETT) (Bezzia) 30, 93 blantoni (LANE & WIRTH ) (Johannsenomyia) 90 blantoni (LANE & WIRTH) (Downeshelea) 80 blantoni (LANE) (Pellucidomyia) 89 blantoni LANE & FORATTINI (Stilobezzia) 86 blantoni SORIA & BYSTRAK (Forcipomyia) 25, 49 blantoni SPINELLI & WIRTH (Bezzia) 30, 93 blantoni VARGAS & WIRTH (Culicoides) 64 blantoni WIRTH & RATANAWORABHAN (Neobezzia) 91, 114 blantoni WIRTH & RATANAWORABHAN (Stenoxenus) 99 blantoni WIRTH (Echinohelea) 81 blantoni WIRTH (Parabezzia) 84 Blantonia WIRTH & DOW 48 boliviensis (KIEFFER) (Palpomyia) 96

boliviensis KIEFFER (Palpomyia) 95-96 boliviensis SPINELLI & WIRTH (Culicoides) 73 bonaerensis SPINELLI (Brachypogon) 78 borgmeieri WIRTH & WAUGH (Dasyhelea) 60 borinqueni FOX & HOFFMAN (Culicoides) 28, 65 Borkenthelea SPINELLI & GROGAN 40, 78, 101, 116117, 140, 142, 149 borkenti SPINELLI & CAZORLA (Brachypogon) 79 bourioni (CLASTRIER) (Nannohelea) 83, 108 Brachyconops WIRTH & ATCHLEY 43 Brachypogon KIEFFER 32, 78, 101, 107, 115-116, 138-139, 141, 149, 152 brachyrhyncha WIRTH & DOW (Forcipomyia) 56 brasiliae (LANE) (Neobezzia) 91 brasiliana CLASTRIER & WIRTH (Forcipomyia) 54 brasilianum FORATTINI (Culicoides) 68 brasiliensis (LANE) (Nilobezzia) 92 brasiliensis (LUTZ) (Leptoconops) 43-44 brasiliensis LANE (Monohelea) 83 brasiliensis MACFIE (Atrichopogon) 45 brasiliensis MACFIE (Forcipomyia) 26, 52 brasiliensis MACFIE (Palpomyia) 95 brasiliensis MACFIE (Stenoxenus) 99 brasiliensis SPINELLI & GROGAN (Parabezzia) 84 brasiliensis WIRTH (Bahiahelea) 78, 113, 125 bredini WIRTH & BLANTON (Culicoides) 65 breelandi CLASTRIER & WIRTH (Forcipomyia) 54 brevicornis (KIEFFER) (Bezzia) 30, 93 brevilabellata CLASTRIER & WIRTH (Forcipomyia) 54 brevipalpis (INGRAM & MACFIE) (Paradasyhelea) 75 brevipalpis MACFIE (Atrichopogon) 45 bricenoi ORTÍZ (Culicoides) 70 broadheadi CLASTRIER & WIRTH (Forcipomyia) 54 bromeliae SAUNDERS (Forcipomyia) 25, 49 bromeliae SPINELLI & WIRTH (Bezzia) 30, 93 bromelicola (LUTZ) (Forcipomyia) 26, 55-56 brownei SPINELLI (Culicoides) 68 brunnea WIRTH (Parabezzia) 84 brunneipennis SPINELLI & WIRTH (Paryphoconus) 98 bystraki GROGAN & WIRTH (Parabezzia) 109 Cacaohelea WIRTH & GROGAN 40-41, 79, 101, 115, 117, 126, 140-141, 149 cacaoi WIRTH & WAUGH (Dasyhelea) 60 cacaophila RONDEROS & SPINELLI (Forcipomyia) 52

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

cacozelus MACFIE (Culicoides) 65 caerulea SAUNDERS (Forcipomyia) 25, 48 calatheae WIRTH (Forcipomyia) 49 calcarata (COQUILLETT) (Forcipomyia) 25, 49 calchaqui SPINELLI & MARINO (Forcipomyia) 57 calchaqui SPINELLI (Brachypogon) 78 caldasi BROWNE (Culicoides) 72 caliginosa INGRAM & MACFIE (Forcipomyia) 50 caliginosella WIRTH (Forcipomyia) 50 callangana KIEFFER (Palpomyia) 95 Caloforcipomyia SAUNDERS 48 caloptera KIEFFER (Heteromyia) 89 calvescens MACFIE (Dasyhelea) 27, 60 camposi ORTÍZ & LEÓN (Culicoides) 71 cancer HOGUE & WIRTH (Culicoides) 28, 70 capitata WIRTH & GROGAN (Bezzia) 93 caprilesi FOX (Culicoides) 74 caribbea WIRTH & DOW (Forcipomyia) 48 caribbeana CLASTRIER & RACCURT (Parabezzia) 84 caribbeana SAUNDERS (Forcipomyia) 26, 56 caribbeana SPINELLI & WIRTH (Alluaudomyia) 29, 76 caribbeana SPINELLI & WIRTH (Dasyhelea) 60 caribbeanus EWEN (Atrichopogon) 25, 45 caribe LANE & FORATTINI (Stilobezzia) 86 caridei (BRÈTHES) (Culicoides) 17, 64 carioca (TAVARES & SILVA PEREIRA) (Downeshelea) 80 carioca LANE (Bezzia) 93 carioca LANE (Palpomyia) 95 carnatus BORKENT & PICADO (Atrichopogon) 45 carpenteri WIRTH & BLANTON (Culicoides) 71 carpinteroi MARINO & SPINELLI (Atrichopogon) 45 carrerai (LANE) (Palpomyia) 95 carrerai LANE (Stenoxenus) 99 carsiomelas WIRTH & BLANTON (Culicoides) 71 carvalhoi WIRTH & BLANTON (Culicoides) 73 casali CAVALIERI & CHIOSSONE (Atrichopogon) 45 casali CAVALIERI & CHIOSSONE (Leptoconops) 44 castanea LANE (Heteromyia) 89 castanea MACFIE (Palpomyia) 95 castillae FOX (Culicoides) 72 castneri CLASTRIER & WIRTH (Forcipomyia) 54 castroi (TAVARES & SILVA PEREIRA) (Downeshelea) 80 catarina CLASTRIER & WIRTH (Forcipomyia) 54 catarinensis LANE (Palpomyia) 95

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catarinensis MARINO & SPINELLI (Forcipomyia) 51 catarinensis SPINELLI & WIRTH (Alluaudomyia) 76 catarinensis SPINELLI & WIRTH (Bezzia) 93 caucaensis WIRTH & LEE (Culicoides) 72 caudata INGRAM & MACFIE (Macrurohelea) 82 caudispina (Dipetalonema) 33 cayoensis SPINELLI & GROGAN (Parabezzia) 84 cayoensis SPINELLI & WIRTH (Bezzia) 93 cebacoi (LANE & WIRTH) (Downeshelea) 80 Centrorhynchus LUTZ 51-52, 61, 64 Ceratobezzia KIEFFER 41, 88, 98 Ceratoculicoides WIRTH & RATANAWORABHAN 41, 101, 107, 109, 138, 149 Ceratopogon MEIGEN 27, 44, 46-47, 49-51, 54, 56, 59-62, 67-70, 75-76, 79, 83, 86-87, 9298, 118 cerifera SAUNDERS (Forcipomyia) 26, 53 cervicalis (Onchocerca) 33 chacoensis SPINELLI & WIRTH (Culicoides) 65 chaconi MACFIE (Stilobezzia) 30, 86 chaquensis DURET & LANE (Heteromyia) 89 charrua FELIPPE-BAUER & SPINELLI (Downeshelea) 80 charruus SPINELLI & Martinez (Culicoides) 28, 68 chaverrii SPINELLI & BORKENT (Culicoides) 28, 62 chiapasi (LANE & WIRTH) (Downeshelea) 80 chilensis (PHILIPPI) (Forcipomyia) 51 chilensis FORATTINI (Leptoconops) 44, 151 chilensis GROGAN & WIRTH (Notiohelea) 83 chilensis INGRAM & MACFIE (Atrichopogon) 46 chilensis INGRAM & MACFIE (Dasyhelea) 60 chilensis INGRAM & MACFIE (Palpomyia) 96 chilensis SPINELLI & RONDEROS (Bezzia) 93 chirusi (LANE & WIRTH) (Downeshelea) 80 christopheri BORKENT (Baeodasymyia) 29, 77 chrysonotus WIRTH & BLANTON (Culicoides) 62 cincta (COQUILLETT) (Dasyhelea) 27, 59, 60 claripennis LYNCH ARRIBÁLZAGA (Ceratopogon) 99 clastrieri DESSART (Forcipomyia) 55 clastrieri GROGAN & WIRTH (Nannohelea) 83, 151 clastrieri SPINELLI & GROGAN (Parabezzia) 84 Clastrieromyia SPINELLI & GROGAN 32, 41, 95, 102, 122, 128, 143, 145, 149 clavata WILLISTON (Heteromyia) 30, 89 clavipennis SPINELLI & WIRTH (Bezzia) 93

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clavipes WIRTH & RATANAWORABHAN (Neobezzia) 91 Clinohelea KIEFFER 32, 41, 88, 101, 117, 119, 142, 149 coheni CLASTRIER & WIRTH (Forcipomyia) 54 colombiae (LANE & WIRTH) (Downeshelea) 80 colombiae WIRTH (Forcipomyia) 58 colombiana CLASTRIER & WIRTH (Forcipomyia) 54 colossus BORKENT & PICADO (Atrichopogon) 45 columbiana (KIEFFER) (Dasyhelea) 60 columbiana KIEFFER (Palpomyia) 96 columbianus KIEFFER (Atrichopogon) 45 comechingon SPINELLI & MARINO (Atrichopogon) 45 commatis WIRTH & BLANTON (Culicoides) 71 concinna (MEIGEN) (Probezzia) 131-132 concoloripes MACFIE (Bezzia) 94 conifera MACFIE (Palpomyia) 96 contubernalis ORTÍZ & LEÓN (Culicoides) 69 convexipenis WIRTH & SPINELLI (Forcipomyia) 52 coomani SÉGUY (Stenoxenus) 113-114, 121, 125 copanensis UTMAR & WIRTH (Forcipomyia) 48 copiosus ROOT & HOFFMAN (Culicoides) 65, 111 coquilletti KIEFFER (Stilobezzia) 30, 85-86 coracina KIEFFER (Stilobezzia) 87 cornuta SAUNDERS (Forcipomyia) 26, 52 coroicoensis WIRTH (Palpomyia) 96 correntina DURET & LANE (Heteromyia) 89 correntina RONDEROS & DÍAZ (Dasyhelea) 27, 60 costalis MACFIE (Atrichopogon) 45 costalis WIRTH (Parabezzia) 84 costaricae MACFIE (Atrichopogon) 45 costaricae WIRTH & RATANAWORABHAN (Neobezzia) 91 Cotocripus BRÈTHES 40, 64 coutinhoi BARRETTO (Culicoides) 68 covagarciai ORTÍZ (Culicoides) 62 crassicrus KIEFFER (Palpomyia) 96 crepuscularis MALLOCH (Culicoides) 28, 63 crescentis WIRTH & BLANTON (Culicoides) 71 cristata CLASTRIER & WIRTH (Forcipomyia) 54 crucifer CLASTRIER (Culicoides) 66 crudelis (MAYER) (Forcipomyia) 53-54 crudelis KNAB (Forcipomyia) 54 cryptogamus MACFIE (Atrichopogon) 45 cuacuahuitlus HUERTA & BORKENT (Brachypogon) 79 cuiabai WIRTH (Culicoides) 70

Culicoides LATREILLE 17-18, 22, 24, 32-34, 40, 4243, 60-65, 67, 69, 100-101, 103, 105, 133134, 137, 147, 149, 150 cummingi SPINELLI & BORKENT (Culicoides) 71 cunasi LANE & WIRTH (Monohelea) 83 cylindrica WIRTH & DOW (Forcipomyia) 56 cylindricornis WIRTH & BLANTON (Culicoides) 74 daedaloides WIRTH & BLANTON (Culicoides) 71 daedalus MACFIE (Culicoides) 71 dalcyi GROGAN & WIRTH (Amerohelea) 92 dalessandroi WIRTH & BARRETO (Culicoides) 70 damascenoi LANE & DURET (Clinohelea) 88 danaisi (FLOCH & ABONNENC) (Forcipomyia) 58 darlingtonae WIRTH & BLANTON (Culicoides) 66 darwini MARINO & SPINELLI (Forcipomyia) 53 Dasyhelea KIEFFER 15, 22, 24, 32, 40, 59, 75, 101, 103, 105, 129, 133-134, 137, 146-147, 150 dasyophrus MACFIE (Culicoides) 72 davidi SPINELLI (Culicoides) 68 daviesi WIRTH & BLANTON (Culicoides) 70 deanei FELIPPE-BAUER & QUINTELAS (Downeshelea) 80 deanei FELIPPE-BAUER & WIRTH (Culicoides) 75 debilipalpis LUTZ (Culicoides) 17, 28, 66 decor (WILLISTON) (Culicoides) 62 delpontei CAVALIERI & CHIOSSONE (Atrichopogon) 45 delpontei CAVALIERI (Forcipomyia) 50 denisae CLASTRIER (Culicoides) 66 dentifemur SPINELLI & WIRTH (Bezzia) 93 depilis MACFIE (Atrichopogon) 45 desutterae CLASTRIER & WIRTH (Forcipomyia) 54 diabolicus HOFFMAN (Culicoides) 68 Diaphanobezzia INGRAM & MACFIE 40, 80, 101, 110, 139, 150 Dicrohelea KIEFFER 90 dicrourus WIRTH & BLANTON (Culicoides) 28, 70 Didymophleps WEYENBERGH 99, 102 didymothecae MACFIE (Atrichopogon) 45 diffusus SPINELLI (Culicoides) 68 diminuta LANE & FORATTINI (Stilobezzia) 87 diminutus BARBOSA (Culicoides) 69 dimorphus KIEFFER (Stenoxenus) 99 Diphaomyia VARGAS 64 discoloripes MACFIE (Forcipomyia) 52 discrepans ORTÍZ & MIRSA (Culicoides) 70

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disjuncta INGRAM & MACFIE (Isthmohelea) 82, 126, 151 distispinulosa SPINELLI & WIRTH (Alluaudomyia) 29, 76 diversus FELIPPE-BAUER (Culicoides) 66 dominicana DE MEILLON & WIRTH (Forcipomyia) 26, 56 dominicanus WIRTH & BLANTON (Culicoides) 62 dominicii ORTÍZ (Culicoides) 71 domizii SPINELLI (Atrichopogon) 45 donajii VARGAS (Culicoides) 72 donskoffi CLASTRIER & WIRTH (Forcipomyia) 54 dorsalis WIRTH & DOW (Forcipomyia) 56 dorsofasciata (LUTZ) (Stilobezzia) 30, 86 dowi BYSTRAK & WIRTH (Forcipomyia) 49 Downeshelea WIRTH & GROGAN 80, 101, 110, 113, 115-116, 118, 126, 138, 140-141, 150 dryadum MACFIE (Stilobezzia) 87 Drymodesmyia VARGAS 65 duartei TAVARES & LUNA DIAS (Culicoides) 73 dubia MACFIE (Forcipomyia) 52 dubitans LANE (Stilobezzia) 30, 87 dunklei CLASTRIER & WIRTH (Forcipomyia) 54 dunni WIRTH & BLANTON (Culicoides) 71 dureti LANE & FORATTINI (Stilobezzia) 87 dureti RONDEROS & SPINELLI (Culicoides) 66 dycei SPINELLI & GROGAN (Clastrieromyia) 31, 95, 127, 131 eadsi WIRTH & BLANTON (Culicoides) 66 echinodes MACFIE (Atrichopogon) 45 Echinohelea MACFIE 32, 40, 81, 101, 104, 122, 127, 131, 142, 150 Echinoideshelea WIRTH 81 ecuadorensis SPINELLI & GROGAN (Brachypogon) 78 ecuadorensis SPINELLI & WIRTH (Paryphoconus) 98 edeni WIRTH & BLANTON (Culicoides) 28, 64 edmistoni WIRTH & SPINELLI (Forcipomyia) 26, 52 edwardsi (SAUNDERS) (Forcipomyia) 26, 50, 56 edwardsi INGRAM & MACFIE (Forcipomyia) 50 edwardsi INGRAM & MACFIE (Stilobezzia) 85 edwardsiana WIRTH (Forcipomyia) 50 efferus FOX (Culicoides) 62 eldridgei WIRTH & BARRETO (Culicoides) 66 elegantula (JOHANNSEN) (Stilobezzia) 86 elutus MACFIE (Culicoides) 64 endemicus SPINELLI & MARINO (Atrichopogon) 45 enderleini LANE (Paryphoconus) 98

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equatoriensis BARBOSA (Culicoides) 66 eriophora (WILLISTON) (Forcipomyia) 54 esmeralda LANE & FORATTINI (Stilobezzia) 87 espinolai F ELIPPE -B AUER & L OURENCO - DE OLIVEIRA (Culicoides) 66 esteparia MARINO & SPINELLI (Forcipomyia) 49 estevezae RONDEROS & SPINELLI (Culicoides) 72 estevezae SPINELLI & WIRTH (Alluaudomyia) 76 ethelae SPINELLI & GROGAN (Brachypogon) 78 eublepharus MACFIE (Culicoides) 72 eucnemus MACFIE (Atrichopogon) 45 eukosma MACFIE (Forcipomyia) 48 Eukraiohelea INGRAM & MACFIE 86 Euprojoannisia BRÈTHES 49 euthystyla WIRTH & SPINELLI (Forcipomyia) 52 evansi WIRTH & BLANTON (Culicoides) 64 excentricus LANE (Stenoxenus) 99 fairchildi LANE & WIRTH (Monohelea) 83, 110, 138 fairchildi WIRTH & BLANTON (Culicoides) 71 fairfaxensis WIRTH (Forcipomyia) 111 falcifera SAUNDERS (Forcipomyia) 25, 49 fallax KIEFFER (Clinohelea) 88 farri WIRTH & BLANTON (Culicoides) 62 farri WIRTH (Forcipomyia) 56 fasciata GROGAN & WIRTH (Amerohelea) 92 fasciata SAY (Heteromyia) 89, 121 fasciatus (WALKER) (Heteromyia) 89 felippebauerae CLASTRIER & WIRTH (Forcipomyia) 54 femoralis LANE & FORATTINI (Stilobezzia) 87 fernandezi ORTÍZ (Culicoides) 72 fernandoi TAVARES & SOUZA (Culicoides) 68 ferreyrai RONDEROS & SPINELLI (Culicoides) 68 fiebrigi KIEFFER (Atrichopogon) 45 fiebrigi KIEFFER (Stilobezzia) 30, 87 fieldi WIRTH & BLANTON (Culicoides) 73 filarifer HOFFMAN (Culicoides) 22, 33, 68 filibranchia (LUTZ) (Dasyhelea) 27, 60 filiducta SPINELLI & WIRTH (Bezzia) 93 filiductus WIRTH (Culicoides) 66, 93 fimbriatus MACFIE (Atrichopogon) 45 Fittkauhelea WIRTH & BLANTON 41, 81, 101, 110, 115, 139-140, 150 fittkaui SPINELLI & WIRTH (Alluaudomyia) 76 fittkaui SPINELLI & WIRTH (Paryphoconus) 98

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fittkaui WIRTH & BLANTON (Culicoides) 74 fittkaui WIRTH & RATANAWORABHAN (Neobezzia) 91 flava (WILLISTON) (Forcipomyia) 51 flavicauda MACFIE (Dasyhelea) 27, 60 flavicaudae MACFIE (Atrichopogon) 45 flavidus (JOHANNSEN) (Paryphoconus) 31, 98 flavifemoris MACFIE (Forcipomyia) 53 flavifrons (GUÉRIN) (Dasyhelea) 132 flavipes LUTZ (Atrichopogon) 45, 95 flavivenulus COSTA LIMA (Culicoides) 68 flinti SPINELLI & WIRTH (Bezzia) 93 flinti SPINELLI & WIRTH (Paryphoconus) 98 flinti WIRTH (Culicoides) 66 flochabonnenci ORTÍZ & MIRSA (Culicoides) 72 florenciae MESSERSMITH (Culicoides) 72 floridensis BECK (Culicoides) 28, 75 floridensis DOW & WIRTH (Forcipomyia) 57 floridensis WIRTH (Leptoconops) 44, 104 fluminensis FELIPPE-BAUER & QUINTELAS (Downeshelea) 80 fluminensis LANE (Bezzia) 93 fluminensis LANE (Stilobezzia) 87 flumineus MACFIE (Atrichopogon) 45 fluvialis MACFIE (Culicoides) 72 fluviatilis (LUTZ) (Culicoides) 75 forattinii ORTÍZ (Culicoides) 74 forattinii WIRTH & GROGAN (Schizonyxhelea) 30, 85, 112 Forcipomyia MEIGEN 15, 17, 18, 19-21, 22, 24, 3233, 40, 47-49, 53, 55-58, 101-103, 105, 118, 133, 134, 136, 147-148, 150 formosa (LOEW) (Nilobezzia) 135 foxi ORTÍZ (Culicoides) 17, 68 franklini CLASTRIER & WIRTH (Forcipomyia) 54 franklini SPINELLI (Culicoides) 68 freitasi WIRTH & BLANTON (Culicoides) 64 frontispina (DOW & TURNER) (Amerohelea) 92, 127 fuliginosa (MEIGEN) (Forcipomyia) 26, 53-54 fulvus JOHANNSEN (Stenoxenus) 99 fur (JOHNSON) (Forcipomyia) 56 furcifera MACFIE (Forcipomyia) 48 furens (POEY) (Culicoides) 17, 28, 33, 70, 150 furva INGRAM & MACFIE (Stilobezzia) 85 fusca (PHILIPPI) (Forcipomyia) 40, 50

fusca SPINELLI & WIRTH (Bezzia) 93 fuscipennis (LANE & WIRTH) (Downeshelea) 80 fuscipennis SPINELLI & GROGAN (Macrurohelea) 82 fuscipennis WIRTH (Parabezzia) 84 fusciradialis SPINELLI & WIRTH (Paryphoconus) 98 fuscivenosus (LUTZ) (Brachypogon) 79 fusculus (COQUILLETT) (Atrichopogon) 25, 47, 133 fuscus LANE (Paryphoconus) 98 fusicornis (COQUILLETT) (Forcipomyia) 56, 118 fusipalpis WIRTH & BLANTON (Culicoides) 68 gabaldoni ORTÍZ (Culicoides) 73 galapagensis (COQUILLETT) (Forcipomyia) 54 galindoi GROGAN & WIRTH (Amerohelea) 92 galindoi WIRTH & BLANTON (Culicoides) 73 galindoi WIRTH & SORIA (Forcipomyia) 59 galliarii MARINO & SPINELLI (Forcipomyia) 49 gallicus CLASTRIER (Leptoconops) 133, 135 gamboai BORKENT & PICADO (Atrichopogon) 45 gentilii SPINELLI & GROGAN (Macrurohelea) 82 genualis (LOEW) (Forcipomyia) 25, 50 germanus MACFIE (Culicoides) 66 gibbera (COQUILLETT) (Bezzia) 93 gibsoni WIRTH (Culicoides) 72 ginesi ORTÍZ (Culicoides) 66 glabellus WIRTH & BLANTON (Culicoides) 73 glaber MACFIE (Atrichopogon) 45 glabra (COQUILLETT) (Bezzia) 30, 93 glabrior MACFIE (Culicoides) 66 Glaphiromyia VARGAS 65 glauca MACFIE (Forcipomyia) 25, 48, 106 glauca MACFIE (Stilobezzia) 30, 87-88 globulifer MACFIE (Atrichopogon) 45 globulosa SPINELLI & WIRTH (Bezzia) 93 goeldii WIRTH & BLANTON (Culicoides) 74 goianensis LANE (Bezzia) 93 goianensis LANE (Paryphoconus) 98 gordoni MACFIE (Atrichopogon) 45 gorgasi WIRTH & BLANTON (Culicoides) 70 gracile (Dipetalonema) 33 grahambelli FORATTINI (Culicoides) 66 grandcolasi CLASTRIER & WIRTH (Forcipomyia) 54 grandis LANE & FORATTINI (Stilobezzia) 87 grandis MACFIE (Paryphoconus) 98 granditergitus BORKENT & PICADO (Atrichopogon) 45

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

granditibialis BORKENT & PICADO (Atrichopogon) 45 griseola WIRTH (Dasyhelea) 60 Groganhelea SPINELLI & DIPPOLITO 41, 90, 102103, 122, 128, 134, 143, 145, 150 grogani SPINELLI & WIRTH (Bezzia) 94 guadeloupensis DELÉCOLLE & RIEB (Dasyhelea) 60 guadeloupensis FLOCH & ABONNENC (Culicoides) 72 guaimiesi LANE & WIRTH (Monohelea) 83 guamai WIRTH & BLANTON (Culicoides) 74 guamai WIRTH & DOW (Forcipomyia) 57 guarani LANE (Palpomyia) 31, 96 guarani MARINO & SPINELLI (Forcipomyia) 50 guarani RONDEROS & SPINELLI (Culicoides) 72 guarani RONDEROS & SPINELLI (Forcipomyia) 52 guarani SPINELLI (Alluaudomyia) 76 guerrai WIRTH & BLANTON (Culicoides) 72 guianae (MACFIE) (Stilobezzia) 85 guianae (WIRTH) (Downeshelea) 80 guianae MACFIE (Paryphoconus) 98 guianensis MACFIE (Atrichopogon) 45 gustavoi BORKENT (Baeodasymyia) 77 guttatus (COQUILLETT) (Culicoides) 67-68 gutturosa (Onchocerca ) 33 guyana CLASTRIER & WIRTH (Forcipomyia) 54 guyana CLASTRIER (Palpomyia) 96 guyana CLASTRIER (Schizonyxhelea) 85 guyanensis FLOCH & ABONNENC (Culicoides) 28, 74 Haematomyidium GOELDI 66-67 haematopotus MALLOCH (Culicoides) 28, 64, 125 haitiensis CLASTRIER & RACCURT (Parabezzia) 84 haitiensis DELÉCOLLE (Culicoides) 65 harii SPINELLI & GROGAN (Borkenthelea) 78, 108 harpa SPINELLI & BORKENT (Forcipomyia) 57 harpegonata WIRTH & SORIA (Forcipomyia) 25, 50 harrisi MACFIE (Atrichopogon) 45 Hartomyia MALLOCH 85 hatoensis UTMAR & WIRTH (Forcipomyia) 48 hayesi MATTA (Culicoides) 62 heliconiae FOX & HOFFMAN (Culicoides) 68 herediae WIRTH & SPINELLI (Forcipomyia) 26, 53 hermani SPINELLI & BORKENT (Culicoides) 63 hermosa UTMAR & WIRTH (Forcipomyia) 48 hertigi WIRTH & BLANTON (Culicoides) 65 Heteromyia SAY 32, 89, 101, 117, 142, 151-152

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hieroglyphica KIEFFER (Monohelea) 82-83, 113 hippolytae MACFIE (Dasyhelea) 60 hirsuta INGRAM & MACFIE (Stilobezzia) 85 hirtipes (KIEFFER) (Dasyhelea) 60 hispida BORKENT (Palpomyia) 96 hitchcocki SPINELLI & WIRTH (Culicoides) 65 hobbsi WIRTH & SPINELLI (Forcipomyia) 53 hoffmani FOX (Culicoides) 28, 67 Hoffmania FOX 67 Holoconops KIEFFER 43 homofacies SPINELLI (Atrichopogon) 46 hondurensis HOFFMAN (Leptoconops) 43 hondurensis SPINELLI & BORKENT (Culicoides) 64 hondurensis SPINELLI & GROGAN (Parabezzia) 84 hondurensis SPINELLI & WIRTH (Bezzia) 94 hondurensis SPINELLI & WIRTH (Dasyhelea) 60 horacioi LANE (Clinohelea) 30, 88 horticola LUTZ (Culicoides) 67 hortorum WEYENBERGH (Didymophleps) 99 hugoi SPINELLI & GROGAN (Brachypogon) 79 hylas MACFIE (Culicoides) 28, 67-68 iberaensis SPINELLI & CAZORLA (Palpomyia) 96 ichesi RONDEROS & SPINELLI (Culicoides) 66 ignacioi FORATTINI (Culicoides) 68 imitator ORTÍZ (Culicoides) 67 impar (JOHANNSEN) (Brachypogon) 79 impusilloides SPINELLI & WIRTH (Culicoides) 63 inacayali SPINELLI & MARINO (Atrichopogon) 46 inaequalis SPINELLI & GROGAN (Parabezzia) 84 inamollae FOX & HOFFMAN (Culicoides) 67-68 incubans (MACFIE) (Forcipomyia) 56 incultus EWEN (Atrichopogon) 25, 46 inermicollis KIEFFER (Palpomyia) 96 ingrami SPINELLI & GROGAN (Paradasyhelea) 75, 111 insigniforceps MACFIE (Forcipomyia) 52 insignipalpis MACFIE (Atrichopogon) 46 insignipalpis MACFIE (Forcipomyia) 54 insignis LUTZ (Culicoides) 18, 22, 29, 33, 67-68, 111 insigniventris MACFIE (Atrichopogon) 46 insinuatus ORTÍZ & LEÓN (Culicoides) 17, 67 insularis SPINELLI & GROGAN (Brachypogon) 79 insularis SPINELLI & GROGAN (Palpomyia) 96 intermedia SAUNDERS (Forcipomyia) 26, 52 intrepida MACFIE (Forcipomyia) 58

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iquitosensis CLASTRIER & WIRTH (Forcipomyia) 54 iriartei FOX (Culicoides) 64 irwini GROGAN & WIRTH (Macrurohelea) 82 irwini SPINELLI & WIRTH (Culicoides) 64 Isoecacta GARRETT 76 Isohelea KIEFFER 79 Isthmohelea INGRAM & MACFIE 40, 82, 101, 115, 117, 140, 142, 151 ixodoides (FIEBRIG-GERTZ) (Forcipomyia) 18, 53-54 jamaicensis EDWARDS (Culicoides) 65 jamaicensis SPINELLI & WIRTH (Bezzia) 94 jamaicensis SPINELLI & WIRTH (Dasyhelea) 60 jamaicensis WIRTH (Echinohelea) 81 jamaicensis WIRTH (Forcipomyia) 58 jamaicensis WIRTH (Parabezzia) 84 jipajapae WIRTH (Forcipomyia) 58 jocosa SAUNDERS (Forcipomyia) 26, 56 johannseni (LANE) (Palpomyia) 96 johannseni (WIRTH) (Allohelea) 112 Johannsenomyia MALLOCH 32, 90-91, 95, 102, 122123, 129, 143-144, 146, 151 johnsoni COQUILLETT (Stenoxenus) 31, 99 jubata SPINELLI & WIRTH (Bezzia) 94 jurbergi FELIPPE-BAUER (Culicoides) 64 kampa FELIPPE-BAUER (Culicoides) 67 kawensis CLASTRIER & WIRTH (Forcipomyia) 54 keilini (SAUNDERS) (Forcipomyia) 26, 56 kiefferi LANE (Heteromyia) 89 kiefferi LANE (Paryphoconus) 98 kiefferi LANE (Stilobezzia) 87 kintzi WIRTH & BLANTON (Culicoides) 74 knowltoni BECK (Culicoides) 63 knowltoni CLASTRIER & WIRTH (Leptoconops) 43 koenigi DELÉCOLLE & RIEB (Dasyhelea) 60 kremeri SPINELLI & GROGAN (Clastrieromyia) 95 kuanoskeles MACFIE (Forcipomyia) 53 kuscheli WIRTH & BLANTON (Culicoides) 74 kuscheli WIRTH (Macrurohelea) 82 lacajae MACFIE (Atrichopogon) 46 lacrimatorii MACFIE (Forcipomyia) 53 lacustris INGRAM & MACFIE (Dasyhelea) 27, 60 lacustris LANE (Palpomyia) 31, 96 lacustris RONDEROS (Culicoides) 71 lahillei (ICHES) (Culicoides) 17, 29, 33, 67

lamprogaster EDWARDS (Heteromyia) 89 Lanehelea WIRTH & BLANTON 41, 90, 102-103, 122123, 134, 143-144, 151 lanei ORTÍZ (Culicoides) 74 lanei WIRTH & RATANAWORABHAN (Pellucidomyia) 89 lanei WIRTH & RATANAWORABHAN (Stenoxenus) 99 lanei WIRTH (Echinohelea) 29, 81, 112, 118 lanei WIRTH (Johannsenomyia) 90 lanei WIRTH (Monohelea) 83 lanei WIRTH (Paryphoconus) 31, 98 Lasiohelea KIEFFER 17, 20, 22, 26, 51-52, 58-59, 150 latifemoris (INGRAM & MACFIE) (Pachyhelea) 95 latipennis SPINELLI & WIRTH (Paryphoconus) 98 leei SPINELLI & WIRTH (Alluaudomyia) 76 leei SPINELLI & WIRTH (Bezzia) 94 leei SPINELLI & WIRTH (Paryphoconus) 98 leei WIRTH & BLANTON (Lanehelea) 90-91 leei WIRTH & BLANTON (Parastilobezzia) 84, 113, 119, 152 leei WIRTH & RATANAWORABHAN (Forcipomyia) 57 leei WIRTH (Echinohelea) 81 lenkoi LANE (Bezzia) 94 lenti TAVARES & LUNA DIAS (Culicoides) 70 leoni BARBOSA (Culicoides) 73 leopoldoi ORTÍZ (Culicoides) 72 Lepidohelea KIEFFER 26, 48, 52 Leptoconops SKUSE 16-17, 22, 25, 32-33, 43-44, 101, 103, 111, 129, 134, 136, 146-147, 151 Leptohelea WIRTH & BLANTON 40-41, 82, 101, 110, 138-139, 151 lesliei WIRTH (Forcipomyia) 26, 58-59 levis (COQUILLETT) (Atrichopogon) 44, 111, 118 lichyi FLOCH & ABONNENC (Culicoides) 73 limai BARRETTO (Culicoides) 73 limitrofe SPINELLI & FELIPPE-BAUER (Mallochohelea) 91 limnetis INGRAM & MACFIE (Forcipomyia) 58 limnochares (MACFIE) (Palpomyia) 96 limonensis ORTÍZ & LEÓN (Culicoides) 67 limpidus WIRTH & RATANAWORABHAN (Stenoxenus) 99 lineata (MEIGEN) (Palpomyia) 131 lituratus (WILLISTON) (Atrichopogon) 46 lobatoi FELIPPE-BAUER (Culicoides) 73 lobatus BORKENT & PICADO (Atrichopogon) 46 longicornis (WILLISTON) (Palpomyia) 97

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

longicornis EWEN (Atrichopogon) 25, 46 longispina SAUNDERS (Forcipomyia) 25, 49 lopesi BARRETTO (Culicoides) 73 Lophomyidium CORDERO 47 lota (WILLISTON) (Forcipomyia) 49 loughnani EDWARDS (Culicoides) 65 louriei (MACFIE) (Forcipomyia) 59 lutea (MALLOCH) (Stilobezzia) 112, 119 lutealaris WIRTH & BLANTON (Culicoides) 63 luteigenua WIRTH & SPINELLI (Forcipomyia) 26, 53 luteisquamosa WIRTH (Forcipomyia) 54 luteogrisea WIRTH & WILLIAMS (Dasyhelea) 60 luteovenus ROOT & HOFFMAN (Culicoides) 29, 64 lutzi COSTA LIMA (Culicoides) 68 lutzi LANE (Palpomyia) 96 lydiae BORKENT (Baeodasymyia) 77 lyrinotatus WIRTH & BLANTON (Culicoides) 74 Macfiehelea LANE 89 macfiei (COSTA LIMA) (Forcipomyia) 56 macfiei LANE (Echinohelea) 81 macfiei LANE (Paryphoconus) 98 macfiei LANE (Stilobezzia) 87 macfiei SPINELLI & GROGAN (Paradasyhelea) 75 Macfiella FOX 40, 69 macheti CLASTRIER & LEGRAND (Forcipomyia) 58 macieli TAVARES & RUIZ (Culicoides) 29, 70 macrostigma WIRTH & BLANTON (Culicoides) 74 Macrurohelea INGRAM & MACFIE 40, 82, 101, 116, 130, 141, 151 maculata LANE (Stilobezzia) 87 maculata MACFIE (Dasyhelea) 60 maculicrus INGRAM & MACFIE (Palpomyia) 96 maculipennis (COQUILLETT) (Monohelea) 83 maculipennis CLASTRIER (Atrichopogon) 46 maculithorax (WILLISTON) (Culicoides) 70 maculitibia LANE & FORATTINI (Stilobezzia) 86 maculosa INGRAM & MACFIE (Forcipomyia) 53 magna (SAUNDERS) (Forcipomyia) 26, 56 magnipalpis WIRTH & BLANTON (Culicoides) 73 magnus BORKENT & PICADO (Atrichopogon) 46, 95 maia LANE & FORATTINI (Stilobezzia) 85 malariologiensis PERRUOLO (Culicoides) 75 Mallochohelea WIRTH 32, 91, 102, 122, 128-129, 143, 145-146, 151

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manaosensis LANE & FORATTINI (Stilobezzia) 85 mapuche MARINO & SPINELLI (Forcipomyia) 53 mapuche SPINELLI (Brachypogon) 79 marginalis WIRTH & RATANAWORABHAN (Stenoxenus) 99 marini SPINELLI & DIPPOLITO (Forcipomyia) 50 marinkellei WIRTH & LEE (Culicoides) 64 marmosetae (Dipetalonema) 33 marshi WIRTH & BLANTON (Culicoides) 63 martinezi WIRTH & BLANTON (Culicoides) 74 maruim LUTZ (Culicoides) 17, 29, 69 Mataemyia VARGAS 40, 69 mathisi SPINELLI & WIRTH (Bezzia) 94 maya FELIPPE-BAUER (Monohelea) 83 maya SPINELLI & WIRTH (Paryphoconus) 98 maya SPINELLI & WIRTH (Phaenobezzia) 97 mayeri FORATTINI & LANE (Forcipomyia) 54 mayeri ORTÍZ (Monohelea) 83 mayeri WIRTH (Paryphoconus) 31, 98 mazaruni MACFIE (Bezzia) 94 mcateei WIRTH (Forcipomyia) 132 mckeeveri BRICKLE & HAGAN (Culicoides) 65 mediomunda MINAYA (Dasyhelea) 27, 60 Megaconops WIRTH & ATCHLEY 44 megatheca SPINELLI & WIRTH (Bezzia) 94 melleus (COQUILLETT) (Culicoides) 29, 75 mellichroa MACFIE (Palpomyia) 96 mendozae INGRAM & MACFIE (Atrichopogon) 46 menzeli CLASTRIER & WIRTH (Forcipomyia) 54 merceri CAZORLA & SPINELLI (Stilobezzia) 87 mesotibialis SPINELLI & WIRTH (Bezzia) 94 messersmithi WIRTH & DOW (Forcipomyia) 56-57 Metaforcipomyia SAUNDERS 26, 53 metagonatus WIRTH & BLANTON (Culicoides) 63 mexicanus HUERTA (Atrichopogon) 46 michaeli BORKENT (Baeodasymyia) 29, 77 microchela KIEFFER (Palpomyia) 96 Microconops KIEFFER 43 Microhelea KIEFFER 4, 18, 19-21, 53 micronyx WIRTH & BLANTON (Leptohelea) 82, 109, 112, 119, 126, 151 microtoma (KIEFFER) (Forcipomyia) 54 minasensis FELIPPE-BAUER (Culicoides) 65 minisquamosa WIRTH (Forcipomyia) 54 minitheca MARINO & SPINELLI (Forcipomyia) 50

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minor (WIRTH) (Nilobezzia) 130 mirsae ORTÍZ (Culicoides) 65 misionensis SPINELLI (Brachypogon) 79 misionensis SPINELLI (Paryphoconus) 98 miyamotoi WIRTH & BLANTON (Culicoides) 74 modesta CLASTRIER & RACCURT (Baeodasymyia) 77, 108 modesta LANE (Stilobezzia) 87 mojingaensis WIRTH & BLANTON (Culicoides) 69-70 monicae SPINELLI & BORKENT (Culicoides) 63 monicae SPINELLI & GROGAN (Brachypogon) 79 monilicornis (COQUILLETT) (Forcipomyia) 57, 106 monilicornis WIRTH & BLANTON (Rhynchohelea) 8485, 104, 106, 112, 152 Monohelea KIEFFER 32, 46, 77, 80-82, 101, 110, 127, 138, 151 monotheca SPINELLI & GROGAN (Macrurohelea) 82 monticola INGRAM & MACFIE (Dasyhelea) 60 monticola WIRTH & LEE (Culicoides) 73 moorei CLASTRIER & WIRTH (Forcipomyia) 55 morenoi MARINO & SPINELLI (Forcipomyia) 53 morrisoni GROGAN & WIENERS (Dasyhelea) 60 mortuifolii SAUNDERS (Forcipomyia) 25, 49 multidentata RONDEROS & SPINELLI (Forcipomyia) 52 multilineata (LUTZ) (Downeshelea) 80 multipicta INGRAM & MACFIE (Forcipomyia) 50 multispinosa SPINELLI & GROGAN (Palpomyia) 96 musae CLASTRIER & DELÉCOLLE (Forcipomyia) 26, 56 mutabilis (COQUILLETT) (Dasyhelea) 27, 59-60 muzoni MARINO & SPINELLI (Forcipomyia) 50 muzoni SPINELLI & DURET (Clinohelea) 88 Mycterotypus NOÈ 43 nana (MACFIE) (Forcipomyia) 58 nana WIRTH & BLANTON (Baeohelea) 77-78, 108109, 112, 149 Nannohelea GROGAN & WIRTH 83, 101, 107, 109, 138, 151 nanus MACFIE (Atrichopogon) 46 narthekophora MACFIE (Forcipomyia) 59 navaiae BYSTRAK & WIRTH (Forcipomyia) 49 nebulosus MACFIE (Atrichopogon) 46 necrophila SPINELLI & RODRÍGUEZ (Dasyhelea) 24, 27, 60 needhami THOMSEN (Alluaudomyia) 132 neivai LANE (Clinohelea) 88

nelsoni GROGAN & WIRTH (Amerohelea) 92 nemoralis (MACFIE) (Mallochohelea) 91 Neobezzia WIRTH & RATANAWORABHAN 31, 41, 91, 102, 120, 122, 128, 143, 145, 151 Neoceratopogon MALLOCH 76 neoparaensis TAVARES & SOUZA (Culicoides) 67 neopulicaris WIRTH (Culicoides) 29, 64 neotropica (MACFIE) (Nilobezzia) 92 neotropica CLASTRIER & WIRTH (Forcipomyia) 55 neotropica WIRTH (Allohelea) 76, 118, 148 neotropica WIRTH (Echinohelea) 81 neotropicalis (LANE) (Paryphoconus) 98 nerudai SPINELLI & GROGAN (Borkenthelea) 78 neunguis GROGAN & WIRTH (Parabezzia) 84 niger LANE (Stenoxenus) 99 nigerrima INGRAM & MACFIE (Stilobezzia) 85 nigra KIEFFER (Heteromyia) 89 nigrescens MACFIE (Forcipomyia) 48 nigrigenus WIRTH & BLANTON (Culicoides) 62-63 nigrimaxillata CLASTRIER & WIRTH (Forcipomyia) 55 nigripes (MACFIE) (Mallochohelea) 91 nigripes MACFIE (Clinohelea) 30, 88 nigripes MACFIE (Paryphoconus) 98 nigritibialis SPINELLI & WIRTH (Bezzia) 94 nigroflava LANE & FORATTINI (Stilobezzia) 87 nigroflava LANE (Palpomyia) 96 nigroscutellata LANE (Palpomyia) 96 Nilobezzia KIEFFER 32, 91, 102, 122, 128, 143, 145, 151 nobilis (WINNERTZ) (Bezzia) 31, 93-94, 114, 124, 130 nodosa SAUNDERS (Forcipomyia) 26, 58 nothofagus SPINELLI & GROGAN (Borkenthelea) 78, 112 Notiohelea GROGAN & WIRTH 40, 83, 101, 103, 117, 134, 142, 151 novaeteutoniae CLASTRIER & WIRTH (Forcipomyia) 55 novaeteutoniae MACFIE (Atrichopogon) 46 nubeculosa SPINELLI & WIRTH (Alluaudomyia) 76 nubeculosus MACFIE (Atrichopogon) 46 nubifer MACFIE (Paryphoconus) 98 nuncupata MACFIE (Forcipomyia) 50 obesa COSTA LIMA (Forcipomyia) 55 obfuscatus INGRAM & MACFIE (Atrichopogon) 46 obnoxius FOX (Culicoides) 74 obnubilus INGRAM & MACFIE (Atrichopogon) 46

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

obscura LANE & FORATTINI (Stilobezzia) 87 obscurus EWEN (Atrichopogon) 25, 46 ocumare (ORTÍZ) (Atrichopogon) 46 ocumarensis ORTÍZ (Culicoides) 69 Oecacta POEY 70 oedidactyla (INGRAM & MACFIE) (Physohelea) 90, 113 oklahomensis KHALAF (Culicoides) 29, 62 oligarthra SAUNDERS (Forcipomyia) 26, 56 oliveirai (LANE) (Pellucidomyia) 89 oliveirai LANE (Palpomyia) 96 oliveirai LANE (Paryphoconus) 31, 98 oliveri FOX & HOFFMAN (Culicoides) 69 opilionivora (LANE) (Forcipomyia) 58 orellana (ROBACK) (Heteromyia) 89 orjuelai WIRTH & LEE (Culicoides) 63 ornata LANE & FORATTINI (Stilobezzia) 85 ornata WIRTH (Monohelea) 83 ornaticrus INGRAM & MACFIE (Stilobezzia) 85 ornatipennis CLASTRIER (Atrichopogon) 46 ornatipennis MACFIE (Echinohelea) 81 ornatipennis MACFIE (Forcipomyia) 51 ortizi FOX (Culicoides) 71 ozzardi (Dipetalonema) 33 pabloi BROWNE (Culicoides) 72 pachycnemus MACFIE (Atrichopogon) 46 Pachyhelea WIRTH 32, 41, 95, 102, 123, 128, 144145, 152 Pachyleptus WALKER 89 pachymera (WILLISTON) (Pachyhelea) 31, 95, 124, 152 pachymerus LUTZ (Culicoides) 74 Padrosia RAFINESQUE 61 painteri FOX (Culicoides) 68 pallescens LANE & FORATTINI (Stilobezzia) 87 pallida SPINELLI & GROGAN (Parabezzia) 84 pallidipennis SPINELLI & GROGAN (Brachypogon) 79 pallidipes KIEFFER (Atrichopogon) 46 palpalis MACFIE (Atrichopogon) 46 palpalis MACFIE (Culicoides) 69 Palpomyia MEIGEN 23, 30, 32, 79-80, 86, 89, 90, 92, 95, 102, 122-123, 128, 143-145, 152 pampaensis SPINELLI & WIRTH (Culicoides) 67 pampana LANE (Palpomyia) 96 pampoikilus MACFIE (Culicoides) 71 panamensis (LANE & WIRTH) (Downeshelea) 80

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panamensis BARBOSA (Culicoides) 65 panamensis LANE & FORATTINI (Stilobezzia) 30, 87 panamensis RONDEROS & SPINELLI (Leptoconops) 44 panamensis WIRTH (Echinohelea) 81 panamensis WIRTH (Parabezzia) 84 panamericanus FOX (Culicoides) 71 pancensis BROWNE (Culicoides) 75 Parabezzia MALLOCH 32, 83, 101, 104, 109, 110, 115-116, 139-141, 150, 152-153 paracaudata GROGAN & WIRTH (Macrurohelea) 82 paracincta WIRTH (Dasyhelea) 27, 60 Paradasyhelea MACFIE 32, 40, 75, 101, 105, 137, 152 paraensis (GOELDI) (Culicoides) 17, 24, 29, 33, 66-67 paraensis (LANE) (Nilobezzia) 92 paraensis LANE (Palpomyia) 96 paraensis LANE (Stenoxenus) 99 paraensis WIRTH & BLANTON (Brachypogon) 79, 112 parahybae MACFIE (Dasyhelea) 60 paraignacioi SPINELLI (Culicoides) 69 paramaruim WIRTH & BLANTON (Culicoides) 69 paranaensis SPINELLI & WIRTH (Paryphoconus) 98 parascopus WIRTH & BLANTON (Culicoides) 65 Parastilobezzia WIRTH & BLANTON 40-41, 84, 101, 110, 139 Paryphoconus ENDERLEIN 32, 41, 97, 102, 123, 128129, 144-146, 152 patagonica INGRAM & MACFIE (Dasyhelea) 60 patagonica INGRAM & MACFIE (Forcipomyia) 50 patagonica INGRAM & MACFIE (Palpomyia) 96 patagonica INGRAM & MACFIE (Stilobezzia) 85 patagonica SPINELLI & GROGAN (Diaphanobezzia) 80 patagoniensis RONDEROS (Leptoconops) 43 patagoniensis RONDEROS & SPINELLI (Culicoides) 17, 64 patulipalpis WIRTH & BLANTON (Culicoides) 72 paucienfuscatus BARBOSA (Culicoides) 74 paulistana FORATTINI & RABELLO (Dasyhelea) 27, 61 paulistensis LANE (Palpomyia) 96 paulistensis LANE (Paryphoconus) 98 paulistensis LANE (Stilobezzia) 87 pectinatus MACFIE (Atrichopogon) 46 Pedilohelea DE MEILLON & WIRTH 48, 55 pellucida INGRAM & MACFIE (Diaphanobezzia) 80 Pellucidomyia MACFIE 32, 89, 101, 117, 142, 152 penicillatus DELÉCOLLE & RIEB (Atrichopogon) 46

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penthesileae MACFIE (Dasyhelea) 27, 60 penultimata WIRTH (Forcipomyia) 55 perstans (Dipetalonema ) 33 peruviana CLASTRIER & WIRTH (Forcipomyia) 55 peruvianus FELIPPE-BAUER (Culicoides) 67 peruvianus KIEFFER (Atrichopogon) 46 petiolata MALLOCH (Parabezzia) 83, 113 petrocchiae SHANNON & DEL PONTE (Leptoconops) 44, 151 Phaenobezzia HAESELBARTH 32, 97, 102, 120, 122, 128, 143, 145, 152 phaeonotus WIRTH & BLANTON (Culicoides) 71 Phasmidohelea MAYER 53-55 philotherma MACFIE (Dasyhelea) 61 phlebotomus (WILLISTON) (Culicoides) 17-18, 29, 33, 69 Physohelea GROGAN & WIRTH 40, 90, 102, 117, 142, 152 Phytohelea REMM 26, 55 picadoae SPINELLI & BORKENT (Culicoides) 72 piceiventris KIEFFER (Atrichopogon) 46 pichindensis BROWNE (Culicoides) 73 picta (COQUILLETT) (Stilobezzia) 87 pictipennis (PHILIPPI) (Culicoides) 70-71 pictipennis CLASTRIER (Atrichopogon) 46 pictoni MACFIE (Forcipomyia) 26, 51 pifanoi ORTÍZ (Culicoides) 72, 74 pilosa SPINELLI & GROGAN (Notiohelea) 83 pilosus WIRTH & BLANTON (Culicoides) 65 pinamarensis SPINELLI (Forcipomyia) 51 piroskyi CAVALIERI (Forcipomyia) 50 pistiae (INGRAM & MACFIE) (Phaenobezzia) 97, 114, 130 platensis (BRÈTHES) (Forcipomyia) 49 plaumanni CLASTRIER & WIRTH (Forcipomyia) 55 plaumanni SPINELLI & WIRTH (Alluaudomyia) 76 plaumanni SPINELLI (Culicoides) 69 plebeja (LOEW) (Palpomyia) 114, 124 poikilonotus MACFIE (Culicoides) 65 pollex BORKENT & FORSTER (Dasyhelea) 61 polypori WIRTH & BLANTON (Culicoides) 69 polystictus KIEFFER (Culicoides) 75 poncai LANE & WIRTH (Monohelea) 83 popayanensis WIRTH & LEE (Culicoides) 63 poulaineae INGRAM & MACFIE (Forcipomyia) 50-51 prima CLASTRIER (Alluaudomyia) 76

propinqua (WILLISTON) (Forcipomyia) 50 propinquus MACFIE (Culicoides) 75 propriipennis MACFIE (Culicoides) 72 Protersesthes KIEFFER 43 pseudocrescentis TAVARES & LUNA DIAS (Culicoides) 72 Pseudoculicoides MALLOCH 59 pseudodiabolicus FOX (Culicoides) 17, 69 pseudofasciata GROGAN & WIRTH (Amerohelea) 92 pseudofulvus SPINELLI (Stenoxenus) 99 pseudogibbera SPINELLI & WIRTH (Bezzia) 94 pseudoincisurata WAUGH and WIRTH (Dasyhelea) 106, 111, 118, 125 pseudolacustris DIPPOLITO & SPINELLI (Palpomyia) 96 pseudoparaensis SPINELLI & GROGAN (Brachypogon) 79 pseudovenustula SPINELLI & WIRTH (Bezzia) 94 pseudunguis SPINELLI & GROGAN (Parabezzia) 84 Psychophaena PHILIPPI 40, 70-71 Pterobosca MACFIE 20, 56 pulchripennis MACFIE (Culicoides) 73 pulchripennis MACFIE (Sphaeromias) 76 pulchripes KIEFFER (Bezzia) 31, 94 pulicaris (LINNAEUS) (Culicoides) 132 pullata (WIRTH) (Mallochohelea) 91 pumila MACFIE (Dasyhelea) 61 punctipennis (WILLISTON) (Bezzia) 93-94 punctipes WIRTH (Stilobezzia) 87 punctiradialis SPINELLI & WIRTH (Alluaudomyia) 77 punctivenosa WIRTH & GROGAN (Alluaudomyia) 77 punctulata LANE (Stilobezzia) 87 puracensis WIRTH & DOW (Forcipomyia) 57 puracensis WIRTH & LEE (Culicoides) 17, 63 pusilla (LUTZ) (Dasyhelea) 27, 61 pusilloides WIRTH & BLANTON (Culicoides) 63 pusillus LUTZ (Culicoides) 22, 33, 63 pygmaea (WILLISTON) (Dasyhelea) 61 pyrsonota MACFIE (Dasyhelea) 61 quartibrunneus BORKENT & PICADO (Atrichopogon) 46 quasicornuta SAUNDERS (Forcipomyia) 26, 52 quasidentica FELIPPE-BAUER & QUINTELAS (Downeshelea) 81 quasiingrami MACFIE (Forcipomyia) 49 quasiparaensis CLASTRIER (Culicoides) 67 quatei SPINELLI & GROGAN (Borkenthelea) 78 quatei WIRTH (Forcipomyia) 50

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

quechua MARINO & SPINELLI (Forcipomyia) 151 rabelloi LANE (Stilobezzia) 30, 87 raccurti SPINELLI & GROGAN (Parabezzia) 84 rachoui TAVARES & SOUZA (Culicoides) 67 raleighi MACFIE (Forcipomyia) 25, 50 rangeli ORTÍZ & MIRSA (Culicoides) 72 raposoensis CLASTRIER & WIRTH (Forcipomyia) 55 raposoensis SPINELLI & WIRTH (Bezzia) 94 raposoensis WIRTH & BARRETO (Culicoides) 64 raposoi DE MEILLON & WIRTH (Forcipomyia) 55 rava INGRAM & MACFIE (Stilobezzia) 86 recifei BARBOSA (Culicoides) 69 recifensis BARBOSA (Culicoides) 74 reconditus CAMPBELL & PELHAM-CLINTON (Culicoides) 135 redactus BORKENT & PICADO (Atrichopogon) 46 remigatus EWEN (Atrichopogon) 25, 46 remigera UTMAR & WIRTH (Forcipomyia) 48 reticulatus LUTZ (Culicoides) 29, 74 rettenmeyerorum CLASTRIER & WIRTH (Forcipomyia) 55 reynoldsi INGRAM & MACFIE (Dasyhelea) 61 Rhynchoforcipomyia WIRTH & DOW 40, 48, 56 Rhynchohelea WIRTH & BLANTON 84, 101, 103, 107, 134, 137-138, 152 ricardoi RONDEROS & SPINELLI (Leptoconops) 44 richardsi MACFIE (Echinohelea) 81 ringueleti SPINELLI (Brachypogon) 79 riojana SPINELLI & MARINO (Forcipomyia) 58 rioplatensis MARINO & SPINELLI (Forcipomyia) 26, 51 rodriguezi ORTÍZ (Culicoides) 72 roldani SPINELLI & WIRTH (Bezzia) 31, 94 ronderosae SPINELLI & BORKENT (Culicoides) 65 ronderosi GROGAN & WIRTH (Amerohelea) 92 rondoniensis SPINELLI & DIPPOLITO (Groganhelea) 90, 114, 120, 150 roraimensis FELIPPE-BAUER & SPINELLI (Monohelea) 83 rostratus WIRTH & BLANTON (Culicoides) 25, 63 roubaudi CLASTRIER & DELÉCOLLE (Forcipomyia) 58 rozeboomi BARBOSA (Culicoides) 68-69 rubriceps KIEFFER (Clinohelea) 88 rufa KIEFFER (Heteromyia) 89 ruizi FORATTINI (Culicoides) 69 rusticus MACFIE (Atrichopogon) 47 sabalitensis UTMAR & WIRTH (Forcipomyia) 48

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saintjusti TAVARES & RUIZ (Culicoides) 68 Saliohelea WIRTH & RATANAWORABHAN 48, 57 saltaensis SPINELLI & WIRTH (Culicoides) 65 saltanensis LANE & DURET (Clinohelea) 88 saltensis (CAVALIERI) (Forcipomyia) 52 sanctaeclarae MACFIE (Atrichopogon) 47 sanctaeclarae WIRTH (Forcipomyia) 57 sanctibernardini KIEFFER (Stilobezzia) 86 sanctilaurentii KIEFFER (Atrichopogon) 47 sanmartini WIRTH & BARRETO (Culicoides) 67 santaemarthae (KIEFFER) (Dasyhelea) 61 santanderi BROWNE (Culicoides) 73 saundersi EWEN (Atrichopogon) 25, 47, 132 sayhuequei MARINO & SPINELLI (Forcipomyia) 58 scabra (COQUILLETT) (Palpomyia) 96 Schizoforcipomyia CHAN & LEROUX 57 Schizonyxhelea CLASTRIER 30, 32, 41, 85, 101, 107, 138, 152 schmitzi SPINELLI & GROGAN (Brachypogon) 79 schmitzorum DIPPOLITO & SPINELLI (Bezzia) 94 schnacki SPINELLI & GROGAN (Clastrieromyia) 95, 114 schnacki SPINELLI (Alluaudomyia) 29, 77 schwarzii (COQUILLETT) (Nilobezzia) 30, 92 scissurae MACFIE (Dasyhelea) 61 scopus ROOT & HOFFMAN (Culicoides) 65 scorzai ORTÍZ (Culicoides) 75 scutata LANE & FORATTINI (Stilobezzia) 87 seminole WIRTH (Forcipomyia) 26, 52-53 sentior (INGRAM & MACFIE) (Austrosphaeromias) 90 sequax (WILLISTON) (Atrichopogon) 47 setifer (LUTZ) (Culicoides) 64 setiger MACFIE (Stenoxenus) 99 setigera SAUNDERS (Forcipomyia) 25, 49 setigera SPINELLI & WIRTH (Bezzia) 94 setosa WIRTH (Macrurohelea) 82 setosicrus (KIEFFER) (Forcipomyia) 49 setosilateralis BORKENT & PICADO (Atrichopogon) 47 seudoobfuscatus SPINELLI (Atrichopogon) 47 sexpunctata SPINELLI & WIRTH (Alluaudomyia) 77 sexvittata WIRTH (Forcipomyia) 51 shannoni (INGRAM & MACFIE) (Forcipomyia) 58 shannoni (WIRTH & BLANTON) (Austrohelea) 77, 113, 148 shannoni INGRAM & MACFIE (Dasyhelea) 61

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silvicola MACFIE (Stilobezzia) 87 similans LANE & FORATTINI (Stilobezzia) 87 similis SPINELLI & GROGAN (Macrurohelea) 82 similis SPINELLI & MARINO (Atrichopogon) 47 similis SPINELLI (Amerohelea) 92 simplex LANE & FORATTINI (Stilobezzia) 87 sinclairi BORKENT (Dasyhelea) 61 sinuosa DOW & WIRTH (Forcipomyia) 58 smarti MACFIE (Echinohelea) 81 smithi (LEWIS) (Mallochohelea) 114 snowi LANE (Bezzia) 31, 94 soibelzoni MARINO & SPINELLI (Forcipomyia) 57 somuncurensis MARINO & SPINELLI (Forcipomyia) 51 sonorensis WIRTH & RATANAWORABHAN (Paryphoconus) 98 sordidipes (MACFIE) (Amerohelea) 30, 92 soriai WIRTH & WAUGH (Dasyhelea) 61 soriai WIRTH (Forcipomyia) 51 spangleri DE MEILLON & WIRTH (Forcipomyia) 55 spangleri WIRTH (Parabezzia) 84 spathicerca WIRTH (Dasyhelea) 61 spatulifera SAUNDERS (Forcipomyia) 25, 49 spatuliformis SPINELLI & GROGAN (Brachypogon) 79 spatuligera MACFIE (Forcipomyia) 51 Sphaerohelea SPINELLI & FELIPPE-BAUER 41, 92, 102-103, 122-123, 134, 143-144, 152 spilmani DE MEILLON & WIRTH (Forcipomyia) 55 spinellii GROGAN & WIRTH (Amerohelea) 82 spinellii WIRTH & GROGAN (Alluaudomyia) 77 spinellii WIRTH & GROGAN (Diaphanobezzia) 80 spinifemur (LANE) (Palpomyia) 96 spinifemur WIRTH & BLANTON (Lanehelea) 91 spinosa LUTZ (Palpomyia) 96 spinosa SAUNDERS (Forcipomyia) 26, 59 spinosus BORKENT & PICADO (Atrichopogon) 47 spurius WIRTH & BLANTON (Culicoides) 67 squamithorax CLASTRIER (Forcipomyia) 53 squamitibia LUTZ (Forcipomyia) 49, 51 squamosa LUTZ (Forcipomyia) 55 steineri SPINELLI & WIRTH (Paryphoconus) 98 Stenoxenus COQUILLETT 32, 99, 102, 122-123, 129, 143-144, 146, 152 stigmalis (COQUILLETT) (Brachypogon) 111 stigmalis WIRTH (Culicoides) 75

Stilobezzia KIEFFER 17, 24, 32, 40, 84-86, 101103, 115-117, 126, 134, 140-142, 149, 152 stonei (WIRTH) (Downeshelea) 80-81 stubalensis FOX (Culicoides) 74 stylifer (LUTZ) (Forcipomyia) 26, 51-52 suarezi RODRÍGUEZ & WIRTH (Culicoides) 63 suarezi SPINELLI & RONDEROS (Dasyhelea) 61 subaspera (COQUILLETT) (Palpomyia) 31, 96 subflavus MACFIE (Paryphoconus) 98 subfusca MACFIE (Bezzia) 94 subfuscula INGRAM & MACFIE (Palpomyia) 97 subsessilis KIEFFER (Stilobezzia) 86 succinea INGRAM & MACFIE (Stilobezzia) 86 Synthyridomyia SAUNDERS 57 taeniatus MACFIE (Atrichopogon) 47 talarum SPINELLI (Atrichopogon) 47 tamboensis WIRTH & LEE (Culicoides) 72 tamioi LANE (Palpomyia) 97 tanycornis BORKENT (Palpomyia) 97 tapantiensis BORKENT & PICADO (Atrichopogon) 47 taragui MARINO (Forcipomyia) 50 taragui SPINELLI (Paryphoconus) 98 tarapaca SPINELLI & WIRTH (Culicoides) 65 tavaresi FELIPPE-BAUER & WIRTH (Culicoides) 74 tehuelche MARINO & SPINELLI (Forcipomyia) 58 telesfordi SPINELLI & GROGAN (Brachypogon) 79 telmatophilus (MACFIE) (Paryphoconus) 98 tenuiannulata SPINELLI & WIRTH (Alluaudomyia) 77 tenuicrus KIEFFER (Palpomyia) 97 tenuiforceps MACFIE (Forcipomyia) 57 tenuilobus WIRTH & BLANTON (Culicoides) 73 tenuisquamipes WIRTH (Forcipomyia) 51 teretipalpis WIRTH & BARRETO (Culicoides) 71 terminalis (COQUILLETT) (Paryphoconus) 98 termophila (SPINELLI) (Mallochohelea) 30, 91 terrestris SAUNDERS (Forcipomyia) 26, 59 Tersesthes TOWNSEND 43-44 Tetraphora PHILIPPI 48, 50 tetrathyris WIRTH & BLANTON (Culicoides) 73 tettigonaris WIRTH & CASTNER (Forcipomyia) 55 thalestris MACFIE (Dasyhelea) 61 thersites (WILLISTON) (Atrichopogon) 47 thomasi CLASTRIER & WIRTH (Forcipomyia) 55 thomsenae WIRTH (Stilobezzia) 87

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae

thoracica INGRAM & MACFIE (Macrurohelea) 82 Thyridomyia SAUNDERS 57 tibialis LANE & FORATTINI (Stilobezzia) 86 tidwelli SPINELLI (Culicoides) 69 tirzae BORKENT & PICADO (Atrichopogon) 47 todatangae WIRTH & BLANTON (Culicoides) 67 torrens (TOWNSEND) (Leptoconops) 43, 125 townesi LANE (Clinohelea) 88 townsendi KNAB (Forcipomyia) 51 townsendi LANE (Clinohelea) 88 transferrans ORTÍZ (Culicoides) 72 transversa LANE & FORATTINI (Stilobezzia) 87 trapidoi WIRTH & BARRETO (Culicoides) 63 travassosi FORATTINI (Culicoides) 69 travassosi LANE (Paryphoconus) 98 travassosi LANE (Stilobezzia) 88 Trichohelea GOETGHEBUER 58 tricoloratus WIRTH & BLANTON (Culicoides) 74 trifidus SPINELLI & BORKENT (Culicoides) 73 trilineatus FOX (Culicoides) 75 trinidadensis HOFFMAN (Culicoides) 69 trinidadensis MACFIE (Atrichopogon) 47 trinidadensis SAUNDERS (Forcipomyia) 26, 58 tripunctata SPINELLI & WIRTH (Alluaudomyia) 77 tropica (KIEFFER) (Forcipomyia) 54 tuberculata SAUNDERS (Forcipomyia) 27, 59 tuberculatus EWEN (Atrichopogon) 25, 47 turgidipes (INGRAM & MACFIE) (Physohelea) 90 ugandae MACFIE (Pellucidomyia) 119 umbella MACFIE (Palpomyia) 97 umbratilis MACFIE (Atrichopogon) 47 uncinata (JOHANNSEN) (Parabezzia) 84 undecimpunctatus KIEFFER (Culicoides) 67 unetensis PERRUOLO (Culicoides) 75 unguis WIRTH (Parabezzia) 84 unica BYSTRAK & WIRTH (Forcipomyia) 49 unimaculatus MACFIE (Paryphoconus) 98 uniradialis WIRTH & BLANTON (Culicoides) 74 univesicula MACFIE (Forcipomyia) 58 uramaensis SPINELLI & DIPPOLITO (Forcipomyia) 50 urracaisi LANE & WIRTH (Monohelea) 83 uruguayensis (CORDERO) (Atrichopogon) 47 uruguayensis FELIPPE-BAUER & SPINELLI (Monohelea) 83 uruguayensis RONDEROS (Culicoides) 65

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uruguayensis SPINELLI & GROGAN (Clastrieromyia) 95 utae KNAB (Forcipomyia) 51 utricularis MACFIE (Atrichopogon) 47 valleensis CLASTRIER & WIRTH (Forcipomyia) 55 vargasi GROGAN & WIRTH (Amerohelea) 92 vargasi WIRTH & BLANTON (Culicoides) 64 varia INGRAM & MACFIE (Stilobezzia) 86 varicolor SAUNDERS (Forcipomyia) 25, 48 variipennis COQUILLETT (Culicoides) 5 varipennis WIRTH & WILLIAMS (Forcipomyia) 53 venezuelensis ORTÍZ & MIRSA (Culicoides) 29, 70-71 venezuelensis ORTÍZ (Leptoconops) 44 venezuelensis ORTÍZ (Stilobezzia) 88 venustula (WILLISTON) (Bezzia) 93-94 verecundus MACFIE (Culicoides) 69 vernoni CLASTRIER & WIRTH (Forcipomyia) 55 vernoni WIRTH & BLANTON (Culicoides) 73 versicolor MACFIE (Palpomyia) 97 villosipes (KIEFFER) (Dasyhelea) 61 viridans FORATTINI & RABELLO (Dasyhelea) 61 volcanensis WIRTH & BLANTON (Culicoides) 70 wagneri (SÉGUY) (Forcipomyia) 55 wallacei WIRTH & BLANTON (Culicoides) 70 Warmkea SAUNDERS 58 warmkei WIRTH (Atrichopogon) 47 weemsi WIRTH & SPINELLI (Forcipomyia) 53 williamsi CAZORLA & SPINELLI (Stilobezzia) 88 williamsi MARINO & SPINELLI (Forcipomyia) 53 williamsi SPINELLI (Culicoides) 73 williamsi WIRTH & WAUGH (Dasyhelea) 61 willistoni WIRTH & BLANTON (Culicoides) 69 willistoni WIRTH (Forcipomyia) 55 winderi WIRTH & WAUGH (Dasyhelea) 61 winderi WIRTH (Forcipomyia) 53 wirthi (LANE) (Pellucidomyia) 89 wirthi CLASTRIER & RACCURT (Parabezzia) 84 wirthi LANE & FORATTINI (Stilobezzia) 88 wirthi LANE (Palpomyia) 31, 97 wirthi LANE (Paryphoconus) 99 wirthi SPINELLI & FELIPPE-BAUER (Neobezzia) 91 wirthi SPINELLI & GROGAN (Macrurohelea) 82 wirthi SPINELLI (Austrosphaeromias) 90 wirthi SPINELLI (Brachypogon) 79 wirthomyia VARGAS (Culicoides) 65

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wokei BARBOSA (Culicoides) 69 wokei FOX (Culicoides) 69, 75 wokei WIRTH & GROGAN (Heteromyia) 30, 89 woodfordi MACFIE (Atrichopogon) 47 woodruffi SPINELLI & GROGAN (Brachypogon) 79 woodruffi SPINELLI & WIRTH (Bezzia) 94 wygodzinskyi CAVALIERI (Forcipomyia) 50 wygodzinskyi LANE (Clinohelea) 89 wygodzinskyi LANE (Paryphoconus) 99 wygodzinskyi LANE (Stilobezzia) 30, 88 xanifer WIRTH & BLANTON (Culicoides) 69 yamana MARINO & SPINELLI (Forcipomyia) 51

yamana SPINELLI & GROGAN (Macrurohelea) 82 yaracuyensis ORTÍZ (Culicoides) 73 yolancae BORKENT & PICADO (Atrichopogon) 47 youngi SPINELLI & WIRTH (Alluaudomyia) 77 youngi WIRTH & BARRETO (Culicoides) 67 youngi WIRTH & GROGAN (Cacaohelea) 79, 111, 119 youngi WIRTH (Forcipomyia) 50 yukonensis HOFFMAN (Culicoides) 104 zeteki WIRTH & DOW (Forcipomyia) 57 zonogaster INGRAM & MACFIE (Forcipomyia) 51 zumbadoi SPINELLI & BORKENT (Culicoides) 72