Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species (Ephemeroptera, Plecoptera, and Trichoptera) [1 ed.] 2017933502

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TB 1109 March 2017

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Larvae Larvae of of the the Southeastern Southeastern USA USA Mayfly, Mayfly, Stonefly, Stonefly, and and Caddisfly Caddisfly Species Species (Ephemeroptera, Plecoptera, and Trichoptera) John C. Morse, W. Patrick McCafferty, Bill P. Stark, and Luke M. Jacobus, Editors

South Carolina Agriculture and Forestry Research System

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species (Ephemeroptera, Plecoptera, and Trichoptera) John C. Morse, W. Patrick McCafferty, Bill P. Stark, and Luke M. Jacobus, Editors

Clemson University Clemson, South Carolina 2017

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Published by Clemson University Public Service Publishing Copyright © 2017 Clemson University 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 publisher. Printed in the United States. Find Clemson University Public Service Publishing on the World Wide Web at https://secure.touchnet.net/C20569_ustores/web/store_main.jsp?STOREID=168&SINGLESTORE=true. Library of Congress Control Number: 2017933502 First Printing: March 2017 Front Cover: Photographers Eric Fleek (insects), Coleson Wrege (backgrounds); Editor Morgan Summerlin Top center: Drunella lata (Morgan, 1911) Bottom left: Agnetina capitata (Pictet, 1841) Bottom right: Ceraclea ophioderus (Ross, 1938) Back Cover: Mouth of Injun Creek, tributary of Little Pigeon River, near Greenbrier Ranger Station, Great Smoky Mountains National Park, Sevier County, Tennessee--Habitat for a very diverse community with mayflies, stoneflies, and caddisflies. Photo taken about 1:30 pm, 2 Aug 2000, during a foggy, drizzling rain. A number 14 Hare’s Ear was the fly of the day for a fisherman I met. Photographer: Luke M. Jacobus. Image Manipulation, Layout, and Design: Virginia Winn, Columbia, South Carolina Julie Sellers, Columbia, South Carolina

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BIOTA OF SOUTH CAROLINA Volume 9 A Publication Series of the Biodiversity Initiative College of Agriculture, Forestry, and Life Sciences Clemson University Clemson, S.C. EDITORIAL BOARD Peter H. Adler Stephen H. Bennett Patrick D. McMillan Paula Levin Mitchell David W. Tonkyn John F. Townsend Alfred G. Wheeler, Editor

Clemson University South Carolina Department of Natural Resources Clemson University Winthrop University Clemson University Virginia Natural Heritage Program Clemson University

INFORMATION FOR AUTHORS The Biota of South Carolina publication series is intended to promote the study and public understanding of biodiversity at the genetic, species, ecosystem, and landscape scales of integration. Papers for this series should make a substantial contribution to a taxonomic group or to a geographic region and ordinarily should contain keys and illustrations. All submissions are peer reviewed. Authors are encouraged to discuss proposed contributions with the Editor before submitting a manuscript and to consult previous papers in the series for style and format. Three copies of the manuscript, including illustrations, should be sent to the Editor: Alfred G. Wheeler, Department of Plant & Environmental Sciences, Clemson University, Clemson, SC 29634-0310 ([email protected]). SUGGESTED CITATION FORM Morse, J.C., W.P McCafferty, B.P. Stark, & L.M. Jacobus, Editors. 2017. Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species. Biota of South Carolina. Vol 9. Clemson University Public Service Publishing, Clemson University, Clemson, South Carolina, USA. 482 pp. PURCHASING COPIES Copies of this publication may be purchased from the Public Service Bulletin Room, 96 Poole Agricultural Center, Clemson University, Clemson, S.C. 29634-0129 OR on the World Wide Web: https://secure. touchnet.net/C20569_ustores/web/store_cat.jsp?STOREID=168&CATID=298&SINGLESTORE=true for $40.00 each. Make checks payable to Clemson University.

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BIOTA OF SOUTH CAROLINA Volumes in the Series Volume 1 Ground Beetles and Wrinkled Bark Beetles of South Carolina (Coleoptera: Geadephaga: Carabidae and Rhysodidae) Volume 2 Scarab Beetles (Coleoptera: Scarabaeidae) of South Carolina Volume 3 Water Beetles of South Carolina (Coleoptera: Gyrinidae, Haliplidae, Noteridae, Dytiscidae, Hydrophilidae, Hydraenidae, Scirtidae, Elmidae, Dryopidae, Limnichidae, Heteroceridae, Psephenidae, Ptilodactylidae, and Chelonariidae) Volume 4 Leaf and Seed Beetles of South Carolina (Coleoptera: Chrysomelidae and Orsodacnidae) Volume 5 Rhynchospora (Cyperaceae) of South Carolina and the Eastern United States Volume 6 Weevils of South Carolina (Coleoptera: Nemonychidae, Attelabidae, Brentidae, Ithyceridae, and Curculionidae) Volume 7 Jewel Beetles (Coleoptera: Buprestidae) of South Carolina Volume 8 Tenebrionoidea of South Carolina (Coleoptera: Mycetophagidae, Archeocrypticidae, Tetratomidae, Melandryidae, Mordellidae, Ripiphoridae, Zopheridae, Tenebrionidae, Synchroidae, Oedemeridae, Stenotrachelidae, Meloidae, Mycteridae, Boridae, Pythidae, Pyrochroidae, Salpingidae, Anthicidae, Ischaliidae, and Aderidae) Volume 9 Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species (Ephemeroptera, Plecoptera, and Trichoptera)

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Table of Contents

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species (Ephemeroptera, Plecoptera, and Trichoptera)

John C. Morse, W. Patrick McCafferty, Bill P. Stark, and Luke M. Jacobus, Editors

Table of Contents Chapter 1. INTRODUCTION, J.C. Morse, B.P. Stark, and L.M. Jacobus...............................................10 Chapter 2. EPHEMEROPTERA, W.P. McCafferty, L.M. Jacobus, A.V. Provonsha, and N.A. Wiersema............................................................................................................................15 INTRODUCTION and major characteristics................................................................................15 LIST of families, genera, and species............................................................................................18 KEY to families (some genera and species)...................................................................................24 AMELETIDAE...............................................................................................................................40 Ameletus, key to species.....................................................................................................40 BAETIDAE, key to genera and species..........................................................................................42 Acentrella, key to species....................................................................................................54 Acerpenna, key to species...................................................................................................56 Anafroptilum, key to species...............................................................................................56 Apobaetis, key to species....................................................................................................58 Baetis, key to species..........................................................................................................58 Baetopus, key to species.....................................................................................................60 Callibaetis, key to species...................................................................................................62 Heterocloeon, key to species..............................................................................................62 Iswaeon, key to species.......................................................................................................64 Labiobaetis, key to species.................................................................................................66 Neocloeon, key to species...................................................................................................66 Paracloeodes, key to species..............................................................................................66 Plauditus, key to species.....................................................................................................68 Procloeon, key to species....................................................................................................70 BAETISCIDAE..............................................................................................................................74 Baetisca, key to species......................................................................................................74 CAENIDAE genera and species.....................................................................................................76 Brachycercus, key to species..............................................................................................78 Caenis, key to species.........................................................................................................78 Cercobrachys, key to species..............................................................................................80 Sparbarus, key to species...................................................................................................82 EPHEMERELLIDAE, key to genera and species..........................................................................82 Attenella, key to species......................................................................................................84 Dannella, key to species.....................................................................................................84 Drunella, key to species......................................................................................................86 Ephemerella, key to species................................................................................................88 Eurylophella, key to species...............................................................................................90 Serratella, key to species....................................................................................................94

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EPHEMERIDAE genera and species.............................................................................................96 Ephemera, key to species....................................................................................................96 Hexagenia, key to species...................................................................................................98 HEPTAGENIIDAE, key to genera and species............................................................................100 Epeorus, key to species.....................................................................................................104 Heptagenia, key to species...............................................................................................106 Leucrocuta, key to species................................................................................................108 Maccaffertium, key to species...........................................................................................110 Macdunnoa, key to species...............................................................................................118 Nixe, key to species...........................................................................................................118 Rhithrogena, key to species..............................................................................................120 Stenacron, key to species..................................................................................................124 ISONYCHIIDAE..........................................................................................................................126 Isonychia, key to species...................................................................................................126 LEPTOHYPHIDAE, key to genera and species...........................................................................130 Tricorythodes, key to species............................................................................................132 LEPTOPHLEBIIDAE, key to genera and species........................................................................136 Habrophlebiodes, key to species......................................................................................138 Leptophlebia, key to species.............................................................................................138 Paraleptophlebia, key to species......................................................................................140 METRETOPODIDAE..................................................................................................................146 Siphloplecton, key to species............................................................................................146 NEOEPHEMERIDAE..................................................................................................................148 Neoephemera, key to species............................................................................................148 OLIGONEURIIDAE....................................................................................................................150 Homoeoneuria, key to species..........................................................................................150 POLYMITARCYIDAE, key to genera and species......................................................................150 Ephoron, key to species....................................................................................................150 POTAMANTHIDAE....................................................................................................................152 Anthopotamus, key to species...........................................................................................152 SIPHLONURIDAE......................................................................................................................154 Siphlonurus, key to species...............................................................................................154 REFERENCES cited.....................................................................................................................156 Chapter 3: PLECOPTERA, B.P. Stark...................................................................................................161 INTRODUCTION and major characteristics..............................................................................161 LIST of families, genera, and species..........................................................................................168 KEY to families (some genera and species).................................................................................180 PLECOPTERA, key to families...................................................................................................180 CAPNIIDAE, key to genera and species.....................................................................................182 CHLOROPERLIDAE, key to genera and species......................................................................184 LEUCTRIDAE, key to genera and species.................................................................................184 NEMOURIDAE, key to genera and species...............................................................................186 PELTOPERLIDAE, key to genera and species..........................................................................188 PERLIDAE, key to genera and species.......................................................................................188 PERLODIDAE, key to genera and species.................................................................................192

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TAENIOPTERYGIDAE, key to genera and species..................................................................196 CAPNIIDAE................................................................................................................................198 Allocapnia, key to species.................................................................................................198 CHLOROPERLIDAE.................................................................................................................202 Alloperla, key to species...................................................................................................202 Haploperla, key to species................................................................................................204 Sweltsa, key to species......................................................................................................206 LEUCTRIDAE............................................................................................................................208 Leuctra, key to species......................................................................................................208 Megaleuctra, key to species..............................................................................................210 Zealeuctra, key to species.................................................................................................210 NEMOURIDAE..........................................................................................................................212 Amphinemura, key to species...........................................................................................212 Ostrocerca, key to species................................................................................................212 Prostoia, key to species....................................................................................................214 Soyedina, key to species...................................................................................................214 Zapada, key to species......................................................................................................214 PERLIDAE..................................................................................................................................216 Acroneuria, key to species................................................................................................216 Agnetina, key to species....................................................................................................218 Beloneuria, key to species................................................................................................220 Hansonoperla, key to species...........................................................................................220 Neoperla, key to species...................................................................................................220 Paragnetina, key to species..............................................................................................222 Perlesta, key to species.....................................................................................................224 Perlinella, key to species..................................................................................................226 PERLODIDAE............................................................................................................................226 Cultus, key to species........................................................................................................226 Diploperla, key to species.................................................................................................226 Helopicus, key to species..................................................................................................228 Hydroperla, key to species................................................................................................228 Isogenoides, key to species...............................................................................................230 Isoperla, key to species.....................................................................................................230 Remenus, key to species....................................................................................................236 Yugus, key to species.........................................................................................................236 PTERONARCYIDAE................................................................................................................238 Pteronarcys, key to species...............................................................................................238 TAENIOPTERYGIDAE.............................................................................................................238 Strophopteryx, key to species............................................................................................238 Taeniopteryx, key to species.............................................................................................240 REFERENCES cited.....................................................................................................................244

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Chapter 4. TRICHOPTERA, J.C. Morse, R.W. Holzenthal, and O. Yadamsuren..................................248 INTRODUCTION and major characteristics..............................................................................248 LIST of families, genera, and species..........................................................................................253 KEY to families (some genera and species).................................................................................284 APATANIIDAE, key to genera and species..................................................................................294 Manophylax, key to species..............................................................................................294 BRACHYCENTRIDAE, key to genera and species....................................................................296 Brachycentrus, key to species...........................................................................................296 Micrasema, key to species................................................................................................300 CALAMOCERATIDAE, key to genera and species....................................................................304 DIPSEUDOPSIDAE.....................................................................................................................306 Phylocentropus, key to species.........................................................................................306 GLOSSOSOMATIDAE, key to genera and species.....................................................................306 Agapetus, species..............................................................................................................308 Glossosoma, key to species...............................................................................................308 Protoptila, species.............................................................................................................310 GOERIDAE, key to genera...........................................................................................................310 Goera, key to species........................................................................................................310 Goerita, key to species......................................................................................................312 HELICOPSYCHIDAE.................................................................................................................312 Helicopsyche, key to species.............................................................................................312 HYDROPSYCHIDAE, key to genera and species.......................................................................312 Cheumatopsyche, key to species.......................................................................................318 Diplectrona, key to species...............................................................................................322 Homoplectra, key to species.............................................................................................324 Hydropsyche, key to species.............................................................................................324 Macrostemum, key to species...........................................................................................344 Parapsyche, key to species...............................................................................................344 HYDROPTILIDAE, key to genera and species...........................................................................346 Hydroptila, species............................................................................................................350 Ithytrichia, species............................................................................................................352 Mayatrichia, species.........................................................................................................352 Neotrichia, species............................................................................................................352 Ochrotrichia, species........................................................................................................352 Orthotrichia, species.........................................................................................................353 Oxyethira, species.............................................................................................................353 Stactobiella, species..........................................................................................................353 LEPIDOSTOMATIDAE, key to genera.......................................................................................354 Lepidostoma, species........................................................................................................354 Theliopsyche, species........................................................................................................354 LEPTOCERIDAE, key to genera and species..............................................................................354 Ceraclea, key to species...................................................................................................358 Mystacides, species...........................................................................................................366 Nectopsyche, key to species..............................................................................................368 Oecetis, key to species......................................................................................................370 Setodes, key to species......................................................................................................378 Triaenodes, key to species................................................................................................380

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LIMNEPHILIDAE, key to genera and species.............................................................................388 Anabolia, species..............................................................................................................390 Frenesia, key to species....................................................................................................392 Ironoquia, key to species..................................................................................................392 Limnephilus, key to species..............................................................................................392 Platycentropus, species.....................................................................................................394 Pseudostenophylax, species..............................................................................................394 Pycnopsyche, key to species.............................................................................................394 MOLANNIDAE...........................................................................................................................398 Molanna, key to species....................................................................................................398 ODONTOCERIDAE, key to genera and species.........................................................................398 Psilotreta, key to species..................................................................................................400 PHILOPOTAMIDAE, key to genera and species.........................................................................404 Chimarra, key to species..................................................................................................406 Wormaldia, key to species................................................................................................408 PHRYGANEIDAE, key to genera and species.............................................................................408 Agrypnia, key to species...................................................................................................410 Banksiola, species.............................................................................................................410 Oligostomis, key to species...............................................................................................410 Phryganea, species...........................................................................................................410 Ptilostomis, species...........................................................................................................410 POLYCENTROPODIDAE, key to genera and species................................................................412 Cernotina, species.............................................................................................................414 Holocentropus, key to species..........................................................................................414 Neureclipsis, species.........................................................................................................414 Nyctiophylax, key to species.............................................................................................414 Plectrocnemia, species......................................................................................................414 Polycentropus, species......................................................................................................416 PSYCHOMYIIDAE, key to genera and species...........................................................................416 Psychomyia, key to species...............................................................................................418 RHYACOPHILIDAE....................................................................................................................418 Rhyacophila, key to species..............................................................................................418 SERICOSTOMATIDAE, key to genera and species....................................................................424 Agarodes, key to species...................................................................................................424 THREMMATIDAE.......................................................................................................................428 Neophylax, key to species.................................................................................................428 REFERENCES cited.....................................................................................................................434 INDEX..........................................................................................................................................443

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Chapter 1

INTRODUCTION

John C. Morse, Bill P. Stark, and Luke M. Jacobus The unusually high density of plant and animal species in the southeastern United States has contributed to the excellent quality of life in this region for humans and other organisms. Diverse biotic and abiotic factors here combine to create the remarkable variety of habitats and niches that these species occupy (Bryer et al., 2000). The highest species richness in North America is found in the Southeast, with freshwater species especially well represented. Our region’s freshwater habitats, however, are under increasing anthropogenic pressures (Benz & Collins, 1997; Chaplin et al., 2000). Southeastern ecoregions, especially the Southern Appalachian Mountains and Florida Panhandle, are well known for their species richness, including the occurrences of many rare species (Morse et al., 1997; Chaplin et al., 2000; Parker et al., 2007; Graves & Ward, 2011). Interest in the biology of these many species has prompted research on their habits, habitat requirements, trophic relationships, physiologies, life histories, and other traits. We are becoming more aware of the differences among species with respect to their physiologies and functional roles in the environment (e.g., Poteat & Buchwalter, 2014) and how those differences reflect ecosystem diversity and health at the species level of taxonomy. A better understanding of the environmental constraints on our biota has been accompanied by an appreciation of its potential for indicating environmental changes, including those caused by human activity, both locally and elsewhere in the biosphere. Concomitant with an increasing use of freshwater macroinvertebrates for monitoring water quality (e.g., Barbour et al., 1999) is the need for more-reliable and efficient sampling and analytical methods. Among research goals is a desire to recognize taxa that are most informative for changed environmental conditions and to determine the most appropriate taxonomic level for assessing those conditions. The mayflies (Ephemeroptera), stoneflies (Plecoptera), and caddisflies (Trichoptera) rank high among environmentally sensitive freshwater macroinvertebrates. Immature stages of these insect orders live in the water and their adults emerge from the water into a terrestrial environment to mate and lay eggs. As a consequence, the ecological integrity of both the freshwater and the surrounding riparian habitat is important for them to flourish. Mayflies have egg and larval (nymph) life history stages living in fastflowing to standing water; subimaginal and imaginal (adult) stages are found in nearby shoreside habitats (Edmunds et al., 1976; Waltz & Burian, 2008). Stoneflies have egg and larval (nymph) stages in the water (typically streams), with adults in nearby habitats (Stewart & Stark, 2002, 2008), typically along the water’s edge. Caddisflies are holometabolous insects with eggs, larvae, and pupae thriving in many types of surface-water habitats; adults occur mostly in riparian habitats (Wiggins, 1996, 2004). Collectively, these three orders of insects often are called “EPT” (Lenat & Penrose, 1996). The density and diversity of EPT in freshwater habitats, especially smaller-order streams, are often high, contributing greatly to the capacity of these ecosystems to process available nutrients and in turn to transfer nutrients to a wide range of invertebrate and vertebrate predators and terrestrial ecosystems. Taxonomic resolution of EPT taxa should consider several issues. For example, does species-level resolution improve the precision and accuracy of biological assessments over family- or genus-level resolution and, if so, is that improvement worth the effort and cost (Lenat & Resh, 2001)? Answers to those questions depend on the ability to identify the biota to the species level. Associating a distinctive name with a taxon is a critical step for the operation of the international system of biological information storage and retrieval. Information about a taxon’s morphology and biology—its unique set of structural and functional features—is “stored” in many different printed and digital resources using the scientific name of the taxon as a key word. This key word allows information to be “retrieved” from those resources. General references for applying names accurately to the orders, families, and genera of aquatic

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insects in North America include the book by Merritt et al. (2008a) and publications listed in Chapter 1 of that resource. For more than 30 years, the principal single reference for lowest-possible-taxonomiclevel identification (species-level where possible) of freshwater insects in North and South Carolina and surrounding regions has been the work of Brigham et al. (1982). That volume, commonly called “the Duke manual,” was sponsored by Duke Power Company to facilitate and standardize identifications by Duke’s biologists monitoring the water quality of effluent and its effects downstream of the company’s facilities. That volume included not only illustrated diagnostic keys, but also remarkably complete syntheses of the known biological characteristics of the taxa, syntheses that are outside the scope of the present volume. The book by Brigham et al. (1982) was published in two formats, a 3-ring binder and a 3-post binder, which allowed pages to be inserted and removed, anticipating opportunities to update the work. Sufficient copies were produced that it became a standard reference for benthologists throughout the southeastern United States. Soon after the manual’s appearance, Duke Power Company (later Duke Energy) was honored by the National Wildlife Federation with its Corporate Leadership Award (1984 and 2000) for the company’s progressive attention to environmental quality in the region, no doubt considering the value of “the Duke Manual” in its decision. Researchers over the past 35 years have associated and described larvae of many more species of EPT than were known in 1982. The main purpose of the present publication, therefore, is to facilitate up-todate species-level identification of mayflies, stoneflies, and caddisflies, to the extent currently possible, not only for the Carolinas, but for the southeastern United States. “Southeastern” here will refer to the states of the U.S. Environmental Protection Agency Region IV: Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, South Carolina, and Tennessee. Inclusion of a species in one of the taxonomic keys does not necessarily indicate its confirmed presence in all or any of these southeastern states. Several species from neighboring states are included to facilitate their identification should they be discovered in the eightstate study region. A few extralimital species also are included to emphasize the diagnostic characters of the local species in cases where we anticipate discovery of several new species, due to a poor state of knowledge for that group in the Southeast. We hope similar identification guides can be made available soon for other major freshwater taxa and for other regions of North America. Since 1982, research has expanded our knowledge of the diversity of freshwater insect and oligochaete species of the southeastern United States, both in terms of discovering, recognizing, describing, and naming species new to science and in terms of discovering and describing previously unknown life history stages, mainly larvae, of species with formally established names. These descriptions are scattered in many scholarly works, a majority of which are out of print, difficult to find, or otherwise inaccessible for modern benthologists. For example, many descriptions and identification tools languish in theses, dissertations, workshop manuals, and unpublished manuscripts or private research notes. The information they contain has never been formally published. We have compiled as much of that mostly inaccessible knowledge as possible to make it available to a broader audience. In addition, several species have been collected and reared just for this study; although formal descriptions have not been published, we include the species in the keys after thorough study. Our publication thus synthesizes the published and unpublished results of otherwise disparate and often unavailable taxonomic research. Our keys, in many cases, separate identifiable morphs for which we assign provisional names for tracking purposes. Some of these are new species that await formal scientific description; some are simply distinct ecological varieties of species; and some are forms that remain unassociated with their namebearing metamorphic stages. In a few cases, especially where more research on a genus or species-group is needed, we refer to specific epithets that recently have been considered junior synonyms of other older names, but which historically have been associated with a particular morphotype, in anticipation of their revalidation in light of new evidence (e.g., Webb et al., 2012). Another major purpose of this work is to indicate data gaps where species-level taxonomic research should be undertaken. In many cases, some basic research is needed, especially for stoneflies and caddisflies, to associate yet-unidentified life history stages with identifiable adults and then to discover

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and describe diagnostic differences useful for species-level identification. Specifically, we encourage users of this resource to make the associations by one or more of at least three approaches: (1) rearing poorly known taxa with methods such as those by Jacobus & McCafferty (2004) for mayflies or those referenced by Merritt et al. (2008b) for all three orders, (2) collecting and preserving mature caddisfly pupae with their shed larval sclerites (“the metamorphotype method” of Milne, 1938; Wiggins, 1996), and (3) comparing sequences of DNA, especially those of mitochondrial cytochrome C oxidase subunit I (mtCOI), for vouchered specimens of all identifiable and unidentifiable life history stages (e.g., Zhou et al., 2007a, 2007b, 2011; Webb et al., 2012; Ruiter et al., 2013). Searches for diagnostic characters are best accomplished by direct comparison of the last-instar larvae of different species that have been associated confidently with one or more of these methods. Their diagnostic characters will be those (1) that are heritable, (2) that sort unambiguously without intermediates, and (3) that are correlated with similarly heritable and distinctive features not likely to be functionally related or genetically linked. Written descriptions of resulting diagnostic characters should be accompanied by high-quality photographs and/or drawings. This research also will make available more characters for inferring the phylogenetic relationships of EPT taxa, facilitating use of the relationships for predictive research on other, correlated biological features of interest. We anticipate that this volume will be useful to both amateur and professional benthologists desiring to know the species of southeastern EPT. Although technical language might discourage casual users, it generally is necessary at this level of taxonomic refinement. The diagnostic characters mentioned in the keys typically are expressed only or most clearly in late or final instar larvae; in some cases, these characters are relatively subtle. For mayflies and stoneflies, the final instar larva is most readily recognized by dark or black wingpads containing developing subimaginal or adult wings following apolysis but preceding ecdysis. For caddisflies, the final instar larva is recognized by reference to a complete set of larval instars (most caddisflies have 5) or by capture of larvae that have prepared pupal shelters and assumed a typical prepupal posture—a shortened and thickened body shape with head tucked against the prosternum and legs folded against the thorax. More-subtle species-level differences may be detected best by reference to long series of specimens and side-by-side comparisons of the user’s different hypothesized species. We have attempted to provide quality illustrations of the various characters currently and historically used to separate species. Keys for many of the taxa in this volume are introduced here for the first time, so that they have not been tested by many users and the range of possible variations has not been fully explored. Readers who discover specimens that are inconsistent with the keys are encouraged to share their observations with the authors. Consistent with Brigham et al.’s (1982) original intent to provide updates for their book, we anticipate future revisions of this volume. Inasmuch as ours is a tax-supported publication and therefore largely in the public domain, we most likely will provide updates via the worldwide web, subject to consent by copyright holders. Important supplementary resources for this volume are the sets of frequently updated manuals provided by state regulatory agencies in the Southeast, such as those of the Florida Department of Environmental Protection Bureau of Laboratories (http://www.dep.state.fl.us/labs/cgi-bin/sbio/keys.asp) and the North Carolina Division of Water Quality, Biological Assessment Branch (http://ncdenr.gov/web/ wq/taxonmanual). Although specific for these states, much of the fauna treated in these manuals occurs throughout the Southeast. The production of this volume was supported by funding from a United States Environmental Protection Agency (USEPA) Section 106(A) supplemental monitoring grant, through its Water Protection Division, to the South Carolina Department of Health and Environmental Control (SCDHEC). We are grateful to these agencies and especially Dr. James B. Glover, Manager of the Aquatic Biology Section, SCDHEC, Bureau of Water, for their encouragement and support. Dr. Al Wheeler, Editor for the Biota of South Carolina series, graciously helped to edit the book. Ms. Virginia Winn assembled final copy in Adobe® InDesign® 5.5 software. We thank Ms. Charlene Mayfield, Director of PSA Publishing, for her continuing support of the Biota of South Carolina series and her help to produce and distribute this volume. Acknowledgements for contributors to specific parts of the book are noted in the respective chapters.

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Introduction

References Cited Barbour, M.T., J. Gerritsen, B.D. Snyder, & J.B. Stribling. 1999. Rapid Bioassessment Protocols for Use in Streams and Wadeable Rivers: Periphyton, Benthic Macroinvertebrates and Fish, Second Edition. EPA 841-B-99-002. U.S. Environmental Protection Agency, Office of Water, Washington, D.C. Available from (Accessed 07 October 2014). Benz, G.W., & D.E. Collins. 1997. Preface. In: Benz, G.W., & D.E. Collins (Eds.), Aquatic Fauna in Peril: The Southeastern Perspective. Southeast Aquatic Research Institute Special Publication 1, Lenz Design & Communications, Decatur, Georgia, USA, pp. xi–xv. Brigham, A.R., W.U. Brigham, & A. Gnilka (Eds.) 1982. Aquatic Insects and Oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. Bryer, M.T., K. Maybury, J.S. Adams, & D.H. Grossman. 2000. More than the sum of the parts: Diversity and status of ecological Systems. In: Stein, B.A., L.S. Kutner, & J.S. Adams (Eds.), Precious Heritage: The Status of Biodiversity in the United States. Oxford University Press, Oxford, United Kingdom, pp. 201–238. Chaplin, S.J., R.A. Gerrard, H.M. Watson, L.L. Master, & S.R. Flack. 2000. The geography of imperilment: Targeting conservation toward critical biodiversity areas. In: Stein, B.A., L.S. Kutner, & J.S. Adams (Eds.), Precious Heritage: The Status of Biodiversity in the United States. Oxford University Press, Oxford, United Kingdom, pp. 159–199. Edmunds, G.F., Jr., S.L. Jensen, & L. Berner. 1976. The Mayflies of North and Central America. University of Minnesota Press, Minneapolis, Minnesota, USA. Graves, P.H., III, & G.M. Ward. 2011. Mayfly and stonefly distribution in the mainstem Cahaba River, Alabama. Southeastern Naturalist 10: 477–488. Jacobus, L.M., & W.P. McCafferty. 2004. Contribution to the morphology and descriptive biology of Caurinella idahoensis (Ephemeroptera: Ephemerellidae). Western North American Naturalist 64: 101–108. Lenat, D.R., & D.L. Penrose. 1996. History of the EPT taxa richness metric. Bulletin of the North American Benthological Society 13: 305–306. Lenat, D.R., & V.H. Resh. 2001. Taxonomy and stream ecology—The benefits of genus- and species-level identifications. Journal of the North American Benthological Society 20: 287-298. Merritt, R.W., K.W. Cummins, & M.B. Berg (Eds.) 2008a. An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa, USA. Merritt, R.W., V.H. Resh, K.W. Cummins, & D.P. Batzer. 2008b. Sampling aquatic insects: Collection devices, statistical considerations, and rearing procedures. In: Merritt, R.W., K.W. Cummins, & M.B. Berg (Eds.) 2008a. An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa, USA, pp. 1–6. Milne, M.J. 1938. The “metamorphotype method” in Trichoptera. Journal of the New York Entomological Society 46: 435–437. Morse, J.C., B.P. Stark, W.P. McCafferty, & K.J. Tennessen. 1997. Southern Appalachian and southeastern streams at risk: Implications for mayflies, dragonflies and damselflies, stoneflies, and caddisflies. In: Benz, G.W., & D.E. Collins (Eds.), Aquatic Fauna in Peril: The Southeastern Perspective. Southeast Aquatic Research Institute Special Publication 1, Lenz Design & Communications, Decatur, Georgia, USA, pp. 17–42. Parker, C.R., O.S. Flint, Jr., L.M. Jacobus, B.C. Kondratieff, W.P. McCafferty, & J.C. Morse. 2007. Ephemeroptera, Plecoptera, Megaloptera, and Trichoptera of Great Smoky Mountains National Park. Southeastern Naturalist, Special Issue 1: 159–174. Poteat, M.D., & D.B. Buchwalter. 2014. Phylogeny and size differentially influence dissolved Cd and Zn bioaccumulation parameters among closely related aquatic insects. Environmental Science & Technology 48: 5274–5281.

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Ruiter, D.E., E.E. Boyle, & X. Zhou. 2013. DNA barcoding facilitates associations and diagnoses for Trichoptera larvae of the Churchill (Manitoba, Canada) area. BMC Ecology 13: 5 (39 pp.) Available from (Accessed 13 October 2014). Stewart, K.W., & B.P. Stark. 2002. Nymphs of North American Stonefly Genera (Plecoptera), Second Edition. Caddis Press, Columbus, Ohio, USA. Stewart, K.W., & B.P. Stark. 2008. Plecoptera. In: Merritt, R.W., K.W. Cummins, & M.B. Berg (Eds.) 2008a. An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa, USA, pp. 311–384. Waltz, R.D., & S.K. Burian. 2008, Ephemeroptera. Merritt, R.W., K.W. Cummins, & M.B. Berg (Eds.) 2008a. An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa, USA, pp. 181–236. Webb, J.M., L.M. Jacobus, D.H. Funk, X. Zhou, B. Kondratieff, C.J. Geraci, R.E. DeWalt, D. Baird, B. Richard, I. Phillips, & P.D.N. Hebert. 2012. A DNA Barcode Library for North American Ephemeroptera: Progress and Prospects. PLoS ONE 7(5): e38063. doi: 10.1371/journal.pone.0038063. Available from

(Accessed 13 October 2014). Wiggins, G.B. 1996. Larvae of the North American Caddisfly Genera (Trichoptera), Second Edition. University of Toronto Press, Toronto, Ontario, Canada. Wiggins, G.B. 2004. Caddisflies: The Underwater Architects. University of Toronto Press, Toronto, Ontario, Canada. Zhou, X., K.M. Kjer, & J.C. Morse. 2007a. Associating larvae and adults of Chinese Hydropsychidae caddisflies (Insecta: Trichoptera) using DNA sequences. Journal of the North American Benthological Society 26: 719–742. Zhou, X., K.M. Kjer, & J.C. Morse. 2007b. An introduction to the species delimitation, larval-adult association of Chinese Hydropsychidae using independent DNA sequences and adult morphology. In: Bueno-Soria, J., R. Barba-Alvarez, & B. Armitage (Eds.), Proceedings of the XIIth International Symposium on Trichoptera, June 18–22, 2006. Caddis Press, Columbus, Ohio, U.S.A., pp. 355–368. Zhou, X., J.L. Robinson, C.J. Geraci, C.R. Parker, O.S. Flint, Jr., D. Etnier, D. Ruiter, R.E. DeWalt, L.M. Jacobus, & P.D.N. Hebert. 2011. Accelerated construction of a regional DNA barcode reference library: Caddisflies (Trichoptera) in the Great Smoky Mountains National Park. Journal of the North American Benthological Society 30: 131–162 + 3 suppl.

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Chapter 2 ~ Key to Mayflies

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Chapter 2

EPHEMEROPTERA

W. Patrick McCafferty, Luke M. Jacobus, Arwin V. Provonsha, and Nick A. Wiersema With respect to mayflies, the southeastern United States is one of the richest and most diverse area of such size in the world. The approximately 300 species in the area (McCafferty et al., 2010; Webb et al., 2012) represent more than half of those known from the United States, very nearly half of those known from North America (McCafferty & Jacobus, 2015) and about 10% of the world fauna (Barber-James et al., 2008, 2013; Kluge, 2015). Our key is intended primarily as a guide to the identification of the aquatic, larval life stage of mayfly families, genera, and species known from Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, South Carolina, and Tennessee—states that make up USEPA Region 4. A total of 287 nominal species and another 15 that apparently represent undescribed species (indicated within a genus by informal lettering or numbering) are addressed in the key; these totals include some species from nearby areas that we consider likely to be found in the coverage area. We also discuss a few taxa informally. Nominal taxa of families, genera, and species addressed in the key are listed in Table 2.1. The table includes species known as larvae that have been reported from Region 4, those from adjacent areas that are known as larvae and eventually might be found in Region 4 (noted by a plus sign), twelve for which larvae are unknown (indicated by a dagger), and three presumed to be extinct that also have never been known as larvae (indicated by a double dagger). These latter 15 species necessarily are excluded from the key, except for limited parenthetical remarks. The twelve extant species known to occur in Region 4 states but are unknown in the larval stage include Apobaetis futilis (McDunnough), Asioplax texana (Traver), Isonychia berneri Kondratieff & Voshell, Iswaeon rubrolaterale (McDunnough), Plauditus veteris (McDunnough), Procloeon intermediale (McDunnough), Procloeon simile (McDunnough), Rhithrogena anomala McDunnough, R. rubicunda Traver, Siphlonurus decorus Traver, Siphloplecton costalense Spieth, and Siphloplecton simile Berner. These species represent only about 4% of those in Region 4, and this low percentage illustrates the extreme degree to which our knowledge of larval Ephemeroptera in North America has improved recently, which now makes such an extensive key possible. We note that we have seen one male adult specimen from mountainous western North Carolina that appears to belong to the Neotropical and western Nearctic baetid genus Cloeodes (Salles et al., 2015), but this genus is not included in the key because the larva of the North Carolina species is not known, and generic attribution is tentative. We also have seen one larval specimen from a swamp stream in coastal eastern North Carolina that may represent a new genus from the Baetodes complex of genera (recently reviewed by Nieto, 2016), but this unnamed taxon is excluded from the key because additional study is needed to determine its identity and diagnostic features. Three of the four recent mayfly species presumed to be extinct in North America (McCafferty, 2001) were collected from the Southeast. They include Isonychia diversa Traver, Pentagenia robusta McDunnough, and Siphlonurus luridipennis (Burmeister). These poorly known species are known only from winged stages. Isonychia diversa was collected late June and July 1916 from near Knoxville, Tennessee (Traver, 1934; Kondratieff & Voshell, 1984). We are uncertain about the date of collection of the other two species. Pentagenia robusta was collected from Cincinnati, Ohio (McDunnough, 1926) and is presumed to have emerged from the Ohio River, which forms Ohio’s southern border with Kentucky; thus it is included in this account. Siphlonurus luridipennis was taken from “Carolina.” McCafferty et al. (2010) listed the extant species known from the Southeast—the Region 4 states mentioned above plus Arkansas, Louisiana, Virginia, and West Virginia—and the exact southeastern states from which they are known based on published records. Additional species from Kentucky were

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

reported by McCafferty (2011b). We do not include complete state and provincial distribution data for the mayfly species treated herein. These data have not yet been compiled for all species, and doing so is beyond the time limitations of the current work. Some taxonomic changes among baetid genera were proposed by Jacobus & Wiersema (2014) subsequent to the listings provided by McCafferty et al. (2010) and the new records provided by McCafferty (2011b). Southeastern records of Arthropleidae have been questioned (Jacobus 2013), but the single North American species of this family is included in this work. For those unfamiliar with recent names, generic combinations or synonyms for the mayflies treated herein, a complete history and continually updated name equivalencies for North American Ephemeroptera can be found at the Mayfly Central Website (McCafferty, 1996; McCafferty & Jacobus, 2015). Keys to families, genera, and species are sequenced alphabetically. In this way, users who may already know the family or the genus of the specimens they are working with may proceed to the appropriate section. For the few families that are monospecific in the coverage area, family, genus, and species are taken up only in the key to families. Similarly, for those genera that are monospecific, the genus and species are taken up only within the respective genus key. All other species are keyed within the multispecific keys for a particular genus. Couplets and figure numbers are continuous throughout the mayfly keys. Figures are positioned near the couplets where they are referenced, usually on the opposing page. Morphological characters used in the key, when possible, are those that are most user-friendly and consistent in populations across the entire coverage area. Other characters may be useful at local levels. Any specialized morphology or technical terms used in key sections are explained at the initial point of usage within the appropriate couplets or as appended notes to the couplets. Key characteristics vary considerably among families and genera, and they are taken up at their point of usage. Such self-contained couplets make any additional explanation or introduction to general morphology in this preface unnecessary and, more importantly, eliminate the need to move away from the couplet at hand to understand its usage. In addition to explaining characters, we suggest ideal relative magnification for viewing particular characters and note when it is advisable or necessary to make slide mounts. Other items of information may be included within key couplets or appended as notes when deemed useful for identifying southeastern mayflies. For example, species with restricted distributions will be noted, and at any taxonomic level, some general habitat information may be appended if it is specific enough to facilitate identification. The reported distribution of species not currently known from the area of coverage will always be given, as well as notes on species unknown as larvae. For some portions of the key, citations to more detailed references are given, but other portions are strictly original to this work and therefore have no such citations. The prevailing generic and species taxonomy is used in this key (McCafferty & Jacobus, 2015). We point out certain possibly invalid species that cannot be keyed definitively and require further study. Particularly in the genera Drunella and Ephemerella (Ephemerellidae), certain morphological variants that may or may not prove to be valid species will key to the species names that have historically been associated with them even if they currently are considered synonyms. This anticipates any possible revalidation of certain names in light of other synonyms in these genera having recently been shown to be valid, genetically distinct, cryptic species (e.g., Funk et al., 2008b; Webb et al., 2012: Supporting Information Text S1). Limited discussions of these nomenclatural issues are included as remarks within the key. The key is based on the synthesis and application of diagnostic data and other characterization associated with the applicable mayfly taxa and on new diagnostic and comparative data that have been generated as a result of research for the key. Organization of the key also is based on what has been found to be the most logical sequence of couplets and characters within couplets, with ease of use more important than other considerations. Areas of the key that are weak or possibly unreliable remain, at least in part, because of the lack of consistent comparative data associated with certain species. These areas are few in

Chapter 2 ~ Key to Mayflies

Page 17

number and are pointed out with cautionary statements appended to the key. Users should be aware that species of the genera Leucrocuta (Heptageniidae), Siphloplecton (Metretopodidae), and Tricorythodes (Leptohyphidae) may be particularly troublesome even though their larvae are generally known. Most species of Leucrocuta cannot be reliably keyed to species because some larvae have been erroneously associated with named adults, and comprehensive comparisons of reared or otherwise associated larvae have not been completed. In the case of Siphloplecton, morphological characters for separating several species are unreliable, suggesting the species taxonomy is questionable. In the case of Tricorythodes, a generic revision is needed, because it is not known whether certain potentially diagnostic characteristics might vary intraspecifically, nor which species are valid, nor which species might actually be comprised of complexes of cryptic species. There also remain some pairs of well-known species (at least as adults) that are difficult, if not impossible, to distinguish consistently as larvae, such as Heptagenia dolosa vs. H. marginalis, Hexagenia bilineata vs. H. limbata, Isonychia bicolor vs. I. rufa, and Leptohplebia cupida vs. L. nebulosa (McCafferty, 1975; Kondratieff & Voshell, 1984; Burian, 2001; Jacobus & Webb, 2013). Some complex larval characteristics may work some of the time for such pairs, but have not been demonstrated to be totally reliable, especially across large geographic ranges. Mature male larvae sometimes are required to ascertain diagnostic developing adult characters. In all difficult species pairs or groups, we advise that larvae be reared to the adult stage if a species-level identification is critical, as male adults in these particular groups generally are more reliable for assigning species names. Jacobus & McCafferty (2004a) described the rearing technique we used in field studies for this chapter. The introductory chapter of this book contains additional discussion about methodologies for associating stages of mayflies. Several of the figures are originals by Arwin V. Provonsha presented here for the first time. The others are from various sources, but most are borrowed or variously modified from works involving the authors of this chapter. Specific sources of the latter are not given in the figure legends but are based on the following: Bae & McCafferty (1991), Bednarik & McCafferty (1979), Jacobus (2006), Jacobus & McCafferty (2000, 2004b, 2006, 2008), Lugo-Ortiz & McCafferty (1998a, 1998b), McCafferty (1971, 1975, 1977a, 1977b, 1981a, 1981b, 1985, 1990, 1991, 2010), McCafferty & Lenat (2003), McCafferty & Provonsha (1986), McCafferty & Waltz (1995, 1998), McCafferty et al. (2005, 2009), Morihara & McCafferty (1979a, 1979b, 1979c), Provonsha (1990), Provonsha & McCafferty (1982, 2006), Randolph & McCafferty (1996), Sun & McCafferty (2008), Waltz & McCafferty (1989), Wang & McCafferty (1996), Webb & McCafferty (2006, 2008), Wiersema (1999, 2000), Wiersema & Long (2000), and Wiersema & McCafferty (2000, 2005). We thank Michael Cole for the dorsal habitus illustration of Penelomax. Sources of other figures that do not include authors of this chapter are given as part of the figure legend. To the authors of journal papers or theses containing figures and diagnostic information adopted or modified for use herein, we express our sincere thanks. Virginia Winn contributed many hours and other personal resources towards this project and final figure production, in particular. George Towers, Interim Head of IUPUC Division of Science, is acknowledged for graciously allocating part of the funds necessary to re-create the final figures. We additionally thank Andy Usher for his assistance with figure preparations prior to peer-review of the chapter and Tammy Morton for solving problems associated with the final format of Table 2.1. Dave Lenat, Steven Beaty, Eric Fleek, Victor Holland, John Morse, and students enrolled in the 2015 Highlands Biological Station EPT courses provided critical reviews and trial usages of the key that led to improvements and corrections. Alfred Wheeler improved the clarity and quality of language in this chapter.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Table 2.1. Alphabetical list of families, genera, and species or varieties of Ephemeroptera known or suspected from Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, South Carolina & Tennessee (USEPA Region 4). For a listing of state distributions from within this region for each species, see McCafferty et al. (2010), with additional records given by McCafferty (2011b). + = Species known from nearby states, suspected to occur in Region 4 and addressed in key, either in a couplet or anecdotally. † = Species unknown as larva. ‡ = Species unknown as larva and presumed to be extinct.

Acanthametropodidae

Acanthametropus pecatonica (Burks)

Ameletidae +

Ameletus browni McDunnough Ameletus cryptostimulus Carle Ameletus lineatus Traver Ameletus ludens Needham Ameletus tarteri Burrows Ameletus tertius McDunnough Ameletus sp. 1 Ameletus sp. 2 Ameletus sp. 3

Arthropleidae

Arthroplea bipunctata (McDunnough)

Baetidae

Acentrella alachua (Berner) Acentrella barbarae Jacobus & McCafferty Acentrella nadineae McCafferty, Waltz & Webb Acentrella parvula (McDunnough) Acentrella turbida (McDunnough) Acerpenna macdunnoughi (Ide) Acerpenna pygmaea (Hagen)

Anafroptilum album (McDunnough) Anafroptilum bifurcatum (McDunnough) Anafroptilum minor (McDunnough) Anafroptilum ozarkense (Wiersema & Burian) + Anafroptilum semirufum (McDunnough) Anafroptilum sp. A



Baetidae continued

Apobaetis etowah (Traver) † Apobaetis futilis (McDunnough) Apobaetis sp. A





Baetis brunneicolor McDunnough Baetis flavistriga McDunnough Baetis intercalaris McDunnough Baetis phoebus McDunnough Baetis pluto McDunnough Baetis tricaudatus Dodd

Baetopus trishae Waltz Baetopus sp. A

Barbaetis benfieldi Kennedy

Callibaetis floridanus Banks Callibaetis fluctuans (Walsh) Callibaetis pretiosus Banks

Camelobaetidius musseri (Traver & Edmunds) Cloeon dipterum (Linnaeus) Diphetor hageni (Eaton) +

Fallceon quilleri (Dodds)

Heterocloeon amplum (Traver) Heterocloeon berneri (MüllerLiebenau) Heterocloeon curiosum (McDunnough) Heterocloeon frivolum (McDunnough) Heterocloeon grande (Wiersema & Long)

Chapter 2 ~ Key to Mayflies

Baetidae continued

Heterocloeon petersi (MüllerLiebenau) Heterocloeon sp. A Heterocloeon sp. B + Heterocloeon sp. C + Heterocloeon sp. D

Iswaeon anoka (Daggy) Iswaeon davidi (Waltz & McCafferty) † Iswaeon rubrolaterale (McDunnough)

Labiobaetis dardanus (McDunnough) Labiobaetis ephippiatus (Traver) Labiobaetis frondalis (McDunnough) + Labiobaetis longipalpus (Morihara & McCafferty) Labiobaetis propinquus (Walsh) Neocloeon alamance Traver Neocloeon triangulifer (McDunnough)

Paracloeodes fleeki McCafferty & Lenat Paracloeodes minutus (Daggy)

Plauditus bimaculatus (Berner) Plauditus cestus (Provonsha & McCafferty) Plauditus cingulatus (McDunnough) Plauditus dubius (Walsh) Plauditus gloveri McCafferty & Waltz Plauditus punctiventris (McDunnough) + Plauditus texanus Wiersema † Plauditus veteris (McDunnough) Plauditus virilis (McDunnough)

Procloeon fragile (McDunnough) † Procloeon intermediale (McDunnough) Procloeon nelsoni Wiersema Procloeon pennulatum (Eaton) Procloeon quaesitum (McDunnough) Procloeon rivulare (Traver) Procloeon rubropictum (McDunnough)

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Baetidae continued

Procloeon rufostrigatum (McDunnough) † Procloeon simile (McDunnough) Procloeon simplex (McDunnough) Procloeon viridoculare (Berner) Procloeon sp. A + Procloeon sp. B

Pseudocentroptiloides usa Waltz & McCafferty Waynokiops dentatogriphus Hill, Pfeiffer & Jacobus

Baetiscidae +

Baetisca becki Schneider & Berner Baetisca berneri Tarter & Kirchner Baetisca carolina Traver Baetisca escambiensis Berner Baetisca gibbera Berner Baetisca lacustris McDunnugh Baetisca laurentina McDunnough Baetisca obesa (Say) Baetisca rogersi Berner Baetisca rubescens (Provancher)



Dolania americana Edmunds & Traver

Behningiidae

Caenidae

Amercaenis cusabo Provonsha & McCafferty Brachycercus berneri Soldán Brachycercus nitidus (Traver)

Caenis amica Hagen Caenis anceps Traver Caenis diminuta Walker Caenis eglinensis Pescador & Richard Caenis hilaris (Say) Caenis latipennis Banks Caenis macafferti Provonsha Caenis punctata McDunnough Caenis tardata McDunnough

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Caenidae continued

Cercobrachys etowah Soldán Cercobrachys pomeiok Sun & McCafferty

Sparbarus choctaw Sun & McCafferty Sparbarus lacustris (Needham) Sparbarus maculatus (Berner) Sparbarus miccosukee Sun & McCafferty Sparbarus nasutus (Soldán)

Susperatus prudens (McDunnough)

Ephemerellidae

Attenella attenuata (McDunnough) Attenella margarita (Needham) Dannella lita (Burks) Dannella provonshai (McCafferty) Dannella simplex (McDunnough)



Drunella allegheniensis (Traver) Drunella cornuta (Morgan) Drunella cornutella (McDunnough) Drunella lata (Morgan) Drunella longicornis (Traver) Drunella tuberculata (Morgan) Drunella walkeri (Eaton)

Ephemerella catawba Traver Ephemerella crenula Allen & Edmunds Ephemerella dorothea Needham Ephemerella excrucians Walsh Ephemerella floripara McCafferty Ephemerella hispida Allen & Edmunds Ephemerella invaria (Walker) Ephemerella needhami McDunnough Ephemerella rossi Allen & Edmunds Ephemerella subvaria McDunnough Eurylophella aestiva (McDunnough) Eurylophella bicolor (Clemens) Eurylophella doris (Traver)

Ephemerellidae continued

Eurylophella enoensis Funk Eurylophella funeralis (McDunnough) Eurylophella lutulenta (Clemens) Eurylophella macdunnoughi Funk Eurylophella minimella (McDunnough) Eurylophella oviruptis Funk Eurylophella prudentalis (McDunnough) Eurylophella temporalis (McDunnough) Eurylophella verisimilis (McDunnough) Penelomax septentrionalis (McDunnough)

Serratella frisoni (McDunnough) Serratella serrata (Morgan) Serratella serratoides (McDunnough)

Teloganopsis deficiens (Morgan)

Tsalia berneri (Allen & Edmunds)

Ephemeridae

Ephemera blanda Traver Ephemera guttulata Pictet Ephemera simulans Walker Ephemera varia Eaton

Hexagenia atrocaudata McDunnough Hexagenia bilineata (Say) Hexagenia limbata (Serville) Hexagenia orlando Traver Hexagenia rigida McDunnough

Litobrancha recurvata (Morgan)

Heptageniidae

Cinygmula subaequalis (Banks)

Epeorus dispar (Traver) Epeorus fragilis (Morgan) Epeorus namatus (Burks) Epeorus pleuralis (Banks)

Chapter 2 ~ Key to Mayflies

Heptageniidae continued

+ Epeorus punctatus (McDunnough) Epeorus subpallidus (Traver) Epeorus vitreus (Walker)



Heptagenia dolosa Traver Heptagenia flavescens (Walsh) Heptagenia julia Traver Heptagenia marginalis Banks Heptagenia pulla (Clemens) Heptagenia townesi Traver

Leucrocuta aphrodite (McDunnough) Leucrocuta hebe (McDunnough) Leucrocuta juno (McDunnough) Leucrocuta maculipennis (Walsh) Leucrocuta minerva (McDunnough) Leucrocuta thetis (Traver) Leucrocuta sp. 1 Leucrocuta var. A

Maccaffertium appaloosa McCafferty Maccaffertium bednariki McCafferty Maccaffertium carlsoni (Lewis) Maccaffertium exiguum (Traver) Maccaffertium ithaca (Clemens & Leonard) Maccaffertium lenati McCafferty + Maccaffertium luteum (Clemens) Maccaffertium mediopunctatum (McDunnough) Maccaffertium meririvulanum (Carle & Lewis) Maccaffertium mexicanum (Ulmer) Maccaffertium modestum (Banks) Maccaffertium pudicum (Hagen) Maccaffertium pulchellum (Walsh) Maccaffertium sinclairi (Lewis) Maccaffertium smithae (Traver) Maccaffertium terminatum (Walsh) Maccaffertium vicarium (Walker) Maccaffertium wudigeum McCafferty & Lenat

Macdunnoa brunnea Flowers Macdunnoa persimplex (McDunnough)

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Heptageniidae continued +

Nixe flowersi McCafferty Nixe inconspicua (McDunnough) Nixe lucidipennis (Clemens) Nixe perfida (McDunnough) Nixe rusticalis (Traver) Nixe spinosa (Traver)

Raptoheptagenia cruentata (Walsh) Rhithrogena amica Traver Rhithrogena anomala McDunnough Rhithrogena exilis Traver Rhithrogena fasciata Traver Rhithrogena fuscifrons Traver Rhithrogena impersonata (McDunnough) Rhithrogena manifesta Eaton † Rhithrogena rubicunda Traver Rhithrogena uhari Traver †

Spinadis simplex (Walsh)

Stenacron candidum (Traver) Stenacron carolina (Banks) Stenacron floridense (Lewis) Stenacron gildersleevei (Traver) Stenacron interpunctatum (Say) Stenacron minnetonka (Daggy) Stenacron pallidum (Traver)

Stenonema femoratum (Say)

Isonychiidae

Isonychia arida (Say) † Isonychia berneri Kondratieff & Voshell Isonychia bicolor (Walker) ‡ Isonychia diversa Traver Isonychia georgiae McDunnough + Isonychia hoffmani Kondratieff & Voshell Isonychia obscura Traver Isonychia rufa McDunnough Isonychia sayi Burks Isonychia serrata Traver Isonychia sicca (Walsh) Isonychia similis Traver Isonychia tusculanensis Berner

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Leptohyphidae

Asioplax dolani (Allen) † Asioplax texana (Traver) +

Tricorythodes albilineatus Berner Tricorythodes allectus (Needham) Tricorythodes curvatus Allen Tricorythodes robacki (Allen) Tricorythodes stygiatus McDunnough

Leptophlebiidae

Choroterpes basalis (Banks)

Habrophlebia vibrans Needham

Habrophlebiodes americana (Banks) Habrophlebiodes brunneipennis Berner Habrophlebiodes celetaria Berner

Leptophlebia bradleyi Needham Leptophlebia cupida (Say) Leptophlebia intermedia (Traver) Leptohlebia johnsoni McDunnough Leptophlebia nebulosa (Walker)

Paraleptophlebia adoptiva (McDunnough) Paraleptophlebia assimilis (Banks) + Paraleptophlebia calcarica Rowbotham & Allen Paraleptophlebia debilis (Walker) Paraleptophlebia georgiana Traver Paraleptophlebia guttata (McDunnough) Paraleptophlebia jeanae Berner Paraleptophlebia kirchneri Kondratieff & Durfee Paraleptophlebia moerens (McDunnough) Paraleptophlebia mollis (Eaton) Paraleptophlebia ontario (McDunnough)

Leptophlebiidae continued

Paraleptophlebia praepedita (Eaton) + Paraleptophlebia sticta Burks + Paraleptophlebia strigula (McDunnough) Paraleptophlebia swannanoa (Traver) Paraleptophlebia volitans (McDunnough) Traverella lewisi Allen

Metretopodidae † †

Siphloplecton basale (Walker) Siphloplecton brunneum Berner Siphloplecton costalense Spieth Siphloplecton fuscum Berner Siphloplecton simile Berner Siphloplecton speciosum Traver

+

Neoephemera bicolor McDunnough Neoephemera compressa Berner Neoephemera sp. B Neoephemera purpurea (Traver) Neoephemera youngi Berneri

Neoephemeridae

Oligoneuriidae

Homoeoneuria cahabensis Pescador & Peters Homoeoneuria dolani Edmunds, Berner & Traver

Palingeniidae

‡ Pentagenia robusta McDunnough Pentagenia vittigera (Walsh)

Polymitarcyidae

Ephoron album (Say) Ephoron leukon Williamson

+ Tortopsis primus (McDunnough) Tortopsis puella (Pictet)

Chapter 2 ~ Key to Mayflies

Potamanthidae

Anthopotamus distinctus (Traver) Anthopotamus neglectus (Traver) Anthopotamus myops (Walsh) Anthopotamus verticis (Say)



Pseudiron centralis (McDunnough)

Pseudironidae Siphlonuridae

Siphlonurus alternatus (Say) † Siphlonurus decorus Traver ‡ Siphlonurus luridipennis (Burmeister) Siphlonurus marginatus Traver + Siphlonurus marshalli Traver Siphlonurus minnoi Provonsha & McCafferty Siphlonurus mirus (Eaton) Siphlonurus quebecensis (Provancher) Siphlonurus typicus (Eaton)

Page 23

Page 24

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

KEY TO EPHEMEROPTERA LARVAE FAMILIES 1





1’







2(1’)



2’

Gills on abdominal segment 2 either absent, covered by carapace (thoracic shield) from above, or operculate (enlarged and covering followinggills); in mature larvae, forewingpads fused for more than half inner length; gill lamellae (platelike or leaflike part) usually dorsally recumbent on lateral shelves of abdomen; robust or at least partially flattened, sprawling forms...................................................................................................................................17 [Note: These constitute all pannote mayflies (infraorder Pannota) and armored mayflies (suborder Carapacea). They tend to be slow crawling, robust mayflies associated with a variety of substrates and currents. Many are often found among vegetation, debris, on silt/sand, or partially covered in silt.] Gills on abdominal segment 2 present and variously formed, but never covered by carapace nor operculate; in mature larvae, forewingpads separate for half or more of inner length; abdominal gills dorsal, lateral or ventral oriented; when dorsal, usually not recumbent as above, either raised somewhat or held apart somewhat from abdomen; body shapes various...................................................................................................................................2 [Note: These constitute all mayflies of the suborder Pisciforma (generally the minnowlike, brush-legged, and flat-headed mayflies), as well as the burrowing and prong-gilled mayflies of the suborder Furcatergalia.] Abdominal gills 2–7 appearing double, each with elongate gill lamellae profusely fringed with filaments along lateral margins (Figs. 2.1–2.3) (gills 1 rudimentary or absent); tusks (anterior extensions of the mandibles that protrude forward from the head) usually present; if tusks absent, then legs without claws and gills ventrally oriented....................13 [Note: These constitute the burrowing mayflies, including the four families of tusked burrowing mayflies (infraorder Scapphodonta) in the coverage area and the one family of tuskless burrowers (infraorder Palpotarsa). They burrow variously in clay, silt, sand or mixed substrates of streams where they are sometimes taken in drift, or in silt or silt/sand bottoms of ponds and lakes.] Abdominal gills 2–7 never appearing double, elongate, and fringed in combination as above; if gills fringed with filaments, then shorter and gills 1 well developed; tusks not present in southeastern species.............................................................................................3 [Note: Gill numbers refer to abdominal segments to which they are attached.]

Chapter 2 ~ Key to Mayflies



Fig. 2.1

Page 25

Fig. 2.2

Fig. 2.3

Figures 2.1–2.3, Ephemeroptera famm. genn., left gills 4, left dorsolateral. 2.1, Anthopotamus sp. (Potamanthidae); 2.2, Ephoron sp. (Polymitarcyidae); 2.3, Hexagenia sp. (Ephemeridae).

Page 26

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

3(2’)





3’

4(3’)

4’





Abdominal gills 2–7 double or forked with each lamella or fork of lamella narrowelongate and distally pointed (pronglike) (Figs. 2.4–2.6) or ending in cluster of outspread filamentous branches (Fig. 2.7), or with lamellae elongate but broader for much of length, either gradually narrowing into single terminal prong (Fig. 2.9) or with such medioapical pronglike extension accompanied by one or pair of variously developed lateroapical processes (Figs. 2.8, 2.10, 2.11), or infrequently with short lamellae fringed marginally with filaments (Fig. 2.12)......................................LEPTOPHLEBIIDAE, 252 [Note: These are the prong-gilled mayflies or prong-gills. Six genera and 23 species of the family Leptophlebiidae are known from the coverage area. Species may be found in an array of mainly lotic habitats, and the most diverse genus, Paraleptophlebia, is comprised of somewhat delicate narrow-elongate forms, often with vertically oriented heads. More robust forms, such as larvae of Leptophlebia and Choroterpes, tend to have more horizontal heads, but unlike flat-headed mayflies of the family Heptageniidae (see below), they have a portion of the mandibles commonly visible from above.] Abdominal gills 2–7 not as above; if gill lamellae 2–7 pointed or approaching pronglike, then lamellae not double or forked or if rarely appearing forked, then gill with filamentous tuft basally............................................................................................................................4 Head flattened (Figs. 2.13, 2.14), with compound eyes, antennae, and ocelli dorsal on head plate, occasionally area between eyes and antennae elevated somewhat on head plate and antennae arising from sides of elevation; body moderately flattened to flattened.........5 [Note: Some members of the family Oligoneuriidae (see below) have somewhat flattened horizontal heads, but those varieties are not found in eastern North America.] Head not as above, at least somewhat downward oriented and more capsulelike, with compound eyes originating laterally, dorsolaterally or anterolaterally, and antennae, and often at least median ocellus, originating anteriorly on head capsule or on anterior edge of head capsule; body shape variable, often minnowlike, generally not strongly flattened, but some moderately flattened..............................................................................7

Chapter 2 ~ Key to Mayflies

Fig. 2.5

Fig. 2.6

Page 27

Fig. 2.7

Fig. 2.9

Fig. 2.10

Fig. 2.11

Fig. 2.4

Fig. 2.8

Fig. 2.13

Fig. 2.14

Fig. 2.12

Figures 2.4–2.14, Leptophlebiidae and Heptageniidae genn. spp. 2.4–2.12, Leptophlebiidae: 2.4, Paraleptophlebia sp., habitus, dorsal; 2.5–2.7, left gills 4, left dorsolateral: 2.5, Paraleptophlebia sp.; 2.6, Habrophlebiodes sp.; 2.7, Habrophlebia sp. (Edmunds et al., 1963). 2.8, Leptophlebia sp., habitus, dorsal; 2.9–2.12, left gills 4, left dorsolateral: 2.9, Leptophlebia sp.; 2.10, Leptophlebia sp.; 2.11, Choroterpes sp. (Burks, 1953); 2.12, Traverella sp. 2.13–2.14, Heptageniidae genn. spp., habiti, dorsal: 2.13, Maccaffertium lenati; 2.14, Epeorus sp.

Page 28

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

5(4)

5’ 6(5’)



Legs elongate with long and slender claws nearly as long or longer than respective tarsi (Fig. 2.15); abdominal gills 2–7 each consisting of single attenuated pronglike lamella with basal gill tuft and with pronglike filament emerging ventrally from main lamella (Fig. 2.17); horizontal head elevated between compound eyes and antennae arising laterally from side of elevation (Fig. 2.16)....... PSEUDIRONIDAE, Pseudiron, P. centralis [Note: This family (also known as crabwalkers) and genus are monospecific. This carnivorous mayfly hunts midges on shifting sand substrates of mainly rivers and larger streams. It walks crablike in all directions on the sand with its very long, somewhat bowed legs and folds them beneath its body when it takes to rapid swimming.] Legs variable, but claws always much shorter than respective tarsi; abdominal gills 2–7 not exactly as above; horizontal head without central elevation as above...................................... 6

Maxillary palps unique among mayflies, with extremely long, slender (whiplike), second segment with long setae along margins, bordering head and pronotum, as seen in dorsal view (Fig. 2.18), or extending about as far as fore- or midlegs; abdominal gills large, leaf like and pointed (including gills 7) with single lamella and no basal tuft ................................................................ ARTHROPLEIDAE, Arthroplea, A. bipunctata [Note: The family (also known as palpheads) and genus are represented by one species in North America. In the Southeast, A. bipunctata is known only from South Carolina, but this record is questionable. It inhabits bogs, seeps, ponds, and low-flow channels.] 6’ Maxillary palps not as above, never with very long whiplike segment 2 or extending much if any from beneath head; if gill lamellae pointed then with filamentous tuft at base on at least gills 2–5 (pointed gills usually dorsally oriented, or if rarely ventrally oriented then with large filamentous tuft as in Fig. 2.19)...........................HEPTAGENIIDAE, 175 [Note: This family, the flat-headed mayflies, is large and diverse. Within the coverage area, it is represented by 13 genera and about 60 species that are found in a wide variety of freshwater habitats. Larvae are generally benthic sprawlers, and although the vast majority of species are collector/gatherers of fine detritus or periphyton feeders, Raptoheptagenia cruentata and Spinadis simplex are predators.]

Chapter 2 ~ Key to Mayflies

Page 29

Fig. 2.15



Fig. 2.16

Fig. 2.17

Fig. 2.19

Fig.2.18

Figures 2.15–2.19, Pseudironidae, Arthropleidae, and Heptageniidae genn. spp. 2.15–2.17, Pseudiron centralis (Pseudironidae): 2.15, habitus, left lateral; 2.16, head, dorsal; 2.17, left gill 3, left dorsolateral. 2.18, Arthroplea bipunctata (Arthropleidae), habitus, dorsal (modified from Jensen, 1972). 2.19, Raptoheptagenia cruentata (Heptageniidae), left gill 4, left dorsolateral (Jensen, 1972).

Page 30

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

7(4’)



7’









Legs with dorsal lobe of femora (apical lobe at dorsal edge of femur, also sometimes referred to as posterior or outer edge) with ventral-oriented point and/or curvature, sometimes wrapping (then appearing as entire convex apex of femora with lobes discerned only by fusion line), or partially wrapping, ventral lobe, or separate from ventral lobe but with at least ventrally oriented point or extension (Figs. 2.20–2.23); head with main lateral branches of epicranial suture anterior to (below) lateral ocelli, sometimes contacting the anterior or anteromedial edge of ocelli but usually separated from ocelli and often secondarily branching below or lateral of ocelli (Figs. 2.24– 2.28); median caudal filament rudimentary or vestigial in some (two tailed) (Fig. 2.29); antennae often relatively long, 2–3× width of head; if antennae atypically short, then either two tailed (Fig. 2.29) or rarely three tailed; body highly streamlined (Fig. 2.30) or only somewhat flattened..........................................................................BAETIDAE, 29 [Note: The family is highly diverse and consists of 17 genera and at least 65 species within the coverage area. They can be expected in a wide variety of freshwater habitats. More recently discovered unique characteristics of the family include the development of the dorsal lobe of the femora and the position of the branches of the epicranial suture, as explained above. The femoral character should be easily seen in all cases, and the epicranial suture upon close examination is accessible in most cases, but sometimes is obscured by the developing eyes of very mature male larvae with narrow heads. Other qualified characters are given for convenience, with the odd occurrence of a combination of short antennae together with three tails associated with the rarely taken species Barbaetis benfieldi, which, however, does demonstrate familial characteristics such as a convex femoral apex.] Legs with dorsal lobe of femora not ventrally oriented as above, often dorsal lobe not developed, and ventral lobe sometimes highly developed (e.g., Figs. 2.31–2.34); head with main lateral branches of epicranial suture usually connecting with lateral ocelli near center of ocelli or connecting with them at their posterior edge or running slightly posterior to (above) ocelli (e.g., Figs. 2.35–2.38); never two tailed; antennae variable in length; if antennae over 2× width of head, then two rows of long setae on inner margin of forelegs; generally minnowlike (streamlined) bodies......................................................8 [Note: Some, but not all, of these mayflies have a body length over 12mm at maturity; this would generally distinguish these larger forms from Baetidae. Body length referred to throughout the key never includes caudal filaments (tails).]

Chapter 2 ~ Key to Mayflies

Fig. 2.20

Page 31

Fig. 2.21

Fig. 2.22

Fig. 2.23 Fig. 2.24

Fig. 2.26

Fig. 2.27

Fig. 2.29

Fig. 2.28

Fig. 2.30

Fig. 2.25

Fig. 2.31

Fig. 2.32

Fig. 2.33

Fig. 2.34

Fig. 2.35

Fig. 2.36

Fig. 2.37

Fig. 2.38

Figures 2.20–2.38, Ephemereoptera famm. 2.20–2.23, Baetidae genn. spp., right hindfemora apices (anterior faces, with dorsal margins up, pointers to dorsal lobes). 2.24–2.28, Baetidae genn. spp., heads, anterior. 2.29–2.30, Baetidae, habiti, dorsal: 2.29, Plauditus cestus; 2.30, Baetis intercalaris. 2.31–2.34, Ephemeroptera famm., right hindfemora apices (anterior faces, with dorsal margins up, pointers to dorsal lobes): 2.31, Ameletidae sp.; 2.32, Siphlonuridae sp.; 2.33, Heptageniidae sp.; 2.34, Oligoneuriidae sp. 2.35–2.38, Ephemeroptera famm., heads, anterior: 2.35, Ameletidae sp.; 2.36, Metretopodidae sp.; 2.37, Leptophlebiidae sp.; 2.38, Heptageniidae sp.

Page 32

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

8(7’) 8’

Forelegs with two rows of long setae on inner margin of femur and tibia (Figs. 2.39, 2.40).....9 Forelegs without rows of long setae as above..........................................................................10

9(8)

Forelegs normal with respect to having tarsus and claw (Figs. 2.39, 2.41); abdominal gills 1 dorsolaterally oriented...................................................... ISONYCHIIDAE, Isonychia, 239 [Note: This family and genus, the brush-legged mayflies, are represented in the coverage area by 11 species, only one of which remains unknown as larvae. Mature larvae range in size from 9–17 mm and are passive filter feeders common in riffle zones of streams.] Forelegs lacking claw and tarsus segment as such (Figs. 2.40, 2.42); gills 1 ventral ........................................................................ OLIGONEURIIDAE, Homoeoneuria, 286 [Note: This family (the sand brushlegs) is represented by two species of the genus Homoeoneuria in the area of coverage. Larvae evidently are passive filter feeders in current in depressions on sandy substrates of streams and rivers.]

9’







10(8’) Foreclaws bifid (Figs. 2.43, 2.44)......................... METRETOPODIDAE, Siphloplecton, 278 [Note: This family (also known as the cleft-footed minnows) is represented in the area of coverage by a small number of species of the genus Siphloplecton. Larvae are known from streams and rivers; prior to emergence, they may be found among vegetation in shallows near shore.] 10’ Foreclaws not bifid (e.g., Fig. 2.39)..........................................................................................11 11(10’)

11’





Abdominal terga with median row of hooklike tubercles as shown in Fig. 2.45; abdominal sterna with blunt conical tubercles, best developed on sterna 4–8; legs crablike (bowed with long, slender claws), with hindclaws at least as long as hindtarsi..................... ACANTHAMETROPODIDAE, Acanthametropus, A. pecatonica [Note: This family (also known as speed minnows) and genus are represented by only a single species in eastern North America. Within the coverage, area it has been taken only in Georgia and South Carolina. Much like Pseudiron (Pseudironidae) discussed above (but much less frequently collected), it would appear to be a crablike walker and hunter on sandy substrates of streams and rivers, and it also is a very fast swimmer. This has been observed by us in the only other North American genus and species of the family, Analetris eximia (western North America), and is presumed for Acanthametropus pecatonica based on its similar habitat and general morphology.] Abdomen without dorsal and ventral median tubercles as above; legs and claws not as above...............................................................................................................................12

Chapter 2 ~ Key to Mayflies

Page 33

Fig. 2.39

Fig. 2.41

Fig. 2.40

Fig. 2.42

Fig. 2.43

Fig. 244

Fig. 2.45 Figures 2.39–2.45, Ephemeroptera famm. 2.39–2.40, right forelegs, anterodorsal: 2.39. Isonychia obscura (Isonychiidae; Kondratieff & Voshell, 1984); 2.40, Homoeoneuria sp. (Oligoneuriidae; Edmunds et al., 1958). 2.41–2.42, Habiti, dorsal: 2.41, Isonychia sicca (Isonychiidae); 2.42, Homoeoneuria sp. (Oligoneuriidae; Edmunds et al., 1958). 2.43, Siphloplecton speciosum (Metretopodidae), left foreclaw, dorsal (modified from Berner, 1978). 2.44–2.45, habiti: 2.44, Siphloplecton sp. (Metretopodidae), dorsal; 2.45, Acanthametropus pecatonica (Acanthametropodidae), left lateral.

Page 34

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

12(11’)

12’

13(2)





13’ 14(13’)

14’

Abdominal gills large, leaflike, broad, and well tracheated, with broadly indented distal margin, and appearing double due to dorsal recurved flap on at least gills 1 and 2 (Fig. 2.46, 2.47)..................................................... SIPHLONURIDAE, Siphlonurus, 292 [Note: This family, sometimes referred to as the big minnow mayflies (as opposed to the small minnow mayflies, family Baetidae), is represented in the coverage area by six species of the genus Siphlonurus. More mature larvae of Siphlonurus inhabit shallow pools, swampy areas, ponds, lake margins, and stream edges; however, younger larvae of some species may develop in connector streams. They occur on silted substrates, but are also adept swimmers, and apparently are omnivores.] Abdominal gills relatively small, platelike, and ovate, without tracheation but with riblike outer margin and usually with a similar riblike inner margin or sclerotized rib near inner margin (Fig. 2.48–2.50)..............................................AMELETIDAE, Ameletus, 21 [Note: These painted minnows, as they are sometimes called because of their often striking color patterns, are represented in the coverage area by at least five species of Ameletus (the only genus of Ameletidae in North America). Larvae inhabit streams where they can be found in current or often in small edge pools or quiet waters where they may dart rapidly for short distances. They are also reported from large rivers, lakes, and ponds with rock or gravel bottoms.] Legs aberrant, lacking terminal claw; gills ventrally oriented; tusks not present (Fig. 2.51) ......................................................................... BEHNINGIIDAE, Dolania, D. americana [Note: This family of primitive tuskless burrowers is monogeneric in North America and known primarily from the Southeast. The unusual larvae, with their palplike forelegs, are known from a limited number of populations, and D. americana might represent a species complex. Larvae burrow in clean large-grain sand of streams without creating a burrow. They do create a small chamber, however, under the body with their mid- and hindlegs where gills have been witnessed to beat and are protected from abrasion. They are known to be predatory.] Legs with terminal claw; gills lateral or dorsal; tusks present..................................................14 Gills laterally oriented; body considerably flattened; foretibiae relatively narrow (Fig. 2.52) ......................................................................... POTAMANTHIDAE, Anthopotamus, 289 [Note: This family, the so-called hacklegills, is represented by one genus and four species in North America. All four are known from the coverage area. Larvae are burrowers, mainly in mixed substrates of streams, often along the edges of large partially buried rocks or large boulders.] Gills dorsal; body more or less cylindrical or somewhat flattened; foretibiae at least somewhat modified or expanded........................................................................................15

Chapter 2 ~ Key to Mayflies

Page 35

Fig. 2.46 Fig. 2.47

Fig. 2.48 Fig. 2.49

Fig. 2.50

Fig. 2.51 Fig. 2.52

Figures 2.46–2.52, Ephemeroptera famm. 2.46–2.47, Siphlonurus sp. (Siphlonuridae): 2.46, habitus, left lateral (modified from Burks, 1953); 2.47, left gill 2, left dorsolateral. 2.48–2.50, Ameletus spp. (Ameletidae): 2.48, habitus, left lateral (modified from Burks, 1953); 2.49–2.50, left gills 4, left dorsolateral. Figs. 2.51–2.52, habiti: 2.51, Dolania americana (Behningiidae, left dorsolateral); 2.52, Anthopotamus myops (Potamanthidae), dorsal.

Page 36

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

15(14’)



15’ 16(15’)

16’





Forefemora produced somewhat posteroproximally beyond trochanter (Figs. 2.53, 2.54); tusks (Fig. 2.55) somewhat broadened in middle but attenuated to tips, with outer marginal keel roughened variously and with some heavy spurs, and tips of tusks never convergent or downturned; frontal process bifurcate (Fig. 2.55) .................................... ..................................................................... PALINGENIIDAE, Pentagenia, P. vittigera [Note: This family, also known as the spiny-headed burrowers because of the roughened appearance of the frontal process between the antennal bases and the smaller processes over the antennal bases, is represented in North America by a unique genus and single extant species. Larvae are known from rivers where they form burrows in clay banks. The other North American species of the genus, P. robusta, is presumed to be recently extinct.] Forefemora not produced at its base as above; tusks not broadened in middle, without outer marginal keel with spurs or spines; if outer margin with row of armature, then tusk tips appearing convergent in dorsal view and frontal process not bifid; frontal process either broadly based and bifurcate, broadly based and variously obtuse, narrow and rounded, or not developed ............................................................................................................... 16 Tusk tips inwardly and/or downwardly oriented; tusks with unsocketed spines scattered dorsally on tusks or with one heavy spine subdistally on medial margin; hindtibiae not produced distally well beyond base of tarsus into acute tibial process; frontal process narrow and rounded or not developed (Figs. 2.56, 2.57) . .... POLYMITARCYIDAE, 287 [Note: This family, the pale burrowers, is represented by two genera and three species in the coverage area. Larvae are mainly stream and river inhabitants, with Tortopsis and sometimes Ephoron being clay burrowers, with Ephoron also known to burrow in mixed substrates.] Tusk tips outwardly and/or upwardly oriented (Figs. 2.58, 2.59); tusks with armature (small socketed spurs) only in basal half laterally or dorsolaterally, mostly not apparent in dorsal view of head; hindtibiae produced well beyond base of tarsus into acute process (Fig. 2.60); frontal process either broadly based between antennae and variously obtuse (Fig. 2.59), or bifurcate and forming short anterolateral prongs (Figs. 2.58) ........................................................................................................ EPHEMERIDAE, 167 [Note: This family of common burrowers is represented in the coverage area by three genera and 10 species. Hexagenia larvae are the prevalent silt burrowers of streams, rivers, ponds, and lakes, sometimes known for their mass emergences, whereas larvae of Litobrancha burrow mainly in marl of small streams, and those of Ephemera may burrow in sand/silt or mixed substrates of mainly lotic, but occasionally lentic, habitats.]

Chapter 2 ~ Key to Mayflies

Page 37

Fig.2.53 Fig. 2.54 Fig.2.55

Fig. 2.56

Fig. 2.57

Fig. 2.58 Fig. 2.59

Fig.2.60

Figures 2.53–2.60, Palingeniidae, Polymitarcyidae, and Ephemeridae genn. spp. 2.53–2.55. Pentagenia vittigera (Palingeniidae): 2.53, habitus, left dorsolateral; 2.54, right foreleg, anterodorsal; 2.55, head, dorsal. 2.56, Ephoron album (Polymitarcyidae), habitus, left dorsolateral. 2.57, Tortopsis puella (Polymitarcyidae), head, left dorsolateral. 2.58–2.60, Ephemeridae genn. spp.: 2.58, Ephemera simulans, habitus, left dorsolateral; 2.59–2.60, Hexagenia sp.: 2.59, head, pronotum, foreleg base, dorsal; 2.60, right hind tibia apex and tarsus, anterodorsal.

Page 38

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

17(1)

Thorax modified into carapace covering base of legs and abdomen from above (Fig. 2.61) ........................................................................................... BAETISCIDAE, Baetisca, 100 [Note: This family of armored mayflies is represented in the coverage area by nine of the 11 recognized species, making the Southeast the richest area for the family. Larvae are found in running waters where they tend to be associated with sand, sand/silt or sand/gravel substrates but sometimes are found on debris or other objects. They have been observed to swim in a labored way for short distances.] Thorax not modified into a carapace, although forewingpads of mature larvae are fused for much of their length .......................................................................................................... 18



17’ 18(17’)



18’ 19(18’)

19’ 20(19’)

20’





Gills on abdominal segment 2 absent (Figs. 2.62, 2.63), and sometimes also absent from abdominal segment 3 (either rudimentary or absent on abdominal segment 1); gills on abdominal segments 3 or 4 sometimes operculate or partially operculate . ............................................................................. EPHEMERELLIDAE, 128 [Note: The so-called spiny crawlers are a diverse and extensive family of mayflies. In the coverage area it is represented by nine genera and probably more than 35 species. Larvae generally are found in running water habitats and may be benthic sprawlers, climbers or clingers.] Gills present on abdominal segment 2 and modified into operculate covers of following gills ................................................................................................................................... 19 Operculate gills ovate, subovate, triangulate, or subtriangulate (never square or subquadrate) (Figs. 2.64–2.67) ....................................................................... LEPTOHYPHIDAE, 248 [Note: This family of little stout crawlers is poorly represented in the Southeast compared to the Southwest; two genera and six species are known from the coverage area. Larvae often are associated with silt or silt/sand substrates in streams, where they are generally slow and sprawling forms. Their setae often are packed with silt.] Operculate gills nearly square or subquadrate (Figs. 2.68–2.70) ........................................... 20 Operculate gills not interlocking along medial margins, usually slightly overlapping (Figs. 2.68, 2.69); mesonotum (dorsum of second thoracic segment) without anterolateral processes (Figs. 2.68, 2.69); hindwingpads absent; mature body length usually 2–8 mm ................................................................................................................. CAENIDAE, 109 [Note: These are the small squaregills, represented in the coverage area by six genera and 20 species. Larvae generally are benthic sprawlers or climbers known from a variety of low- to moderate-flow lotic habitats, as well as lentic habitats. Body length never includes caudal filaments (tails).] Operculate gills meeting and interlocking with setae along medial margins (Fig. 2.70); mesonotum with anterolateral processes (Fig. 2.70); hindwingpads present; mature body length 8–17 mm ................................... NEOEPHEMERIDAE, Neoephemera, 283 [Note: These are the large squaregills. Larvae may be found in slow to swift currents under stones or amongst debris, among roots in cut banks, or on moss-covered driftwood. As noted above, their longer body does not include the caudal filaments.]

Chapter 2 ~ Key to Mayflies

Page 39

Fig. 2.61

Fig. 2.62

Fig. 2.63

Fig. 2.65

Fig. 2.66

Fig. 2.64

Fig. 2.67

Fig. 2.68

Fig. 2.69

Fig. 2.70

Figures 2.61–2.70, Ephemeroptera famm. 2.61–2.63, habiti: 2.61. Baetisca sp. (Baetiscidae); 2.62–2.63, Ephemerellidae genn. spp.: 2.62, Ephemerella excrucians (Allen & Edmunds, 1965); 2.63, Eurylophella bicolor (Allen & Edmunds, 1963a). 2.64–2.67, Leptohyphidae genn. spp.: 2.64, Tricorythodes sp., habitus. 2.65–2.67, right operculate gills 2, right dorsolateral. 2.68–2.70, habiti: 2.68, Caenis hilaris (Caenidae); 2.69, Brachycercinae sp. (Caenidae). 2.70, Neoephemera pupurea (Neoephemeridae; Edmunds et al., 1963).

Page 40

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

AMELETIDAE, Ameletus species 21(12’) 21’

Abdominal tergum 2 with posterior marginal microspines (Fig. 2.71) .................................. 22 Abdominal tergum 2 without posterior marginal spines (Fig. 2.72) . ..................................... 28 [Note: Five nominal species of Ameletus represent the described Ameletidae of the coverage area. All five are known as larvae; however, undescribed larval variants or species apparently are represented in the coverage area (see below). Ameletus janetae, from nearby West Virginia, is not known in the larval stage.]

22(21) 22’

Caudal filaments dark from base or from very near base, with pale distal band .................... 23 Caudal filaments pale, with or without dark medial band ...................................................... 24

23(22)



Abdomen with dorsal, pale mid-longitudinal stripe on terga 1–9 .................. Ameletus browni [Note: This species is not known from the coverage area, but it has a nearest confirmed locale in Pennsylvania.] Abdomen without mid-longitudinal stripe as above . ........................................ Ameletus sp. 3 [Note: This larval variety has been found in North Carolina.]

24(22’) 24’

Abdomen with three ventral, dark mid-longitudinal stripes ................................................... 25 Abdomen without such ventral stripes, but sometimes with single indistinct medial mid-longitudinal stripe ..................................................................................................... 26

25(24)

Ventral abdominal stripes distinct and contrasting strongly with background coloration, stripes sometimes coalescing posteriorly on sternum 9 . ........................ Ameletus lineatus [Note: The sharpness of striping is subject to fading. This species might be equivalent to A. ludens.] Ventral abdominal stripes diffuse with edges not well defined, stripes more or less coalescing variously on sterna 9, 8–9, or 7–9 . ........................................................... Ameletus ludens

23’





25’ 26(24’) 26’

Caudal filaments with dark medial band . ............................................................................... 27 [Note: The tail-band width varies among species.] Caudal filaments without distinctive medial band ............................................. Ameletus sp. 2 [Note: This larval variety, possibly a variant of A. tertius or an undescribed species, has been found in North Carolina.]

27(26) 27’

Abdominal terga 3–6 medium brown with distinctively contrasting pale spots; terga 7–8 pale; terga 9–10 dark brown (Fig. 2.73) .................................................... Ameletus tertius Abdominal terga medium brown with faint pale areas; terga 8–10 somewhat darker with less spotting ................................................................................................. Ameletus sp. 1 [Note: This probably valid species, which has been found in North Carolina, may also be differentiated by the possession of elongate pale spots on the anterior face of the femora.]



28(21’) 28’

Abdominal terga 3–10 or 4–10 with spines on posterior margins ................... Ameletus tarteri Terga 5–10 or 6–10 with spines on posterior margins ....................... Ameletus cryptostimulus

Chapter 2 ~ Key to Mayflies

Page 41

Fig. 2.71

Fig. 2.72

Fig. 2.73 Figures 2.71–2.73, Ameletus spp., Ameletidae. 2.71–2.72, Ameletus spp., abdominal terga 2, left half, dorsal: 2.71, A. ludens; 2.72, A. cryptostimulus. 2.73, A. tertius, terga 4–10 + caudal filaments, dorsal.

Page 42

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

BAETIDAE genera 29(7)





29’





30(29’) 30’

Gills relatively large with one or more recurved ventral (posterior) flaps or lobes on at least gills 1–6 (e.g., Figs. 2.74–2.76) (those with large flaps appearing as double or triple gills); branching tracheation always apparent in gills and consisting of midlongitudinal main trunk in at least basal half of gill with bilateral and apical branching (e.g., Figs. 2.74, 2.75); main lamella of at least gills 4–7 longer than broad (e.g., Figs. 2.74, 2.75); claws somewhat long and slender and only slightly curved, ventrally with two rows of well-developed denticles, with more-distal denticles spinelike (e.g., Fig. 2.77) ....................................................................................................................... Callibaetis, 66 [Note: Callibaetis is commonly found in lentic habitats but also can be found in lotic habitats.] Gills variable, but never as above, often simple, without flaps or lobes, or if flap present, then single and dorsal (anterior) on one or more pairs of gills; gills either without branching tracheation, or with variable types of branching such as with mid-longitudinal trunk with bilateral branching, or with mid-longitudinal trunk with asymmetrical branching mostly toward inner expanded area of gill, or with several main trunks emerging from central base of gill; gills varying in shape, sometimes broader than long; claws variable, often short and moderately to strongly curved (Figs. 2.78, 2.79), sometimes long or very long and lanceolate (Figs. 2.80–2.82), rarely rakelike (Fig. 2.83), and with or without denticles; if claws elongate and with two rows of denticles, then denticles not developed (e.g., Fig. 2.82) as above .................................................... 30 [Note: The vast majority of Baetidae that key here are lotic; relatively few are lentic, though these habitats remain underexplored in the Southeast and elsewhere.] Median caudal filament (middle tail) developed, at least half as long as cerci (lateral pair of tails) ...................................................................................................................... 31 Median caudal filament reduced, ranging from minute to not much longer than abdominal tergum 10, usually with ten or fewer segments ................................................................ 40 [Note: A developed middle tail sometimes breaks off near the base, in which case there should be evidence of it from a truncate stub noticeable with close examination. An unbroken, rudimentary or vestigial middle tail will have a pointed apex.]

Chapter 2 ~ Key to Mayflies

Fig. 2.74

Page 43

Fig. 2.75

Fig. 2.76

Fig. 2.77

Fig. 2.78

Fig. 2.81

Fig. 2.79

Fig. 2.82

Fig. 2.80

Fig. 2.83

Figures 2.74–2.83, Baetidae genn. spp. 2.74–2.76, Callibaetis spp., right gills 6, right dorsolateral: 2.74–2.75, ventral flap unfolded (modified from Check, 1982): 2.74, C. fluctuans; 2.75, C. floridanus; 2.76, C. floridanus, ventral flap in normal fold. 2.77, C. floridanus, right foreclaw, posteromesal. 2.78–2.83, right foretarsi and claws: 2.78, Baetis sp.; 2.79, Acentrella sp.; 2.80, Anafroptilum sp.; 2.81, Pseudocentroptiloides usa; 2.82, Cloeon dipterum; 2.83, Camelobaetidius musseri.

Page 44

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

31(30)





31’ 32(31’)

Terminal segment of labial palps simple, without lobes, and usually with more or less straight apical margin (truncate), often slightly concave or slightly rounded (Figs. 2.84– 2.87), distal margin often perpendicular to mid-longitudinal line of terminal segment or sometimes oblique (diagonal to mid-longitudinal line of terminal segment), and labial palps rarely falcate distally (appearing oblique principally because outer aspect produced somewhat distally as in Fig. 2.87); gills simple or with single dorsal flap (or combination of both on individuals), gills sometimes broad and when tracheated, then with asymmetrical branching to inner expanded side or commonly palmate with trunks radiating from central base ............................................................................................... 43 [Note: This grouping is often referred to as the long-clawed baetids because of the slender, long claws associated with many of them—long claws probably are an adaptation for the silted or sandy substrates on which many with such claws are commonly found. Some other baetids not in this grouping also have long claws. In addition, while not exclusive to the longclawed baetids, these mayflies tend to be very slender and fusiform and are among the most minnowlike of the small minnow mayflies.] Terminal segment of labial palps variable, but not as above; gills simple (without dorsal flap), generally longer than broad and, when tracheated, with at least mid-longitudinal trunk for some distance and often with bilateral branching . ............................................ 32

32’

Claws rakelike, expanded laterally and apically, and with transverse apical row of long denticles (e.g., Fig. 2.83); within the Southeast known only from southwestern North Carolina . ................................................................................ Camelobaetidius, C. musseri [Note: This is the only southeastern Nearctic species of the genus and only the third North American species known outside the Southwest and Neotropics. Some questions, however, surround the North Carolina record.] Claws not specialized as above . ............................................................................................. 33

33(32’) 33’

Claws clearly shorter than respective tarsi; labrum with medial indentation along distal margin (e.g., Fig. 2.88), rarely indentation very small (Fig. 2.89); hindwingpads present or absent ............................................................................................................................ 34 Claws as long as respective tarsi; labrum without medial indentation along distal margin (Fig. 2.90); hindwingpads absent; usually associated with sand ................... Apobaetis, 59

34(33) 34’

Labial palps similar to Fig. 2.91; hindwingpads absent ................................. Paracloeodes, 82 Labial palps variable, but not as above; hindwingpads present . ............................................ 35

35(34’)

Mouthparts generally with setose appearance and labial palp distinctively bulbous with distal constriction (Fig. 2.92); terminal segment of maxillary palp usually with inflated appearance; known only rarely, from sandy streams in Kentucky and North Carolina . ......................................................................................................... Baetopus, 65 Mouthparts not exactly as above ............................................................................................ 36

35’

Chapter 2 ~ Key to Mayflies

Fig. 2.84

Page 45

Fig. 2.85

Fig. 2.88

Fig. 2.86

Fig. 2.89

Fig. 2.91

Fig. 2.87

Fig. 2.90

Fig. 2.92

Figures 2.84–2.92, Baetidae genn. spp. 2.84–2.87, left labial palp outlines, ventral: 2.84, Pseudocentroptiloides usa; 2.85, Anafroptilum album; 2.86, Procloeon nelsoni; 2.87, Cloeon dipterum. 2.88–2.90, Labra, anterodorsal: 2.88. Heterocloeon frivolum; 2.89, Paracloeodes minutus (Day, 1955); 2.90, Apobaetis etowah (Day, 1955). 2.91, Paracloeodes minutus, left labial palp outline, ventral. 2.92, Baetopus sp. labium, left lateral half and palp, ventral (modified from Keffermüller, 1960).

Page 46

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

36(35’)



Gills present on abdominal segment 1; coloration and general appearance of specimens variable ............................................................................................................................. 37 [Note: At least gill sockets should be present in cases where gills 1 were dislodged.] Gills and associated gill sockets absent on abdominal segment 1; dorsal color pattern of abdomen sometimes with one pair of submedian pale dots and anteriorly converging pale streaks; posterior terga often paler than anterior terga; fluid-preserved specimens often with hunch-backed appearance not usually seen in other taxa ..... Diphetor, D. hageni

37(36) 37’

Labial palps mittenlike, with large thumblike lobe distomedially oriented (at approximately 50–60 degree angle with palp) (Fig. 2.93); gills 7 relatively sharply pointed ................. ....................................................................................................................... Acerpenna, 54 Labial palps variable, never as above, if labial palp with large lobe, then lobe very broad and medially oriented (at nearly right angle with palp) (e.g., Fig. 2.94); gills 7 more or less rounded apically or at most bluntly pointed .............................................................. 38

38(37’)

Basal segment (scape) of antennae with small notch at distal margin producing small outer distal lobe (e.g., Fig. 2.95); maxillary palps with inner subapical excavation (e.g., Fig. 2.96); labial palps usually with broad medially directed lobe (e.g., Fig. 2.94) ...................................................................................................................... Labiobaetis, 78 Basal segment of antennae without notch or lobe as above; maxillary palps not excavated; labial palps variable, but not exactly as above, with poorly developed or less developed medial lobe . .......................................................................................................................39

36’

38’ 39(38’)

39’ 40(30’)

40’



Antennae 2–3× length of head capsule; forecoxal gill filaments not present ............ Baetis, 60 [Note: Because the genus Fallceon has not been reported from the coverage area, we consider it unlikely to be present. It has been found in nearby Illinois and Louisiana, however, and is common throughout central and western North America. It is distinguished from Baetis by a distinct raised keel on the head, running longitudinally between the antennal bases, and by having a distinct distal excavation on labial palp segment 3.] Antennae subequal in length to head capsule; forecoxal gill filaments present; known only from the Carolinas and Virginia . ..................................................... Barbaetis, B. benfieldi Claws usually with two rows of denticles, with one row having more distinctive denticles and secondary row having less developed (sometimes poorly developed) denticles (Figs. 2.97, 2.98) or secondary row represented only by weak ridge with no denticles; if secondary ridge of claw without denticles, then such individuals relatively large, with mature body length about 7–9 mm, and with femora lacking marginal femoral row of long, fine setae, and with tibiae having deeply forked, short spatulate setae on anterior face (Figs. 2.99, 2.100); otherwise mature body length 3–9 mm ..................................... 41 [Note: Body length never includes caudal filaments.] Claws with one row of denticles and no secondary ridge; mature body length about 3–6 mm ................................................................................................................... 42

Chapter 2 ~ Key to Mayflies

Fig. 2.93

Page 47

Fig. 2.94

Fig. 2.95

Fig. 2.96

Fig. 2.99

Fig. 2.100 Fig. 2.97

Fig. 2.98

Figures 2.93–2.100, Baetidae genn. spp. 2.93, Acerpenna pygmaea, left labial palp outline, ventral. 2.94–2.96, Labiobaetis propinquus: 2.94, left labial palp outline, ventral; 2.95, left scape and pedicel, dorsal; 2.96, left maxillary palp distal segment, posteroventral. 2.97–2.98, right foreclaws, anteroventral: 2.97, Iswaeon davidi; 2.98, Heterocloeon curiosum (Müller-Liebenau, 1974). 2.99–100, Heterocloeon amplum: 2.99, right hind leg, anterior face, dorsal up; 2.100, specialized setae on anterior face of a hind tibia.

Page 48

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

41(40) 41’



42(40’)





42’



43(31)

43’

Primary row of claw denticles becoming progressively longer towards apex of claws as in Figs. 2.98, 2.99; mature body length 4–9 mm, often well over 4 mm; gill filament often present near base of forelegs (e.g., Fig. 2.101); small hindwingpads (Figs. 2.102, 2.103) often present; tibae not expanded as below ............................................. Heterocloeon, 68 Primary row of claw denticles of equal length except for shorter basal denticles (e.g., Fig. 2.97); mature body length 3–4 mm; gill filament never present near base of forelegs; hindwingpads not present (represented by microscopic scale at most); tibiae with expanded appearance (e.g., Fig. 2.104) .................................................... Iswaeon, 77 [Note: Although median dots on abdominal sterna are typical of Iswaeon, such dots may not always be present.] Femora with marginal row of fine, often somewhat long setae (e.g., Fig. 2.105); labium with short palps with segments 2 and 3 sometimes appearing fused with outer and apical margins rounded, segment 3 usually somewhat boxing-glove-fist shaped, and segment 2 usually strongly curved nearly 90 degrees to medial, resulting in palp overlapping paraglossa, and with compact glossae and paraglossae (tightly fitted with paraglossae tending inward over apices of glossae) (similar to Fig. 2.106); hindwingpads present, absent, or vestigial ........................................................................................ Acentrella, 50 [Note: Fine marginal setae of legs can be difficult to detect with improper illumination or contrast because such setae are often pale. Rows of leg setae may differ in density among species. The glossae of the labium are the inner pair of distal lobes on the main body of the labium, and paraglossae are the outer pair of lobes adjacent to the glossae.] Femora without marginal row of fine setae (if row of marginal setae present, then somewhat short and bristlelike); labium with terminal segment 3 of palp distinct, subquadrate or subfalcate apically, and with segment 2 not as curved inwardly as above, and paraglossae and glossae not compacted as above (e.g., Fig. 2.107); hindwingpads absent ................ ......................................................................................................................... Plauditus, 83 [Note: Of the eight species of Plauditus known from the coverage area, only P. veteris remains unknown as larvae.] Abdominal terga 2–5 (Fig. 2.108) with midposterior spines, spines best developed and raised on tergum 2, sometimes poorly developed on 1 and 6; abdomen with well-developed lateral shelflike expansions on segments 3–7 (Fig. 2.108); hindwingpads absent; lentic habitats ............................................................................. Waynokiops, W. dentatogriphus [Note: This species is known only from Kentucky within the area of coverage, but it is also known from Arkansas and Virginia and thus expected to be more widespread in the Southeast, especially considering that lentic habitats have been historically poorly sampled for mayfly larvae.] Abdomen not as above; hindwingpads present or absent; only rarely from lentic habitats . ..... 44

Chapter 2 ~ Key to Mayflies

Page 49

Fig. 2.102

Fig. 2.101

Fig. 2.104

Fig. 2.103

Fig. 2.105

Fig. 106

Fig. 2.108 Fig. 2.107 Figures 2.101–2.108, Baetidae genn. spp. 2.101, Heterocloeon berneri, right foreleg base with gill filament, anterior (modified from Müller-Liebenau, 1974). 2.102–2.103, Heterocloeon spp., hind wingpads: 2.102, H. frivolum; 2.103, H. curiosum. 2.104–2.105, right forelegs, anterodorsal: 2.104, Iswaeon davidi; 2.105, Acentrella turbida. 2.106–2.107, labia, ventral (left) and dorsal (right): 2.106, Acentrella turbida; 2.107, Plauditus sp. 2.108, Waynokiops dentatogriphus abdominal terga, dorsal.

Page 50

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

44(43’)



44’ 45(44’)

45’ 46(45’)



46’





Labrum with broad V-shaped distal margin as in Fig. 2.109; labium with glossae broad, apically blunt, and distinctly shorter than paraglossae (Fig. 2.110); claws somewhat longer than respective tarsi (Fig. 2.111); hindwingpads present; larval habitat usually gravel or sand . ..................................................................... Pseudocentroptiloides, P. usa [Note: Glossae are the innermost lobes of the labium and are bordered laterally by the paraglossae.] Labrum not as above, similar to Fig. 2.112; glossae width variable, but pointed and subequal to, or not much shorter than, paraglossae (e.g., Figs. 2.113, 2.114); claws usually at least somewhat shorter than respective tarsi; hindwingpads present or absent ................ 45 Labial palps falcate distally (Fig. 2.115); gills 1–6 with well-developed dorsal flaps, with dorsal flap on gill 6 subequal to flap on gill 5; hindwingpads absent; lentic habitats ............................................................................................................. Cloeon, C. dipterum [Note: This species is known only from Kentucky within the area of coverage, but it is also known from adjoining states such as Ohio, Indiana, and Virginia. It may be more widespread in the upper Southeast, especially considering that lentic habitats have been historically poorly sampled for mayfly larvae.] Not with exact combination of characteristics given above; hindwingpads present or absent; generally lotic habitats .......................................................................................................46 Both mandibles with incisors fused for half length or more (Figs. 2.116, 2.117); planate mandible (Fig. 2.117) with inner incisor curving inward so apices of denticles rowed more vertically than horizontally, and with prostheca broadened; usually one or more pairs of gills with dorsal flap, or sometimes all gills simple; well-developed lateral spines always present on at least abdominal segments 8 and 9; hindwingpads present or absent ............................................................................................................. Procloeon, 90 [Note: Of the nine nominal species of Procloeon known from the coverage area, seven are known as larvae, with only P. intermediale and P. simile remaining undescribed as larvae. The angulate mandible has a mola region (area with a series of serrations) that forms a distinct angle with the distal margin and typically is the left mandible, whereas the planate mandible tends to have a mola region on the same plane as the distal margin of the mandible, generally the right one. The incisors are the large toothlike structures apically (usually two, sometimes fused). Denticles are the points or serrations associated with the incisors. The prostheca is a large setal or spinelike structure emerging just prior to the incisors on the distal margin.] Angulate mandible (Fig. 2.118) with incisors somewhat fused (slightly less than half length); planate mandible (Fig. 2.119) with incisors separated to base, with inner incisor often narrowing distally and not curved inward, and with prostheca appearing as simple bristle or bristle with some lateral hairlike branches; all gills simple, without dorsal flap; lateral spines on abdomen variable (absent, poorly developed, or well developed); hindwingpads present or absent ........................................................................................ 47

Chapter 2 ~ Key to Mayflies

Page 51

Fig. 2.109

Fig. 2.111 Fig. 2.110

Fig. 2.112 Fig. 2.113

Fig. 2.115

Fig. 2.114

Fig. 2.116

Fig. 2.117

Fig. 118

Fig. 2.119

Figures 2.109–2.119, Baetidae genn. spp. 2.109–2.111, Pseudocentroptiloides usa: 2.109, labrum, anterodorsal; 2.110, left half of labium and left palp outline, ventral. 2.111, right oretarsus and claw, anterior. 2.112, Procloeon nelsoni labrum, anterodorsal. 2.113–2.114, left half of labium and left palp outline, ventral: 2.113, Anafroptilum album; 2.114, Procloeon nelsoni. 2.115, Cloeon dipterum left labial palp, ventral. 2.116–2.117, Procloeon nelsoni: 2.116, angulate (left) mandible; 2.117, planate (right) mandible. 2.118–2.119, Anafroptilum album: 2.118, angulate (left) mandible; 2.119, planate (right) mandible.

Page 52

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

47(46’)



47’





Maxillary palps three segmented, with segment 3 subequal to, or longer than, segment 2; robust setae appressed along outer margin of cerci not changing in essential form or relative size in distal third of cerci compared to more proximal setae; abdominal sternal color pattern without large transverse band anteriorly on sterna 7–9 or 8–9; hindwingpads present or absent ........................................................................................ 48 [Note: Caution should be taken when using the couplet because the demarcation between maxillary palp segment 2 and 3 may require high magnification and/or special lighting to detect.] Maxillary palps apparently two or three segmented, with segment 3 when detectable less than half length of segment 2; robust setae appressed along outer margin of distal third of cerci (Fig. 2.120) becoming spindle shaped and longer than respective cercal segments compared to setae in more proximal region of cerci; abdominal sterna with marginal transverse band between sterna 7–8 and 8–9 (and sometimes between other sterna); hindwingpads present ..................................................... Procloeon (atypical), 90 [Note: Currently, P. viridoculare is the only species of Procloeon from the coverage area that will key here. Essentially a Procloeon with atypical mandibles, it possesses a highly recognizable unique color pattern among long-clawed baetids. It is also keyed comparatively with other Procloeon herein. Slide mounting and high magnification are recommended for viewing the cerci armature character. Generally if structural characters are in doubt, the presence of the marginal transverse bands in the posterior region of the ventral abdomen, as stated, should ensure identification of P. viridoculare; keep in mind, however, that some fluid-preserved specimens, especially young larvae, are subject to fading.]

48(47) 48’

Hind wingpads present .................................................................................. Anafroptilum, 55 Hind wingpads absent ............................................................................................................. 49

49(48’)

Body with dorsal projections; mature body length less than 5 mm, usually 3.0–4.4 mm; legs (Fig. 2.121) with only sparse stout setae and only in ventral aspect (on side of direction of claw curvature); middle abdominal sterna each darkened at anterior margin and with pair of submedian dark dashes adjacent to posterior margin; lateral abdominal spines absent or limited to small spines only in posterior fourth of segments 8 and 9; caudal filaments with dark subdistal band; within coverage area currently known only from Alabama and North Carolina . ........................................ Anafroptilum (atypical), 55 Body without dorsal projections; mature body length at least 5mm; legs (e.g., Fig. 2.122) heavily armored dorsally and ventrally with numerous stout setae; middle abdominal sterna not marked as above; many relatively long lateral spines on abdominal segments 3–9 or 4–9; caudal filaments without subdistal band . .................................. Neocloeon, 81

49’

Chapter 2 ~ Key to Mayflies

Page 53

Fig. 2.120

Fig. 2.121

Fig. 2.122 Figures 2.120–2.122, Baetidae genn. spp. 2.120, Procloeon viridoculare left cercus, dorsal, 2.121–2.122, right forelegs, anterodorsal: 2.121, Anafroptilum minor; 2.122, Neocloeon triangulifer.

Page 54

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Acentrella species 50(42)

Abdominal terga with posteromedial elevations (best developed on terga 2–6) (Fig. 2.123); elevations sometimes are difficult to detect, but region will appear to have layers of blunt spinules on posterior margin; tibiae and tarsi with relatively few short setae along outer margins; currently known only from North Carolina and Tennessee within the coverage area ....................................................................................... Acentrella barbarae Abdominal terga without posteromedial elevations or associated spinule layers; in the rare cases of there being some appearance of elevations on abdominal terga, then tibiae and tarsi with well-developed row of long, fine setae along outer margin (Fig. 2.124)........... 51

50’ 51(50’)



51’



52(51) 52’ 53(51’) 53’

Marginal setation of forelegs similar to Figs. 2.124, 2.125; cerci relatively uniform in coloration (without distinctive light and dark bands), only sometimes graying distally; mature body 4–6 mm; hindwingpads absent or sometimes represented by scalelike or straplike vestiges . ............................................................................................................. 52 [Note: Marginal setae of the tibia and tarsus are sometimes light colored or translucent and therefore may be difficult to detect.] Marginal setation of forelegs more similar to Fig. 2.126; cerci usually with many light and dark bands (variously distinctive, sometimes not apparent); smaller species, with mature body usually less than 4 mm; hindwingpads not developed, but vestigial and straplike as in Fig. 2.127 ................................................................................................... 53 [Note: Alternation of light and dark subregions of cerci is variable and subject to fading.] Forelegs with tibia and tarsus together relatively long (distinctly longer than femur) (Fig. 2.124); gills not pigmented; abdominal tergum 9 and 10 (and sometimes posterior of 8) lighter compared to terga 7 and 8 (or 7 and anterior of 8); fresh specimens very seldom with reddish coloration .................................................................................. Acentrella turbida [Note: A species complex likely is encompassed by the current concept of A. turbida.] Forelegs with tibia and tarsus together relatively short (about as long as femur) (Figs. 2.125, 2.128); gills pigmented basomedially (Fig. 2.128); abdominal terga 8–10 lighter compared to tergum 7 (Fig. 2.128); fresh specimens sometimes with reddish coloration ............................................................................................ Acentrella nadineae Body and cerci with highly contrasting light and dark areas; gills often with some pigmentation (small blotch in distal half of gill); length of vestigial median caudal filament usually less than basal width of cerci ...................................... Acentrella parvula Body and cerci with much less contrasting light and dark areas, sometimes lacking; gills without pigmentation; length of vestigial median caudal filament usually greater than basal width of cerci ............................................................................... Acentrella alachua [Note: Differentiating larvae of A. parvula and A. alachua can be problematic and is best authenticated by association with male adults.]

Chapter 2 ~ Key to Mayflies

Page 55

Fig. 2.123

Fig. 2.125 Fig. 2.124

Fig. 2.126

Fig. 2.127

Fig. 2.128

Figures 2.123–2.128. Acentrella spp. (Baetidae). 2.123, A. barbarae abdominal terga, left lateral. 2.124–2.126, right forelegs, anterodorsal: 2.124, A. turbida; 2.125, A. nadineae; 2.126, A. parvula. 2.127, A. parvula, left half of metanotum with hind wingpad rudiment. 2.128, A. nadineae, habitus, dorsal.

Page 56

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Acerpenna species

(see also Morihara & McCafferty, 1979a) 54(37) 54’



Gills 7 (Fig. 2.129) teardrop shaped and asymmetrical, with posterior (inner) edge somewhat straightened or slightly concave distally, anterior (outer) margin smooth (not serrate), with or without setules (minute setae), and posterior margin serrate, with or without setules .................................................................................... Acerpenna pygmaea Gills 7 spindle shaped and symmetrical, anterior (outer) and posterior (inner) margins serrate and with setules ............................................................ Acerpenna macdunnoughi [Note: Condition of the gill margin is best studied under compound magnification.]

Anafroptilum species 55(48,49) Hindwingpads present; body without dorsal projections . ...................................................... 56 55’ Hindwingpads absent; body with dorsal projections ................................ Anafroptilum minor 56(55)

Caudal filaments with distal third sometimes appearing somewhat darker than basal two-thirds of caudal filaments, but never with dark band of segments near midlength, nor with dark distal band of segments; known only from Kentucky within the coverage area . ............................................................................................ Anafroptilum bifurcatum [Note: This species is also distinguished by a short spineless projection at the middle of the posterior margin of abdominal tergum 9. This characteristic, however, is shared by other species of Anafroptilum outside the coverage area, including A. semirufum, which is known from Pennsylvania and may eventually be found within the coverage area.] 56’ Caudal filaments with dark band of segments near midlength and often with additional band of dark segments distally . ........................................................................................ 57 57(56’) 57’ 58(57’) 58’

Claws relatively long and narrow, about three-fourths as long as respective tarsi (Fig. 2.130); abdominal sterna 8 and 9 with pair of dark quadrate spots submedially at anterior margin; known only from North Carolina ........................... Anafroptilum sp. A [Note: This undescribed but distinct species is known only from mountain streams of western North Carolina.] Claws not as above, stouter and about half as long or somewhat less than half as long as respective tarsi; abdominal sterna 8 and 9 not marked as above ...................................... 58 Cerci with subdistal region (between wide medial dark band and dark tips) with two or three darkened segments separated from each other by about three pale segments; abdominal sterna and usually entire body covered with brown speckling; within coverage area, known only from Tennessee, but expected from Kentucky . ........ Anafroptilum ozarkense Cerci with subdistal region without interspersed individual dark segments; body usually not covered in brown speckling .......................................................... Anafroptilum album

Chapter 2 ~ Key to Mayflies

Page 57

Fig. 2.129

Fig. 2.130 Figures 2.129–2.130, Acerpenna pygmaea and Anafroptilum sp. (Baetidae). 2.129, Acerpenna pygmaea, left gill 7, left dorsolateral. 2.130, Anafroptilum sp., right foretarsus and claw, anterior.

Page 58

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Apobaetis species 59(33’) 59’

Antenna length subequal to head width; sterna 1–9 each with anteromedial spot ........................................................................................................................... Apobaetis sp. A Antenna length about two times head width; no anteromedial spot on each of abdominal sterna 1–9, but sternum 9 sometimes with anterior pigmentation ........... Apobaetis etowah [Note: One additional species of Apobaetis, A. futilis, is known from Mississippi within the coverage area; its larva remains unknown. Additional new species may occur.]

Baetis species

(key modified from Morihara & McCafferty, 1979a; see also Wiersema et al., 2004) 60(39) 60’



61(60)

Labial palps with medioapical lobe of segment 2 poorly developed (Figs. 2.131–2.133); caudal filaments with dark band of segments near midlength .......................................... 61 [Note: Tail banding is subject to some fading.] Labial palps with medioapical lobe of segment 2 moderately well developed (Figs. 2.134, 2.135); caudal filaments without dark medial band .......................................................... 64

61’

Labial palps with relatively long and narrow segment 2, segment 2 at least about twice as long (outer straight-line distance) as basal width of segment 3 (Fig. 2.133); abdominal tergum 5 extensively pale in lateral and sublateral regions, without pair or triad of pale spots near posterior margin (Fig. 2.136) ........................................................... Baetis pluto Labial palps with relatively short segment 2, segment 2 no more than about 1.25 to 1.5× basal width of segment 3 (Figs. 2.131, 2.132); abdominal tergum 5 not as above, more similar to Figs. 2.137–2.139 ............................................................................................. 62

62(61’) 62’

Triad of pale spots near posterior margin of at least abdominal terga 2–4 (Fig. 2.137); abdominal terga 8–10 usually mostly pale ............................................ Baetis intercalaris Pair of pale spots near posterior margin of at least terga 2–4 (Figs. 2.138, 2.139); tergum 8 partly to mostly dark, tergum 9 pale or mostly pale, and tergum 10 variable (Figs. 2.138, 2.139) ........................................................................................................... 63

63(62’) 63’

Pair of pale spots present near posterior margin on abdominal terga 6 and 7, and sometimes pair coalesced on tergum 8 (Fig. 2.138); terga 9 and 10 generally pale (Fig. 2.138); gill tracheation usually dark and distinctive . .................................................. Baetis flavistriga Pair of pale spots absent near posterior margin on abdominal terga 6 and 7, but sometimes weakly present on tergum 8 (Fig. 2.139); tergum 9 generally lighter than 8, and tergum 10 somewhat darkened (Fig. 2.139); gill tracheation usually not dark and distinctive ...................................................................................................................... Baetis phoebus



Chapter 2 ~ Key to Mayflies

Fig. 2.131

Page 59

Fig. 2.132

Fig. 2.133

Fig. 2.134

Fig. 2.135

Fig. 2.136

Fig. 2.137

Fig. 2.138

Fig. 2.139

Figures 2.131–2.139, Baetis spp. (Baetidae). 2.131–2.135, left labial palp, ventral: 2.131, B. intercalaris; 2.132, B. flavistriga; 2.133, B. pluto; 2.134, B. brunneicolor; 2.135, B. tricaudatus. 2.136, B. pluto, abdominal terga 4–6, dorsal. 2.137–2.138, habiti: 2.137, B. intercalaris; 2.138, B. flavistriga. 2.139, B. phoebus, abdomen, dorsal (Ide, 1937).

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

64(60’)





64’





Antennae with pedicel (second segment) with some short robust setae usually somewhat below distal margin (Fig. 2.140), few short spinelike setae (with narrow, sharp points as in Fig. 2.142) sometimes present near distal margin in addition to robust setae; abdominal sterna with robust setae, developed at least along far lateral edge of sterna 4–9, and often also concentrated in groups in far posterolateral corners of sterna (Fig. 2.143) ................................................................................................................. Baetis tricaudatus [Note: Keep in mind that robust and spinelike setae mentioned above are socketed; those referred to here are very small and may or may not be difficult to detect with a dissecting scope and are best seen in slidemounts under compound magnification. Also note that B. tricaudatus and B. brunneicolor can co-occur.] Antennae with pedicel not as above, most without any socketed setae on pedicel, although clusters of minute triangular microspines (not socketed) sometimes present (Fig. 2.141); occasionally individuals with pedicel with about one to four small spinelike setae (with narrow, sharp points for much of length) along apical edge of pedicel (Fig. 2.142); abdominal sterna without robust setae as above, although occasionally few socketed setae coming from margin of some sterna near lateral edge of sterna . ........................... .............................................................................................................. Baetis brunneicolor [Note: Additional similar differences in the presence and absence of robust setae respectively between B. tricaudatus and B. brunneicolor involve the scape (basal segment) of the antenna and the paraprocts (pair of distal plates at the posteroventral end of the abdomen). Also note that color patterns of larvae of B. brunneicolor and eastern populations of B. tricaudatus often are indistinct.]

Baetopus species 65(35) 65’



Head with pair of distinctive lateral horns (Fig. 2.144); known only rarely from North Carolina . ....................................................................................................Baetopus trishae Head without lateral horns as above; known only rarely from Kentucky.......... Baetopus sp. A [Note: The undescribed species resembles Eurasian species of Baetopus and requires further study.]

Chapter 2 ~ Key to Mayflies

Page 61

Fig. 2.142 Fig. 2.140

Fig. 2.141

Fig. 2.143

Fig. 2.144

Figures 2.140–2.144, Baetis spp. and Baetopus trishae (Baetidae). 2.140–2.143, Baetis spp.: 2.140–2.142, right antennal pedicel, dorsal: 2.140, B. tricaudatus; 2.141, B. brunneicolor variant 1; 2.142, B. brunneicolor variant 2 (distal margin only); 2.143, B. tricaudatus, abdominal sternum 8, ventral. 2.144, Baetopus trishae head, anterior.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Callibaetis species

(see Check, 1982 for more illustrations and a more comprehensive treatment of the Nearctic fauna) 66(29)



66’ 67(66’) 67’

Gills 7 with small ventral lobe or flap (Fig. 2.145); foretibiae dorsally with one to several robust setae; maxillary palp two-segmented, with basal segment length about two times its width and with apical segment length about equal to the width of the basal segment ............................................................................................... Callibaetis pretiosus [Note: Claws curve away from the dorsal aspect of the tibia.] Gills 7 single without ventral lobe or flap (Fig. 2.146); foretibiae dorsally without robust setae, only fine setae at most; maxillary palps not exactly as above ................................ 67 Middle region of caudal filaments contrastingly darkened at every fourth intersegment; distal margin of labrum (Fig. 2.147) straight to slightly rounded in each half; lateral margins of abdominal segments 7 & 8 concave ............................... Callibaetis floridanus Middle region of caudal filaments concolorous, without interspersed color pattern as above; distal margin of labrum (similar to Fig. 2.148) strongly rounded in each half; lateral margins of all abdominal segments relatively straight ............ Callibaetis fluctuans

Heterocloeon species 68(41)

68’

Gill filament present near base of each foreleg (e.g., Fig. 2.149) . ......................................... 74 [Note: Gill filaments at the base of the forelegs may function as osmoregulatory papillae. They sometimes are called osmobranchiae. They can be difficult to detect because they can be translucent to nearly transparent.] Gill filament not present at base of forelegs ........................................................................... 69

69(68’) 69’ 70(69) 70’ 71(69’) 71’

Claws with two rows of denticles, secondary row moderately developed (e.g., Fig. 2.150) or only poorly developed; tibiae usually without forked spatulate setae below dorsal margin on either side . ....................................................................................................... 70 Claws with primary row of denticles and with secondary ridge lacking denticles; tibiae usually with deeply forked spatulate setae on anterior face below dorsal margin (Figs. 2.151, 2.152) ........................................................................................................... 71 Claws with secondary row of robust denticles; known only from near Jellico, Tennessee .............................................................................................. Heterocloeon sp. A Claws with secondary row of minute denticles, sometimes appearing only as small bumps; widespread in eastern North America ................................................... Heterocloeon sp. B Cerci banded; hind wingpads vestigial or absent; labial palps (Fig. 2.153) with outer margin of segments 2 and 3 forming continuous curve, and segment 3 subequal in length to segment 2 ........................................................................................................... 72 Cerci unbanded; hind wingpads at least moderately developed; labial palps (Fig. 154) with outer margin nearly straight, curving only in distal half of segment 3, and segment 3 somewhat shorter than segment 2 .................................................................................. 73



Chapter 2 ~ Key to Mayflies

Page 63

Fig. 2.147 Fig. 2.145

Fig. 2.148

Fig. 2.146

Fig. 2.151

Fig. 2.149

Fig. 2.150

Fig. 2.152

Fig. 2.153

Fig. 2.154

Figures 2.145–2.154, Callibaetis spp. and Heterocloeon spp. (Baetidae). 2.145–2.148, Callibaetis spp.: 2.145–2.146, left gills 7, left dorsolateral (Check, 1982): 2.145, C. pretiosus; 2.146, C. floridanus. 2.147–2.148, labra, anterior: 2.147, C. floridanus (Check, 1982); 2.148, C. sp. 2.149–2.154, Heterocloeon spp.: 2.149, H. berneri, gill filament at right foreleg base, anterior (modified from Müller-Liebenau, 1974). 2.150, H. amplum right foreclaw, anterior (Müller-Liebenau, 1974). 2.151–2.152, H. amplum: 2.151, right hind leg, anterior face, dorsal up; 2.152, specialized setae of a hind tibia, anterior. 2.153–2.154, left labial palp outline, ventral: 2.153, H. grande.; 2.154, H. amplum.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

72(71) 72’

Cerci with band at mid-length or slightly proximal; gills nearly symmetrical in shape; abdominal sterna with no lateral tracheation marks .......................... Heterocloeon grande Cerci with band distad of mid-length; gills assymetrical in shape; abdominal sterna with prominent lateral tracheation marks; currently known only from Kansas . ..................... ............................................................................................................... Heterocloeon sp. C

73(71’) 73’

Median caudal filament with 3–5 (very rarely 6) segments; widespread geographic distribution ....................................................................................... Heterocloeon amplum Median caudal filament with 8 or more segments; currently known only from Ohio and Indiana, but extremely likely to occur in Kentucky . ............................ Heterocloeon sp. D

74(68) 74’

Distinct bristled protuberances present ventrally on second and third thoracic segments, and on abdominal sternum 1 and somewhat on sternum 2 (Fig. 2.155) .......................... ........................................................................................................... Heterocloeon berneri Ventral protuberances not present as above ............................................................................ 75

75(74’)



Mature body length 4–5 mm; hindwingpads small and straplike as in Fig. 2.156; secondary row of claw denticles poorly developed (sometimes requiring slide mount for viewing); forefemora with moderately developed, dense marginal row of setae (Fig. 2.158); within coverage area currently known only from Tennessee . ........ Heterocloeon frivolum Mature body length 6–8 mm (usually 7–8 mm); hindwingpads minute as in Fig. 2.157 or smaller; secondary row of claw denticles relatively well developed (similar to Fig. 2.159); forefemora with marginal setal row not as long as above and less dense than above at least in some portions ......................................................................................... 76

76(75’) 76’

Labrum (Fig. 2.160) with row of 6–10 distinct submarginal setae on either half (not including isolated medial seta); abdominal terga with distinct color patterning; abdominal gills diffusely blackened medially ............................... Heterocloeon curiosum Labrum (Fig. 2.161) with row of 3–5 distinct submarginal setae on either half (not including isolated medial seta); abdominal terga without distinct color patterning; abdominal gills essentially grayish ...................................................................... Heterocloeon petersi

75’

Iswaeon species 77(41’) 77’

Abdominal terga with extensive medial spotting as seen in Fig. 2.162 . ........... Iswaeon davidi Abdominal terga without medial spotting, or spot limited to tergum 2 or to 2 and 6 (sometimes spots appearing divided) . ......................................................... Iswaeon anoka [Note: Two of the three species of Iswaeon are known as larvae; I. rubrolaterale remains unknown in this regard.]

Chapter 2 ~ Key to Mayflies

Page 65

Fig. 2.156

Fig. 2.157

Fig. 2.155

Fig. 2.158

Fig. 2.159

Fig. 2.162 Fig. 2.160

Fig. 2.161

Figures 02.155–2.162, Heterocloeon spp. and Iswaeon davidi (Baetidae). 2.155–2.161, Heterocloeon spp.: 2.155, H. berneri, head, thorax, and base of abdomen, left lateral (Müller-Liebenau, 1974); 2.156–2.157, left hind wingpad and left half of metanotum, dorsal: 2.156, H. frivolum; 2.157, H. curiosum 2.158, H. frivolum, right forefemur, anterodorsal. 2.159, H. curiosum, right foreclaw, anterior (MüllerLiebenau, 1974). 2.160–2.161, right halves of labra, dorsal (modified from Müller-Liebenau, 1974): 2.160, H. curiosum; 2.161, H. petersi. 2.162, Iswaeon davidi, abdomen, dorsal.

Page 66

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Labiobaetis species

(key modified from McCafferty & Waltz, 1995) 78(38) 78’



Labrum submarginal setae spatulate (Fig. 2.163), often with broad distal margin appearing fringed ................................................................................................ Labiobaetis frondalis Labrum submarginal setae not spatulate (Figs. 2.164, 2.165) ................................................ 79 [Note: Labiobaetis longipalpus, which does not occur in the coverage area but does occur in surrounding areas, has larvae that are atypical within the genus; they have very long maxillary palps and do not have the broad medial lobe on the labial palps found in other North American species.]

79(78’) 79’

Labrum submarginal setae simple (Fig. 2.164) ................................... Labiobaetis propinquus Labrum submarginal setae branched as in Fig. 2.165 . ........................................................... 80

80(79’) 80’

Planate (right) mandible with peaked prominence between mola and incisors (Fig. 2.166) ....................................................................................................... Labiobaetis ephippiatus Planate mandible either without prominence between mola and incisors or with only slightly rounded prominence similar to Fig. 2.167 ........................... Labiobaetis dardanus

Neocloeon species

(see Funk et al., 2006 for more details) 81(49’)

Gills with dark pigmentation of some trachea broken up (not pigmented) at about twothirds distance from base and at base of basal two or three tracheal side-branches (Fig. 2.169); femora with dark subdistal band or partial band (Fig. 2.168); parthenogenetic species ............................................................................................. Neocloeon triangulifer 81’ Gills with dark pigmentation broken only near base of basal two or three tracheal branches (Fig. 2.170); femora without dark subdistal band, sometimes faint brown mark at about mid-length of femora; sexual species ................................................ Neocloeon alamance

Paracloeodes species

(see McCafferty & Lenat, 2003 for additional comparative treatment) 82(34) 82’



Gills 2 usually 1.5× or less length of abdominal tergum 2; at least terga 2–7 (Fig. 2.171) with dark lateral edge spots at about midlength of terga, and at least terga 6–7 with medial spot and pair of submedian spots near anterior margin; abdominal sterna sometimes with broad transverse intersegmental banding .............. Paracloeodes minutus Gills 2 usually between 2 and 3.5× length of abdominal tergum 2; tergal edge spots not developed as above and terga 6–7 without three anterior spots as above, diffuse clouding at most; never with much or any spotting in pleural area of abdomen, and abdominal tergum 2 with no medial pale spot or with diffuse pale central area only; abdominal sterna at most with thin dark pencil lines at segment margins ............. Paracloeodes fleeki [Note: Gill measurements may not always be reliable in less than nearly mature individuals; in early instar larvae, color pattern characters may become unreliable.]

Chapter 2 ~ Key to Mayflies

Fig. 2.163

Page 67

Fig. 2.164

Fig. 2.165

Fig. 2.169 Fig. 2.166

Fig. 2.170

Fig. 2.167

Fig. 2.168 Fig. 2.171 Figures 2.163–2.171, Labiobaetis spp., Neocloeon spp., and Paracloeodes minutus (Baetidae). 2.163–2.167, Labiobaetis spp.: 2.163–2.165, labra, anterodorsal: 2.163, L. frondalis; 2.164, L. propinquus; 2.165, L. ephippiatus. 2.166–2.167. planate (left) mandibles, ventral: 2.166, L. ephippiatus; 2.167,. L. sp. 2.168, Neocloeon triangulifer, right foreleg, anterodorsal. 2.169–2.170, right gills 4, right dorsolateral: 2.169, N. triangulifer; 2.170, N. alamance. 2.171, Paracloeodes minutus, abdominal terga 5–7, dorsal.

Page 68

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Plauditus species 83(42’)

83’

Antennae relatively very short, about as long as vertical head, and less than 2× width (see previous Fig. 2.29); abdominal segment 5 often contrastingly dark (Fig. 2.172) (sometimes segment 5 only partially darkened and sometimes segment 9 darkened instead of, or in addition to, segment 5) .................................................... Plauditus cestus [Note: This key includes all known larvae from the coverage area, plus P. texanus (from Texas and Oklahoma). The recently discovered larva of P. elliotti (from Minnesota and Manitoba) is most similar to P. dubius (except for having yellow-orange flushing of abdomen and gills) and is the only known larva of the genus that is not included. The larva of P. veteris is not known. The key includes revised concepts of some larvae based on our study of type specimens, rearing experiments, and recent molecular work. The key, however, may need future refinement.] Antennae at least 1.5× length of head and usually more than 2× width, often over 2× length and 3× width of head ........................................................................................................ 84

84(83’) 84’

Abdominal sterna with dark medial marginal spot appearing at least at margin of sterna 5–6 and/or 6–7, often involving most sterna (Figs. 2.173, 2.174) . ......................................... 85 Abdominal sterna without medial spots . ................................................................................ 88

85(84) 85’

Gills 7 darkened at least in basal third; abdominal sterna usually reddish brown .......................................................................................................... Plauditus bimaculatus Gills 7 pale or slightly tinged, and not differing from gills 1–6; abdominal sternum seldom reddish brown ................................................................................................................... 86

86(85’) 86’

Abdominal sterna without lateral tracheation marks (Fig. 2.173) and with one pair of submedian dots ........................................................................................ Plauditus gloveri Abdominal sterna with lateral tracheation marks (e.g., Fig. 2.174) and with one pair, two pair or no pairs of submedian dots or some combination of submedian dot variations . ..... 87

87(86’) 87’

Ventral abdomen with total of one or two median marginal spots present, sometimes very diffuse and sometimes difficult to detect ................................................... Plauditus virilis Ventral abdomen with four or usually more median marginal spots present and generally distinctive (e.g., Fig. 2.174) ........................................................... Plauditus punctiventris

88(84’) 88’

Cerci without middle band of dark segments, but with long hairlike tail setae darker in midregion of cerci (Fig. 2.175); abdominal terga with relatively little patterning in both sexes (Fig. 2.175) . ............................................................................. Plauditus cingulatus Cerci with prominent band of dark segments in midregion; abdomen with contrasting patterns (Fig. 2.176), usually most pronounced in males ................................................. 89

89(88’) 89’

Some abdominal sterna with lateral tracheation marks; legs distinctly banded (Fig. 2.176); widespread distribution . ........................................................................... Plauditus dubius Abdominal sterna without lateral tracheation marks; legs only faintly banded; not currently known from the coverage area . .............................................................. Plauditus texanus





Chapter 2 ~ Key to Mayflies

Fig. 2.172

Fig. 2.173

Page 69

Fig. 2.174

Fig. 2175

Fig. 2.176

Figures 2.172–2.176, Plauditus spp. (Baetidae). 2.172, P. cestus, abdominal terga, dorsal. 2.173–2.174, abdominal sterna, ventral: 2.173, P. gloveri; 2.174, P. punctiventris variant. 2.175, P. cingulatus, abdominal terga and cerci, dorsal (Ide 1937). 2.176, P. dubius variant, habitus, dorsal.

Page 70

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Procloeon species 90(46,47’) Lateral spines present on abdominal segments 8–9 as seen in dorsal view (Fig. 2.177), only rarely additionally with one to four spines also on segment 7; hindwingpads present .... 91 [Note: In addition to the seven nominal species from the coverage area that are known as larvae, this key includes an unnamed species (sp. A) that occurs commonly in the coverage area. Procloeon simplex and another unnamed species (sp. B) also are included; both are known from the Southeast and eventually should be found in the coverage area.] 90’ Lateral spines present on at least abdominal segments 5–9, often also on segments 2–4; hindwingpads present or absent ........................................................................................ 94 91(90)

91’

Gills simple, without dorsal flap; not known from coverage area ................... Procloeon sp. B [Note: This unnamed species is distinct and has been taken in Arkansas, Kansas, Iowa, and Virginia. Thus, it likely will be found in the coverage area.] Gills with dorsal flap (e.g., Fig. 2.178) on some or most abdominal segments (size of flap somewhat variable) ........................................................................................................... 92

92(91’) 92’

Gills 1 only with dorsal flap; caudal filaments without wide medial band of darkened segments ...................................................................................... Procloeon rufostrigatum Gills 1–3 (rarely), 1–4, 1–5 or 1–6 with dorsal flap; caudal filaments with wide medial band of darkened segments . ...................................................................................................... 93

93(92’) 93’

Gills 1–3, 1–4 or 1–5 with dorsal flap; claws without denticles or microdenticles ................................................................................................................ Procloeon rivulare Gills 1–6 with dorsal flap; claws (Fig. 2.179) with microdenticles and usually some (larger) denticles; within coverage area known only from Tennessee and South Carolina ......................................................................................................... Procloeon pennulatum [Note: Claws generally require slide mounting to view denticulation.]

94(90’) 94’

Hindwingpads present; robust setae appressed along outer edge of cerci becoming longer than respective cercal segments and becoming thickened and spindle shaped in distal third of cerci (e.g., Fig. 2.180) .......................................................................................... 95 [Note: Robust setae of cerci are best viewed under compound magnification with comparisons along the entire length of cerci.] Hindwingpads absent; cerci distally with outer setae not enlarging as above ........................ 99

95(94) 95’

All gills simple, without dorsal flap; claws two-thirds of, to subequal to, respective tarsal length ................................................................................................................................ 96 Gills 1–4, 1–5 or 1–6 with dorsal flap; tarsal claws about one-half to two-thirds of respective tarsal length . ..................................................................................................................... 98



Chapter 2 ~ Key to Mayflies

Page 71

Fig. 2.178

Fig. 2.177

Fig. 2.179

Fig. 2.180

Figures 2.177–2.180, Procloeon spp. (Baetidae). 2.177, P. rufostrigatum, abdominal terga 6–10, dorsal. 2.178, P. sp., right gill 1, right dorsolateral. 2.179, P. pennulatum, right foreclaw, anterior. 2.180, P. viridoculare, left cercus, dorsal.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

96(95) 96’



97(96’) 97’



Planate (right) mandible with incisors separated at least in distal half; abdominal sterna with transverse brown bands at margins between sterna 7–8 and 8–9 (most prominent between 7–8) as in Fig. 2.181, sometimes also weakly present on preceding sterna involving 4–7 and also between sterna 9–10, and usually with prominent pair of sublateral dark spots on some sterna ................................................................................ Procloeon viridoculare [Note: Fluid-preserved specimens are subject to fading.] Planate mandible with incisors fused in much of distal half (e.g., Fig. 2.182); abdominal sterna not marked as above ............................................................................................... 97 Abdominal terga 3–8 or 3–9 (Fig. 2.183) each with pair of dark submedian dots, and lateral margins of each tergum with darkened areas at about midlength; tergum 10 and some anterior terga with dark medial marking; femora with dark medial markings on outer face . ................................................................................................ Procloeon fragile Abdominal terga 3–9 without paired submedian dots, and lateral margins of terga without darkened areas as above; tergum 10 without dark medial marking as above, but anterior terga often with medial marking; femora without dark medial markings as above (sometimes faint coloration present) . ......................................................... Procloeon sp. A [Note: Procloeon sp. A is generally less common than P. fragile but is known from more states in the coverage area; the two species often co-occur.]

98(95’) 98’

Gills 1–6 with dorsal flaps; maxillary palps three segmented; within coverage area currently known only from South Carolina ...................................................... Procloeon quaesitum Gills 1–4 or 1–5 with dorsal flaps; maxillary palps two segmented; in the Southeast known only from Tennessee ............................................................................... Procloeon nelsoni

99(94’)

Dorsal flap usually on gills 1–6 (flap of gill 6 very small or occasionally absent); abdominal terga uniformly greenish (fresh specimens) or yellowish (degraded or mounted specimens) with pale submedian oblique markings followed by pair of pale submedian dots and small sublateral pale areas (Fig. 2.184) (relatively large medial black spot sometimes present on some terga); not currently known from coverage area ................................................................................................................ Procloeon simplex [Note: Within the Southeast, this species is known from Louisiana and Arkansas and may eventually be found in the coverage area.] Dorsal flap on gills 1 only; abdominal terga not marked as above but with ornate light and dark patterning ............................................................................... Procloeon rubropictum

99’



Chapter 2 ~ Key to Mayflies

Page 73

Fig. 2.182

Fig. 2.181

Fig. 2.184

Fig. 2.183 Figures 2.181–2.184, Procloeon spp. (Baetidae). 2.181, P. viridoculare, abdominal sterna, ventral. 2.182, P. rubropictum, planate (left) mandible, ventral. 2.183, P. fragile, abdominal terga 6–10, dorsal. 2.184, P. simplex, abdominal terga 5–6, dorsal.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

BAETISCIDAE, Baetisca species

(see Pescador & Berner, 1981 for additional details) 100(17) 100’

Genal spines absent (anterolateral corners of head capsule not developed into spinelike processes) although frontal (anterior medial) process well developed in some (Figs. 2.185–2.190) ......................................................................................................... 101 Genal spines present (anterolateral corners of head developed into spinelike processes) (Figs. 2.191–2.194) ......................................................................................................... 106

101(100) Frontal process of head strongly produced anteriorly into large bifurcate process as in Fig. 2.185 ...................................................................................................................... Baetisca obesa 101’ Frontal process of head not strongly produced anteriorly as above (Figs. 2.186–2.190) ..... 102 102(101’) Lateral spines (pair of processes) of thoracic carapace (carapace) bluntly truncated or roundly pointed and relatively short (e.g., Fig. 2.186); carapace relatively broad (Fig. 2.186) ............................................................................................... Baetisca gibbera 102’ Lateral spines of thoracic carapace pointed (although subject to some wear) and variable in relative length (Figs. 2.187–2.190) (if short with rounded tip then carapace relatively narrow as in e.g., Fig. 2.189); carapace variable in relative width (Figs. 2.187–2.190) .... 103 103(102’) Carapace (Fig. 2.187) moderately broad (length to width ratio 1.3–1.0), with lateral spines elongate with relatively narrow base ............................................................ Baetisca becki 103’ Carapace variable (length to width ratio 1.6–1.0 to 1.2–1.0) with lateral spines not narrow elongate, either shorter or with broad base as (Figs. 2.188–2.190) ................................ 104 104(103’) Carapace (Fig. 2.188) broad (length to width ratio of 1.2–1.0), with lateral spines long with relatively broad base; posteromedian elevation and posterolateral processes on abdominal tergum 9 well developed .......................................................... Baetisca rogersi 104’ Carapace (Figs. 2.189, 2.190) relatively narrow (length to width ratio 1.6–1.0 to 1.4–1.0), with lateral spines relatively short to moderately long; posteromedian elevation and posterolateral processes of tergum 9 poorly developed .................................................. 105 105(104’) Carapace (Fig. 2.189) narrow (length to width ratio 1.6–1.0), with lateral spines relatively short with variable tip; in posterior half of carapace pair of dorsal projections prominent and divergent and higher than posterior medial rise (medial hump) . .... Baetisca laurentina 105’ Carapace (Fig. 2.190) somewhat broader (length to width ratio 1.4–1.0), with lateral spines moderately long and sharply pointed; in posterior half of carapace pair of dorsal projections poorly developed, being distinctly lower than posterior medial rise (medial hump) . ........................................................................................ Baetisca lacustris

Chapter 2 ~ Key to Mayflies

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Fig. 2.185

Fig. 2.188

Fig. 2.186

Fig. 2.189

Fig. 2.187

Fig. 2.190

Figures 2.185–2.190, Baetisca spp. (Baetiscidae), habiti, dorsal (Pescador & Berner 1981). 2.185, B. obesa; 2.186, B. gibbera; 2.187, B. becki; 2.188, B. rogersi; 2.189, B. laurentina; 2.190, B. lacustris.

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106(100’) Abdominal segments 6–9 with long, dense hairlike setae laterally (Fig. 2.191); genal spines very well developed (Fig. 2.191) ..................................................... Baetisca escambiensis [Note: In fresh specimens, the eyes of this species are diagnostically striped.] 106’ Abdominal segments 6–9 with at most few minute setae laterally (Figs. 2.192–2.194); genal spines moderately developed (Figs. 2.192–2.194) . .............................................. 107 107(106’) Lateral spines of thoracic carapace blunt as in Fig. 2.192 .......................... Baetisca rubescens [Note: This more typically northeastern species has not been reported from the coverage area, but it is found in nearby Virginia and West Virginia.] 107’ Lateral spines of thoracic carapace sharply pointed (Figs. 2.193, 2.194) . ........................... 108 108(107’) Lateral spines moderately long (with length about 1.5× basal width) (Fig. 2.193); caudal filaments not darkened in basal fourth (Fig. 2.193) ................................. Baetisca carolina 108’ Lateral spines short (with length subequal to basal width) (Fig. 2.194); caudal filaments darkly banded in basal fourth (Fig. 2.194) . ............................................... Baetisca berneri [Note: B. carolina and B. berneri are either very closely related species or represent variants of B. carolina.]

CAENIDAE genera

(see Sun & McCafferty, 2008 for complete treatment of Brachycercinae) 109(20) 109’

Head without ocellar tubercles (Fig. 2.195); distance between base of forelegs about half or less of distance between bases of second pair of legs ................................................ 110 [Note: Basal leg separation should be viewed with legs spread laterally.] Head with three distinct ocellar tubercles ranging from narrowly acute to broadly rounded (Figs. 2.196, 2.197); distance between base of forelegs more than half of distance between bases of second pair of legs .............................................................................. 111

110(109) Inner margin of foretibiae edged variously with spines or short spines, and sometimes with sparse long setae also (Figs. 2.198, 2.199); operculate gills surface bare or with few hairlike setae, and with posterior margin edged with long hairlike setae (Figs. 2.200, 2.201) ........................................................................................... Caenis, 115 110’ Inner margin of foretibiae with brush of long setae (Fig. 2.202); operculate gills surface covered with short, spatulate setae, with posterior margin edged with short, spatulate setae, not hairlike setae (Fig. 2.203) ............................................... Amercaenis, A. cusabo 111(109’) Operculate gills asymmetrical, with posterolateral corner protruding compared to posteromedian corner (Fig. 2.204) . ....................................................... Brachycercus, 114 111’ Operculate gills not as above (Figs. 2.205, 2.206) ............................................................... 112

Chapter 2 ~ Key to Mayflies

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Fig. 2.191

Fig. 2.192

Fig. 2.196

Fig. 2.193

Fig. 2.198

Fig. 2.194

Fig. 2.199

Fig. 2.197 Fig. 2.202

Fig. 2.195

Fig. 2.200

Fig. 2.201

Fig. 2.204

Fig. 2.205

Fig. 2.203

Fig. 2.206

Figures 2.191–2.206, Baetisca spp. (Baetiscidae) and Caenidae genn. spp. 2.191–2.194, Baetisca spp. (Baetiscidae), habiti, dorsal (Pescador & Berner, 1981): 2.191, B escambiensis; 2.192, B. rubescens; 2.193, B. carolina; 2.194, B. berneri. 2.195–2.206, Caenidae genn. spp.: 2.195, Caenis macafferti, habitus, dorsal. 2.196–2.197, heads, dorsal: 2.196, Brachycercus sp. (modified from Soldán, 1986); 2.197, Cercobrachys sp. 2.198–2.201, Caenis spp.: 2.198–2.199, right forelegs, anterodorsal: 2.198, C. amica; 2.199, C. hilaris; 2.200–2.201, left operculate gills, left dorsolateral: 2.200, C. hilaris; 2.201, C. tardata. 2.202–2.203, Amercaenis cusabo: 2.202, right foreleg, anterodorsal; 2.203, left operculate gill, left dorsolateral. 2.204–2.206, Brachycercinae left operculate gills, left dorsolateral: 2.204, Brachycercus nitidus; 2.205, Sparbarus lacustris; 2.206, Cercobrachys etowah.

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112(111’) Medial processes present on both meso- and metasternum (venter of second and third thoracic segments) (Fig. 2.207) ...................................................... Susperatus, S. prudens [Note: Although this species has been reported from Alabama, we cannot confirm its presence in the area of coverage; it is known mostly from northwestern North America.] 112’ Medial processes absent on meso- and metasternum ........................................................... 113 113(112’) Abdominal segment 6 posterolateral projection apically directed posteriorly, not curved medially (Fig. 2.208) .................................................................................. Sparbarus, 124 113’ Abdominal segment 6 posterolateral projection apically curving toward midline of abdomen (Fig. 2.209) ............................................................................................ Cercobrachys, 123

Brachycercus species

(modified from Sun & McCafferty 2008) 114(111) Hindclaws more than one-fourth of combined length of hindtibiae and hindtarsi (Fig. 2.210) ....................................................................................... Brachycercus berneri 114’ Hindclaws less than one-fourth of combined length of hindtibiae and hindtarsi (Fig. 2.211) ............................................................................................................ Brachycercus nitidus

Caenis species

(see Provonsha, 1990 and Pescador & Richard, 2006 for more complete, comparative treatments) 115(110) Dorsal face of forefemora with transverse row of spatulate setae in distal half (e.g., Fig. 2.212); posterior margin of abdominal sternum 9 strongly to barely notched medially (Figs. 2.213, 2.214); abdominal terga 9 and 10 without triad of black dots; abdominal sterna without submedian pairs of black dots ................................................................. 116 115’ Dorsal face of forefemora with scattered simple setae only (e.g., Fig. 2.215); posterior margin of abdominal sternum 9 convexly rounded or straight (Figs. 2.216, 2.217); abdominal terga 9 and 10 usually with triad of black dots; abdominal sterna often with pairs of submedian black dots . .................................................................................................... 119 116(115) 116’ 117(116’) 117’

Middle third of caudal filaments (tails) with whorls of short setae on every second or third segment, with setae much shorter than distance between base of whorls (Fig. 2.218) ....................................................................................................................... Caenis anceps Middle third of caudal filaments with long lateral setae on every segment, with setae much longer than distance between their bases ........................................................................ 117 Posteromedian projection of abdominal tergum 2 narrowly triangular (viewed dorsally), at least twice as long as wide (Fig. 2.219); abdominal terga 7–9 uniformly brown ................................................................................................................. Caenis macafferti Posteromedian projection of abdominal tergum 2 broadly triangular (viewed dorsally), width subequal to length (Fig. 2.220); abdominal terga 7–9 pale medially with brown longitudinal stripes or blotches laterally ......................................................................... 118

Chapter 2 ~ Key to Mayflies

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Fig. 2.208

Fig. 2.210

Fig. 2.207 Fig. 2.209

Fig. 2.212

Fig. 2.213

Fig. 2.211

Fig. 2.215

Fig. 2.217

Fig. 2.214

Fig. 2.218

Fig. 2.216

Fig. 2.219

Fig. 2.220

Figures 2.207–220, Caenidae genn. spp. 2.207, Susperatus prudens, thorax, left lateral; 2.208–2.209, left halves of abdominal terga 5–6, dorsal: 2.208, Sparbarus nasutus; 2.209, Cercobrachys etowah. 2.210–2.211, Brachycercus spp., right hind tibiae, tarsi, and claws, posterior: 2.210, B. berneri; 2.211, B. nitidus. 2.212–2.220, Caenis spp.: 2.212, C. anceps, right foreleg, anterodorsal. 2.213–2.214, abdominal sterna 8–9, ventral: 2.213, C. hilaris; 2.214, C. macafferti. 2.215, C. amica right foreleg, anterodorsal.; 2.216–2.217, abdominal sterna 8–9, ventral: 2.216, C. amica; 2.217, C. latipennis; 2.218, C. anceps, median caudal filament middle section, dorsal. 2.219–2.220, abdominal terga 2, dorsal: 2.219, C. macafferti; 2.220, C. hilaris.

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118(117’) Y-ridge of operculate gills forked at, or somewhat basad of, midlength of gill, and with median branch with five to six setae (Fig. 2.221); operculate gills with brown and white patterns . ......................................................................................................... Caenis hilaris 118’ Y-ridge of operculate gills forked distad of midlength of gill, median branch with 10–12 setae (Fig. 2.222); operculate gills uniformly brown . ................................. Caenis tardata 119(115’) Posterior margin of abdominal sternum 9 straight (Fig. 2.223); hindfemora with inner marginal setae almost as long as outer marginal setae (Fig. 2.224); thorax usually with mottled pattern ......................................................................................... Caenis latipennis 119’ Posterior margin of abdominal sternum 9 convexly rounded (Fig. 2.225); hindfemora with inner marginal setae much shorter than outer marginal setae (e.g., Fig. 2.226); thorax usually uniformly colored ............................................................................................... 120 120(119’) Posteromedian projection of abdominal tergum 2 moderately long and almost vertical (Fig. 2.227); lateral margins of pronotum strongly divergent anteriorly from posterior margin as seen in dorsal view (Fig. 2.228); known from Florida only .................................................................................................................. Caenis eglinensis 120’ Posteromedian projection of abdominal tergum 2 curved somewhat as seen in lateral view (e.g., Fig. 2.229); lateral margins of pronotum generally subparallel in dorsal view, if bulging anteriorly, never appearing as above ................................................................. 121 121(120’) Hindclaws with 9–11 denticles of uniform size or progressively larger toward tip (Fig. 2.230) .............................................................................................................................. Caenis amica 121’ Hindclaws with 16–20 denticles, with basal four to six denticles larger than other denticles (Figs. 2.231, 2.232) ......................................................................................................... 122 122(121’) Base of antennae (scape and pedicel) usually stained with dark brown; hindtarsi with row of 9–11 fimbriate spurs on ventral face . ................................................... Caenis diminuta [Note: Fimbriate spurs are covered with small branches.] 122’ Base of antennae pale; hindtarsi with row of six to eight fimbriate spurs on ventral face .................................................................................................................... Caenis punctata

Cercobrachys species

(modified from Sun & McCafferty, 2008) 123(113’) Foretarsi with row of long and short setae along inner margin (Fig. 2.233); maxillae galealacinia somewhat sickle shaped (Fig. 2.234) . .......................... Cercobrachys etowah [Note: The galealacinia is the main body of the maxilla.] 123’ Foretarsi with row of only long setae along inner margin; maxillae galealacinia relatively short, nearly triangulate (Fig. 2.235) .............................................. Cercobrachys pomeiok

Chapter 2 ~ Key to Mayflies

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Fig. 2.221

Fig. 2.224

Fig.2.222 Fig. 2.222 Fig. 2.223

Fig. 2.227

Fig. 2.230

Fig. 2.233

Fig. 2.226

Fig. 2.225 Fig. 2.223

Fig. 2.225

Fig. 2.228

Fig. 2.229

Fig. 2.231

Fig. 2.232

Fig. 2.234

Fig. 2.235

Figures 2.221–235, Caenis spp. and Cerobrachys spp. (Caenidae). 2.221–2.232, Caenis spp.: 2.221–2.222, left operculate gills, left dorsolateral: 2.221, C. hilaris; 2.222, C. tardata. 2.223–2.224, C. latipennis: 2.223, sterna 8–9; 2.224, right hind leg, anterior face, dorsal up. 2.225–2.226, C. amica: 2.225, sterna 8–9; 2.226, right hind leg, anterior face, dorsal up. 2.227–2.228, C. eglinensis (Pescador & Richard, 2006): 2.227, abdominal segments1–5 (segment 2 with posterior dorsomedian projection), left lateral; 2.228, pronotum. 2.229, C. amica, abdominal segments 1–5 (segment 2 with posterior dorsomedian projection), left lateral. 2.230–232, right hindclaws, anterior: 2.230, C. amica; 2.231, C. diminuta; 2.232, C. punctata. 2.233–2.235, Cercobrachys spp.: 2.233, C. etowah, left foreleg, anterodorsal. 2.234–235, left maxilla, ventral: 2.234, C. etowah; 2.235, C. sp.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Sparbarus species

(modified from Sun & McCafferty, 2008) 124(113) Middle ocellar tubercle at least 2× length of lateral tubercles ..................... Sparbarus nasutus 124’ Middle ocellar tubercle subequal to, or shorter than, lateral ocellar tubercles ..................... 125 125(124’) Middle ocellar tubercle short-triangulate, with length and basal width subequal; lateral projection of abdominal tergum 2 not developed, with length less than basal width and more or less straight sided laterally ................................................................................ 126 125’ Middle ocellar tubercle relatively elongate, with length 1.5–2.0× basal width; lateral projection of abdominal tergum 2 moderately developed and triangulate (pointed laterally) .......................................................................................................................... 127 126(125) Abdominal tergum 8 without mid-longitudinal brown stripe (Fig. 2.236); femora with black apical marking dorsally; known from Florida only . ............. Sparbarus miccosukee 126’ Abdominal tergum 8 with mid-longitudinal brown stripe (Fig. 2.237); femora without black apical marking dorsally ........................................................... Sparbarus maculatus 127(125’) Posterolateral projections on abdominal tergum 7 about two-thirds length of posterolateral projections on abdominal tergum 6; known from Mississippi and Louisiana only ............................................................................................................... Sparbarus choctaw 127’ Posterolateral projections on abdominal tergum 7 half or less length of posterolateral projections on abdominal tergum 6 ..................................................... Sparbarus lacustris

EPHEMERELLIDAE genera

(see Jacobus & McCafferty, 2008 for comprehensive treatment of genera) 128(18) 128’

Gills present on abdominal segment 3 .................................................................................. 129 Gills absent on abdominal segment 3 ................................................................................... 134

129(128) Forefemora with forward edge jagged or roughened with spurs or spines (e.g., Fig. 2.238) ........................................................................................................................ Drunella, 139 129’ Forefemora with forward edge not as above ........................................................................ 130 130(129’) Abdominal terga with pairs of large arching spines (Fig. 2.239); forefemora without robust or bristlelike setae on upper face ............................................................. Tsalia, T. berneri 130’ Abdominal terga without large arching spines as above; forefemora almost always with at least some bristlelike or large robust setae on upper face (e.g., Fig. 2.240) ................... 131 131(130’) Forefemora narrow, nearly as narrow as foretibiae (Fig. 2.241); abdominal terga with single posteromedian spine ........................................................... Penelomax, P. septentrionalis 131’ Forefemora at least about twice a broad as foretibiae (e.g., Fig. 2.240); abdominal terga with pair of submedian spines or lacking spines entirely ............................................... 132

Chapter 2 ~ Key to Mayflies

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Fig. 2.236

Fig. 2.237

Fig. 2.238

Fig. 2.240 Fig. 2.239

Fig. 2.241

Figures 2.236–2.241, Sparbarus spp. (Caenidae) and Ephemerellidae genn. spp. 2.236–2.237, Sparbarus spp. (Caenidae), abdominal terga 8: 2.236, S. miccosukee; 2.237, S. maculatus. 2.238–2.241, Ephemerellidae genn. spp.: 2.238, Drunella lata, right foreleg, anterodorsal. 2.239, Tsalia berneri (modified from Allen & Edmunds, 1965), abdomen, dorsal. 2.240, Ephemerella subvaria (Allen & Edmunds, 1965), right foreleg, anterodorsal. 2.241, Penelomax septentrionalis, habitus, dorsal.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

132(131’) Caudal filaments (tails) with well-developed interfacing hairlike setae (e.g., Fig. 2.242); maxillary palps developed (e.g., Fig. 2.243); main part of ventral lamella of abdominal gills 6 not deeply cleft in the middle, when viewed from the dorsal aspect..................... ................................................................................................................. Ephemerella, 145 132’ Caudal filaments without well-developed interfacing hairlike setae as above, sparse at most; maxillary palps not present or only poorly developed (e.g., Fig. 2.244); ventral lamella of abdominal gills 6 deeply cleft in middle when viewed from the dorsal aspect, with cleft extending most of length of lamella ....................................................................... 133 133(132’) Abdominal terga without paired posterior submedian spines; maxillary palps not present ..................................................................................................... Teloganopsis, T. deficiens 133’ Abdominal terga with paired posterior submedian spines; maxillary palps present ...................................................................................................................... Serratella, 165 134(128’) Gills 4 not operculate (each covering less than half of gill 5) (Fig. 2.245); abdominal posterolateral projections longer than midlength of their respective terga on only two or fewer terga ..................................................................................................... Attenella, 136 [Note: Numbered gills correspond with the abdominal segment to which they are attached; thus, gills 4 refers to the pair of gills on abdominal segment 4.] 134’ Gills 4 more or less operculate (covering more than half of gills 5), reaching to about posterior margin of tergum 7 or beyond (Figs. 2.246, 2.247); abdominal posterolateral projections longer than midlength of their respective terga on three or more terga ....... 135 135(134’) Abdominal tergum 9 greatly elongated compared to tergum 7 as in Fig. 2.246; claws with denticles; maxillae without palp ............................................................. Eurylophella, 154 135’ Abdominal tergum 9 not greatly elongated compared to tergum 7 as in Fig. 2.247; claws without denticles; maxillae with palp ........................................................... Dannella, 137

Attenella species

(see Jacobus & Fleek, 2010 for additional comparative illustrations) 136(134) Pronotum with well-developed pair of spines (Fig. 2.248); head between eyes appearing twin peaked in anterior view (Fig. 2.249) . ........................................... Attenella attenuata 136’ Pronotum with poorly developed pair of spines; head between eyes appearing slightly convex or biconvex at most (e.g., Fig. 2.250); within the Southeast known only from North Carolina ..................................................................................... Attenella margarita

Dannella species

(modified from McCafferty, 1977a) 137(135’) Abdominal terga 2–9 with prominent posterolateral projections (Fig. 2.251); within coverage area known only from the Carolinas ............................................... Dannella lita 137’ Posterolateral projections absent or only weakly developed on abdominal terga 2 and 3 (Figs. 2.252, 2.253) (best evaluated based on later instars; some variation possible; additional undescribed species may be present but require significant further study) ..... 138

Chapter 2 ~ Key to Mayflies

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Fig. 2.243

Fig. 2.244

Fig. 2.242

Fig. 2.245

Fig. 2.246

Fig. 2.248

Fig. 2.249

Fig. 2.247

Fig. 2.250

Figures 2.242–2.250, Ephemerellidae genn. spp. 2.242, Ephemerella invaria (Allen & Edmunds, 1965), caudal filaments, dorsal. 2.243–2.244, right maxillae, ventral: 2.243, Ephemerella subvaria (modified from Allen & Edmunds, 1965); 2.244, Serratella serrata (Allen & Edmunds, 1963b). 2.245–2.246, abdomens, dorsal: 2.245, Attenella margarita; 2.246, Eurylophella funeralis. 2.247, Dannella provonshai, habitus, dorsal. 2.248–2.250, Attenella spp.: 2.248, A. attenuata (modified from Allen & Edmunds, 1961), pronotum, dorsal. 2.249–2.250, outlines of fronts of heads, dorsal: 2.249, A. attenuata; 2.250, A. margarita.

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138(137’) Posterolateral projections absent from terga 2 & 3 (Fig. 2.252); most terga of abdominal terga 2–7 usually with submedian pair of dark spots (e.g., Fig. 2.252) .... Dannella simplex 138’ Posterolateral projections weakly developed on abdominal tergum 3 (Fig. 2.253) and sometimes on tergum 2; abdominal terga 2–7 without submedian pairs of dark spots as above, although less distinct submedian pair of markings sometimes present on terga 4–7 (Fig. 2.253) ................................................................................. Dannella provonshai

Drunella species

(see also Allen & Edmunds, 1962 for historic treatment of these species and their junior synonyms, with emendments by Funk et al., 2008b) 139(129) Pair of submedian spines present on abdominal terga and well developed on at least terga 5–7; head roughened with pair of spines or tubercles between eyes above ocelli (Figs. 2.254–2.256) ......................................................................................................... 140 139’ Pair of submedian spines absent on abdominal terga (sometimes ridges without spines present); head relatively smooth between eyes and above ocelli, not roughened as above ............................................................................................................................... 142 [Note: The following couplets should be used cautiously because general species diversity and intraspecific variation in this geographic region are poorly understood for this genus.] 140(139) Moderately long, somewhat coarse setae densely rowed at distal margin of clypeus (e.g., Fig. 2.256) . ............................................................................................................ 141 [Note: Do not confuse frontal shelf with clypeus (at base of labrum), which is sometimes not apparent under frontal shelf in dorsal or facial view (e.g., it is not apparent in Figs. 2.254, 2.255).] 140’ Clypeus not as above (if setae present, then sparse) (incl. D. wayah) . ......... Drunella walkeri 141(140) Posterior margins of abdominal terga 8 and 9 with long hairlike setae protruding dorsally; head (e.g., Fig. 2.255) legs (Figs. 2.257, 2.258) and abdominal terga with dense to somewhat dense marginal setae; claws with few denticles situated basally (incl. D. conestee & D. cherokee) .................................................................... Drunella tuberculata 141’ Posterior margins of abdominal terga 8 and 9 without long hairlike setae protruding dorsally; head (Fig. 2.256) and legs (Fig. 2.259) with sparse setae; claws with denticles along most of length ..................................................................... Drunella allegheniensis 142(139’) Frontolateral projections of head (near inner base of antennae) poorly developed (not curved and not extending as far as margin of frontal shelf), with length subequal to basal width (Fig. 2.260); femora with few or no long, fine setae on posterior margin ........................................................................................................................ Drunella lata 142’ Frontolateral projections more strongly developed (straight or variously curved and extending almost to margin of frontal shelf to well beyond margin of frontal shelf), about twice as long as wide (Figs. 2.261–2.263); femora with dense row of long, fine setae on posterior margin ................................................................................................ 143

Chapter 2 ~ Key to Mayflies

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Fig. 2.251

Fig. 2.254

Fig. 2.255

Fig. 2.257

Fig. 2.260

Fig. 2.252

Fig. 2.258

Fig. 2.261

Fig. 2.262

Fig. 2.253

Fig. 2.256

Fig. 2.259

Fig. 2.263

Fig. 2.264

Figures 2.251–2.264, Dannella spp. and Drunella spp. (Ephemerellidae). 2.251–2.253. Dannella spp., abdomens, dorsal: 2.251, D. lita; 2.252, D. simplex; 2.253, D. provonshai. 2.254–2.264, Drunella spp.: 2.254–2.256, heads, frontal (modified from Allen & Edmunds, 1962): 2.254, D. walkeri; 2.255, D. tuberculata; 2.256, D. allegheniensis. 2.257–2.259. right forefemora, anterodorsal (Allen & Edmunds, 1962): 2.257, D. tuberculata variant 1; 2.258, D. tuberculata variant 2; 2.259, D. allegheniensis. 2.260–2.263, heads, frontal (Allen & Edmunds, 1962): 2.260, D. lata; 2.261, D. sp.; 2.262, D. sp.; 2.263. D. sp. 2.264, D. sp., apex of right foretibia with projection, tarsus, and claw, anterior.

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143(142’) Foretibial projections long and distinctly curved (Fig. 2.264) ................ Drunella longicornis [Note: This species has been considered a synonym of D. lata, but in light of recent analyses it may prove to be a variant of D. cornuta. However, it is a distinct morphotype, and so it is keyed here] 143’ Foretibial projections not developed as above . .................................................................... 144 [Note: Morphological variation among Southeast populations of the cornutacornutella complex requires further exploration to elucidate regional species boundaries. At least one putative new species of this complex remains undescribed. Use the following couplet with caution.] 144(143’) Hindtibiae usually greater than three-fourths length of hindfemora; mature body length usually greater than 8mm (7.8–11.4 mm), with midtibiae and hindtibiae length 1.6–2.2 mm; median ocellar tubercles more developed, with median tubercle distinctly longer than lateral tubercles ........................................................................................ Drunella cornuta 144’ Hindtibiae usually less than three-fourths length of hindfemora; mature body length usually less than 8mm (6.4–8.5 mm), with midtibiae and hindtibiae length 1.1–1.5 mm; median ocellar tubercles less developed, with median tubercle only slightly longer than lateral tubercles ............................................................................................... Drunella cornutella

Ephemerella species

(see Allen & Edmunds, 1965 for historic treatment of these species and their junior synonyms) 145(132) Abdominal terga with distinctive paired submedian spines (most well developed with some curvature) as in Figs. 2.265, 2.266 . ..................................................... Ephemerella needhami 145’ Abdominal terga with or without paired submedian spines, if present, not as developed as above .......................................................................................................................... 146 146(145’) Lateral margins of abdominal segments slightly serrate, with setae relatively few but stout (Fig. 2.267) ........................................................................................................................... 147 146’ Lateral margins of abdominal segments usually relatively smooth, with numerous weaker, but spatulate, setae .......................................................................................................... 148 147(146) Abdominal terga often heavily spiculate and with distinctive paired spines as in Fig. 2.268 ............................................................................................................ Ephemerella hispida 147’ Abdominal terga often glabrous or glossy and with paired spines not developed as above, but often with paired convexities of the posterior margins, giving rise to somewhat sinuate appearance (e.g., Fig. 2.269) ................................................ Ephemerella catawba 148(146’) Paired abdominal tergal spines moderately developed and sharp as in Fig. 2.270; tibiae and tarsi with highly contrasted dark bands (Fig. 2.271) ....................... Ephemerella subvaria 148’ Abdominal terga with or without variously developed paired spines; tibiae and tarsi with no dark bands or with only faint bands . ......................................................................... 149 [Note: Preliminary DNA analyses suggest that certain of the Ephemerella species below might represent complexes of species in the Southeast. In this key what appear to be morphological variants or extremes of ranges of variation are keyed.]

Chapter 2 ~ Key to Mayflies

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Fig. 2.266 Fig. 2.265

Fig. 2.268

Fig. 2.267

Fig. 2.269

Fig. 2.270

Fig. 2.271

Figures 2.265–2.271, Ephemerella spp. (Ephemerellidae). 2.265–2.266, Ephemerella needhami: 2.265, abdomen, dorsal (modified from Allen & Edmunds, 1965); 2.266, abdominal terga, left lateral. 2.267, E. catawba, right margin of abdominal tergum 5, dorsal. 2.268–2.271, Ephemerella spp., (modified from Allen & Edmunds, 1965): 2.268–2.270, abdomens, dorsal: 2.268, E. hispida; 2.269, E. catawba; 2.270, E. subvaria. 2.271, E. subvaria, right foretibia, tarsus, and claw, anterodorsal.

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149(148’) Paired spines present on some abdominal terga, with spines minute or small, sharp, and somewhat flattened and triangulate (e.g., Figs. 2.272–2.275) ........................................ 150 149’ Paired spines absent on abdominal terga or if armature present on some terga, then appearing as small blunt protrusions or spiculate ridges ................................................................. 151 150(149) Dorsal color pattern as in Fig. 2.276 . .................................................... Ephemerella floripara [Note: This species, sometimes considered a synonym of E. invaria, represents a distinct color variant atypical of other E. invaria variants or E. invaria complex species.] 150’ Color pattern variable, not as above (e.g., Figs. 2.272–2.275) ................. Ephemerella invaria [Note: This species may represent a complex of species in the Southeast.] 151(149’) Abdominal terga usually with no paired protrusions or prominent spiculate ridges; claws usually only gradually curved (Fig. 2.277) . .................................... Ephemerella dorothea [Note: This name very likely is being applied to a complex of species in the Southeast. As a word of caution, some western forms associated recently by rearing have paired protrusions.] 151’ Abdominal terga with or without paired protrusions or ridges (e.g., Figs. 2.279, 2.280); claws usually more strongly bent (Fig. 2.278) . .............................................................. 152 152(151’) Abdominal terga with posterior margins relatively smooth, with no indication of protrusions or ridges ....................................................................................... Ephemerella excrucians [Note: This species may be a complex of species in the Southeast, including, but not limited to, two commonly encountered nominal variants represented in the following couplet. Six additional Nearctic variants are considered synonyms.] 152’ Some abdominal terga with pairs of small protrusions or short longitudinal ridges at posterior margins (e.g., Figs. 2.279, 2.280) .................................................................... 153 153(152’) Abdominal terga usually with some paired submedian, spiculate ridges (e.g., Fig. 2.279); abdominal tergum 8 with obliquely oriented pale stripes ....................... Ephemerella rossi 153’ Abdominal terga without spiculate ridges (e.g., Fig. 2.280); abdominal tergum 8 without such stripes . ....................................................................................... Ephemerella crenula

Eurylophella species

(for more comprehensive treatments, see Funk & Sweeney, 1994 and Funk et al., 2008a) 154(135) Operculate gills with outer branch of underlying soft gill portion (found below dorsal gill plate) with upper lobes subequal in size to lower lobes (e.g., Fig. 2.281); submedian spines on abdominal tergum 7 relatively narrowly spaced compared to midlength of tergum (e.g., Fig. 2.284) ................................................................................................. 155 154’ Operculate gills with outer branch of underlying soft gill portion with upper lobes absent or highly reduced compared to lower lobes (e.g., Figs. 2.282, 2.283); submedian spines on abdominal tergum 7 relatively widely spaced compared to midlength of tergum (e.g., Fig. 2.285) . ............................................................................................................ 158

Chapter 2 ~ Key to Mayflies

Fig. 2.272

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Fig. 2.273

Fig. 2.274

Fig. 2.275

Fig. 2.276

Fig. 2.277

Fig. 2.278 Fig. 2.279

Fig. 2.281

Fig. 2.280

Fig. 2.282

Fig. 2.284

Fig. 2.283

Fig. 2.285

Figures 2.272–2.285, Ephemerella spp. and Eurylophella spp. (Ephemerellidae). 2.272–2.280. Ephemerella spp.: 2.272–2.275, abdomens, dorsal (modified from Allen & Edmunds, 1965): 2.272, E. sp.; 2.273, E. sp.; 2.274; E. sp.; 2.275, E. sp. (Allen & Edmunds, 1965). 2.276, E. floripara, habitus, dorsal. 2.277–2.278, right foreclaws, anterior: 2.277, E. dorothea; 2.278. E. sp. 2.279–2.280, abdomens, dorsal (Allen & Edmunds, 1965): 2.279, E. sp.; 2.280, E. sp. 2.281–2.285. Eurylophella spp.: 2.281–2.283, left operculate gills 2, left dorsolateral (dorsal plate above, outer branch of ventral, soft portion below; modified from Funk & Sweeney, 1994): 2.281, E. temporalis; 2.282, E. aestiva; 2.283, E. verisimilis. 2.284–2.285, abdominal terga 5–7, dorsal (modified from Allen & Edmunds, 1963a): 2.284, E. prudentalis; 2.285, E. lutulenta.

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155(154) Operculate gills with outer branch of underlying soft gill portion with six upper lobes; head with dorsal pair of tubercles between eyes poorly developed (e.g., Fig. 2.286) ............ 156 155’ Operculate gills with outer branch of underlying soft gill portion with nine or more dorsal lobes; head with dorsal pair of tubercles between eyes well developed (e.g., Fig. 2.287) . ............................................................................................................ 157 156(155) Submedian spines on abdominal terga 1–3 relatively straight in lateral view (Fig. 2.288); mature body length 6.5–8.0mm .................................................. Eurylophella prudentalis 156’ Submedian spines on abdominal terga 1–3 arched as seen in lateral view (similar to Fig. 2.289); mature body length 8.0–9.7mm; known only from North Carolina coastal plain streams, but may be found in South Carolina ......................... Eurylophella oviruptis [Note: These are larvae of a parthenogenetic Eurylophella with aberrant alates that demonstrate an unusual oviposition strategy of abdominal bursting.] 157(155’) Posterolateral projections on abdominal tergum 3 with length of inner margin about one-third midlength of tergum; forefemora width usually greater than one-third length ............... ................................................................................................................ Eurylophella doris 157’ Posterolateral projections on abdominal tergum 3 with inner length about one- fifth midlength of tergum; forefemora width usually about one-third or less length .............. ....................................................................................................... Eurylophella temporalis [Note: Many historical records of this species in the Southeast may be attributable to E. doris, especially east of the Appalachians.] 158(154’) Operculate gills with outer branch of underlying soft gill portion with two to five small dorsal lobes ..................................................................................................................... 159 158’ Operculate gills with outer branch of underlying soft gill portion with no dorsal lobes or only one or two minute dorsal lobes ............................................................................... 162 159(158) Body brown dorsally with only faint if any contrasting color pattern and no speckling; submedian pairs of spines on terga 1–4 relatively elongate with space between spines of each pair progressively increasing from tergum 1 to tergum 4 (Fig. 2.290) ............... ......................................................................................................... Eurylophella funeralis 159’ Body dorsally with some form of speckling or well-contrasted color patterning; submedian pairs of spines on terga 1–4 short and blunt, and more equally spread .......................... 160 160(159’) Head with dorsal pair of tubercles developed as in Fig. 2.291; mature body (prevalent during summer emergence) less than 8.0 mm .................................... Eurylophella aestiva 160’ Head with dorsal pair of tubercles generally not as well developed as above (e.g., Figs. 2.292, 2.293) or not detectable; mature body (prevalent during early spring emergence time) more than 8.5 mm ............................................................................... 161

Chapter 2 ~ Key to Mayflies

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Fig. 2.288 Fig. 2.286

Fig. 2.287 Fig. 2.289

Fig. 2.291

Fig. 2.292

Fig. 2.290 Fig. 2.293 Figures 2.286–2.293, Eurylophella spp. (Ephemerellidae). 2.286–2.287, heads, frontal (anterior): 2.286, E. prudentalis; 2.287, E. temporalis. 2.288–2.289, abdominal terga 1–3, left dorsolateral (modified from Funk & Sweeney, 1994): 2.288, E. prudentalis; 2.289, E. sp. 2.290, E. funeralis, abdominal terga 1–4, dorsal. 2.291–2.293, heads, frontal (anterior): 2.291, E. aestiva; 2.292, E. lutulenta; 2.293, E. enoensis.

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161(160’) Head with dorsal pair of tubercles small (e.g., Fig. 2.292) to absent; spines on abdominal terga 6 and 7 emerging from ridges at base .................................... Eurylophella lutulenta 161’ Head with dorsal pair of tubercles developed as in Fig. 2.293; spines on abdominal terga 6 and 7 with relatively flat base .......................................................... Eurylophella enoensis 162(158’) Head with dorsal pair of tubercles developed as in Fig. 2.294; mature body length always greater than 6.5 mm ...................................................................... Eurylophella verisimilis 162’ Head with dorsal pair of tubercles barely detectable in males and minute in females; mature body length variable ....................................................................................................... 163 163(162’) Space between submedian spines on abdominal tergum 5 distinctly greater than space between submedian spines on tergum 4 ............................................ Eurylophella bicolor 163’ Space between submedian spines on abdominal tergum 5 subequal to space between submedian spines on tergum 4 ........................................................................................ 164 [Note: Morphology in the following couplet may be reliable only when keying mature larvae.] 164(163’) Posterolateral projections somewhat developed on abdominal tergum 2 (Fig. 2.295); mature body (prevalent in spring emergence time) usually greater than 7.5 mm . ...................... ............................................................................................... Eurylophella macdunnoughi 164’ Posterolateral projections not developed on tergum 2, represented by small angle only (Fig. 2.296); mature body (prevalent during summer emergence time) always less than 7.5 mm .................................................................................. Eurylophella minimella

Serratella species

(for historical treatment of these species and their junior synonyms, see Allen & Edmunds, 1963b) 165(133’) Head, thorax, and legs generally with long hairlike setae (femora with at least some, but often dense hairlike setae marginally) (Figs. 2.297–2.300); if hairlike setae not developed at posterior margin of femora, then four black dots present on abdominal sterna, otherwise such dots present or absent ................................................................. 166 165’ Head, thorax, and legs without long hairlike setae (femora without marginal hairlike setae) (Fig. 2.301); sterna without four black dots ............................................ Serratella frisoni 166(165) Abdominal sterna each with four black dots; abdominal tergum 9 posterolateral projections not extending beyond posterior margin of tergum 10, usually only reaching about midlength of tergum 10 . ................................................................... Serratella serratoides 166’ Abdominal sterna each without four black dots; abdominal tergum 9 posterolateral projections extend to about posterior margin of tergum 10 or beyond . .. Serratella serrata [Note: This name likely is being applied to a group of cryptic species that are sometimes distinguishable at individual locales by coloration, setation of the head, and relative development of abdominal armature, but they are not yet consistently distinguishable across their entire ranges. Nominal variants include S. carolina, S. sordida, and S. spiculosa]

Chapter 2 ~ Key to Mayflies

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Fig. 2.294 Fig. 2.295

Fig. 2.297

Fig. 2.296

Fig. 2.298

Fig. 2.300

Fig. 2.299

Fig. 2.301

Figures 2.294–2.301, Eurylophella spp. and Serratella spp. (Ephemerellidae). 2.294, E. verisimilis, head, frontal (anterior). 2.295–2.296, Eurylophella spp., right halves of abdominal terga 2: 2.295, E. macdunnoughi; 2.296, E. minimella. 2.297–2.301, Serratella spp., right forefemur, anterodorsal (upside down; modified from Allen & Edmunds, 1963b): 2.297, S. serratoides; 2.298, S. serrata variant 1; 2.299, S. serrata variant 2; 2.300, S. serrata variant 3; 2.301, S. frisoni.

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EPHEMERIDAE genera 167(16’) Frontal process of head obtuse (either slightly rounded, roundly pointed, conical or nearly truncate but never bifurcate or deeply concave) (e.g., Figs. 2.302, 2.303); forelegs with tibia produced into large acute process anterodistally as in Fig. 2.305; labium with paraglossae large and strongly produced basally as well as distally (Fig. 2.307) .......... 168 167’ Frontal process of head bifurcate (deeply concave on distal margin giving rise to pair of short lateral prongs) (similar to Fig. 2.304); forelegs with tibia not produced into large acute process anterodistally (Fig. 2.306); labium with paraglossae not strongly produced basally (similar to Fig. 2.308) ..................................................... Ephemera, 169 168(167) Small filamentous gills 1 each bifid from near its base; antennae with whorls of long hairlike setae (Fig. 2.302); frontal process rounded, conical or truncate, usually not angulate ....................................................................................................... Hexagenia, 172 168’ Filamentous gills 1 each consisting of single unbranched filament; antennae without whorls of long hairlike setae (Fig. 2.303); frontal process angulate as in Fig. 2.303 . ................ .....................................................................................................Litobrancha, L. recurvata

Ephemera species

(based in part on McCafferty, 1975) 169(167’) Abdomen usually with some color patterning, at least with longitudinal markings on some sterna, not appearing somewhat flattened and broadened; caudal filaments half or less length of body; wingpads with or without various dark spotting ................................... 170 169’ Abdomen essentially lacking color patterning or dark markings, appearing somewhat flattened and broadened; caudal filaments more than half length of body; wingpads with dark spotting ................................................................................ Ephemera guttulata 170(169) Hindwingpads and forewingpads with dark spotting, developed at least as that shown in Figs. 2.309, 2.310 ................................................................................ Ephemera simulans [Note: Wingpad spotting is usually apparent to at least some degree beginning in middle-instar larvae and more apparent in late instars. Spotting should not be confused with dark margining of veins that is sometimes apparent in relatively mature larvae of several species.] 170’ Hindwingpads without dark spotting .................................................................................... 171 171(170’) Forewingpads with dark spotting . ................................................................... Ephemera varia 171’ Forewingpads without dark spotting, but sometimes some diffuse blotching apparent ................................................................................................................. Ephemera blanda

Chapter 2 ~ Key to Mayflies

Fig. 2.302

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Fig. 2.303

Fig. 2.304

Fig. 2.305

Fig. 2.306 Fig. 2.307

Fig. 2.309

Fig. 2.308

Fig. 2.310

Figures 2.302–2.310, Ephemeridae genn. spp. 2.302–2.304, heads, dorsal: 2.302, Hexagenia limbata; 2.303, Litobrancha recurvata; 2.304, Ephemera sp. 2.305–2.306, right forelegs, anterior: 2.305, Hexagenia limbata; 2.306, Ephemera sp. 2.307–2.308, labia, dorsal (left) and ventral (right): 2.307, Hexagenia limbata; 2.308, Ephemera sp. 2.309–2.310, Ephemera simulans: 2.309, left forewingpad, dorsal. 2.310, left hindwingpad, dorsal.

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Hexagenia species 172(168) Frontal process somewhat conical (Fig. 2.311); mature males with distinctive, nearly straight developing penes (approximating Fig. 2.313) ............................ Hexagenia rigida 172’ Frontal process not conical, some with slightly convergent lateral margins at most; mature males with developing penes with some inward curvature (e.g., Figs. 2.314, 2.315) .... 173 173(172’) Frontal process truncate (Fig. 2.312), sometimes very slightly concave along otherwise straight distal margin; abdominal terga 2–6 often with one or two pairs of outlined pale spots submedially (some spots sometimes slightly oblong) .......... Hexagenia atrocaudata 173’ Frontal process rounded or truncate; abdominal terga not as above, often without pairs of pale markings on abdominal terga 2–6 (at most with single pairs of oblique outlined bars submedially) ............................................................................................................ 174 174(173’) Mature males with developing penes with beaklike apex sharply angled inwardly and angulate along outer margin (approximating Fig. 2.314) ................... Hexagenia bilineata 174’ Mature males with developing penes with broadly pointed apex rounding inwardly at apex, with outer margin curved but not angulate as above (approximating Fig. 2.315) ..................................................................................................... Hexagenia limbata group [Note: Variabilities in color pattern and claw thickness are inconsequential in consistently separating H. bilineata, H. limbata, and H. orlando. Hexagenia bilineata is common in, but not restricted to, larger streams and rivers throughout the coverage area. Hexagenia limbata is common in, but not restricted to, smaller streams and lakes/ponds throughout the coverage area. Hexagenia orlando, which is probably synonymous with H. limbata, is said to be restricted to sand-bottomed lakes of central Florida. Hexagenia munda, a junior synonym of H. limbata, may be a valid species. Association of adults with larvae is critical for precise identification of most larvae of these species. Developing male genitalia of last-instar larvae are similar to those found in subimagos.]

Chapter 2 ~ Key to Mayflies

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Fig. 2.311

Fig. 2.313

Fig.2.312

Fig. 2.314

Fig. 2.315

Figures 2.311–2.315, Hexagenia spp. (Ephemeridae). 2.311–2.312. frontal process of head, dorsal: 2.311, H. rigida; 2.312, H. atrocaudata. 2.313–2.315, larval final instar left penis, ventral: 2.313, H. rigida; 2.314, H. bilineata; 2.315, H. limbata.

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HEPTAGENIIDAE genera

(see Webb & McCafferty, 2008 for world treatment) 175(6’) 175’

Median caudal filament (middle tail) highly reduced (two tailed) ....................................... 176 Median caudal filament well developed (three tailed) . ........................................................ 177

176(175) Setae not developed along mid-length region of cerci (paired lateral tails); head with short dense fringe of setae at anterior margin, and with no dorsal tubercles as described below (Fig. 2.316) . ......................................................................................... Epeorus, 186 176’ Setae well developed along mid-length region of cerci; head without anterior fringe of setae, but with paired tubercles dorsally (Fig. 2.317) ......................... Spinadis, S. simplex [Note: This species is rarely taken and has been reported only from Georgia and the Mississippi River between Mississippi and Louisiana in the coverage area and otherwise only from Arkansas within the Southeast. The presence of tail setae may suggest that Spinadis larvae are at least in part swimmers.] 177(175’) First pair of gills meet or overlap beneath abdomen ..................................... Rhithrogena, 227 [Note: Of the nine species currently known from the coverage area, R. anomala and R. rubicunda remain unknown as larvae.] 177’ First pair of gills do not meet ventrally . ............................................................................... 178 178(177’) Ventral face of maxillae with longitudinal row of setae near median margin and without any groupings of scattered setae (Figs. 2.318, 2.319); gills 1 variously shaped, but not concave lengthwise ......................................................................................................... 179 [Note: Face refers to the main body of the maxilla, which is also known as the galealacinia. The maxillary palp is attached near the base of the lateral (outer) margin of the galealacinia. Figs. 2.318–2.320 thus show the ventral face of the right maxilla. Gill numbers (any of 1 through 7) refer to the abdominal segments to which they are attached.] 178’ Ventral face of maxillae with variously scattered setae, sometimes in addition to rowed setae (similar to Fig. 2.320); gills 1 somewhat concave lengthwise (similar to Fig. 2.321) ..... 185

Chapter 2 ~ Key to Mayflies

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Fig. 2.316

Fig. 2.318

Fig. 317

Fig. 2.319

Fig. 2.320

Fig. 2.321 Figures 2.316–2.321, Heptageniidae genn. spp. 2.316–2.317, heads, dorsal: 2.316, Epeorus sp. (modified from Jensen, 1972); 2.317, Spinadis simplex. 2.318–2.320, right maxillae, ventral: 2.318, Maccaffertium sp.; 2.319, Raptoheptagenia cruentata (Jensen, 1972); 2.320, Ecdyonurinae sp. (Jensen, 1972). 2.321, Ecdyonurinae sp., left gill 1, left dorsolateral.

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179(178) Gill lamellae modified into narrow elongate structures inserted ventrally (generally hidden from above); ventral gill fibrillae (filaments or tufts) from beneath abdomen partially seen in dorsal view as in Fig. 2.322; apical segment of labial palps lanceolate (narrow elongate and sharply pointed as in Fig. 2.323) .................. Raptoheptagenia, R. cruentata [Note: This species has rarely been taken in the Southeast (only from Arkansas and Tennessee). As larvae, it historically was often confused by name with Anepeorus simplex (=Spinadis simplex) because of presumed, but incorrect, associations of larvae with adults.] 179’ Gill lamellae not highly modified as above and inserted laterally or dorsolaterally; apical segment of labial palps not lanceolate as above ............................................................. 180 [Note: Gills of Heptageniidae consist of at least a platelike or leaflike portion (lamella) and often also a basal portion below or behind the lamella, often somewhat hidden and made up of shorter filaments (usually a group or tuft of filaments) referred to as fibrillae.] 180(179’) Gills 7 (originating on abdominal segment 7) slender or vestigial, without fibrillae; gills 1–6 not thickened at outer margins, but with single thickened submarginal rib sometimes detectable (e.g., Fig. 2.324) ............................................................................................ 181 [Note: Gills refer generally to the platelike or leaflike portion (lamella) or to the lamella and fibrillae portions together, which should be clear from the statements within the respective couplets. Gill characters of shape, ribbing, folding or tracheation refer only to lamellae.] 180’ Gills 7 similar in shape to other gills, and with fibrillae; gills 1–6 with thickened anterior and posterior margins (at least in basal half of gill) ....................................................... 184 181(180) Gills 6 subequal in size to gills 1–5; gills 7 developed but smaller than gills 6 ................... 182 181’ Gills 6 much smaller than gills 1–5; gills 7 vestigial (considerably smaller than posterolateral process of segment 7) ............................................................................... Macdunnoa, 222 182(181) Gills 1–6 acutely pointed apically (Fig. 2.324) ................................................. Stenacron, 233 182’ Gills 1–6 not acutely pointed (Figs. 2.325, 2.326) ............................................................... 183 183(182’) Gills 1–6 with distal margin straight for most of margin (Fig. 2.325); gills 7 without median tracheal trunk (Fig. 2.327) ................................................................... Maccaffertium, 199 183’ Gills 1–6 with distal margin rounded (Fig. 2.326); gills 7 with median tracheal trunk (Fig. 2.328) .................................................................................................... Stenonema, S. femoratum

Chapter 2 ~ Key to Mayflies

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Fig. 2.324

Fig. 2.322

Fig. 2.325

Fig. 2.326

Fig. 2.323 Fig. 2.327

Fig. 2.328

Figures 2.322–­2.328, Heptageniidae genn. spp. 2.322–2.323, Raptoheptagenia cruentata: 2.322, abdomen and meso- and metathorax, dorsal (modified from Jensen, 1972); 2.323, right labial palp, ventral. 2.324–2.326, left gills 4, left dorsolateral: 2.324, Stenacron sp.; 2.325, Maccaffertium sp.; 2.326, Stenonema femoratum. 2.327–2.328, left gills 7: 2.327, Maccaffertium sp.; 2.328, Stenonema femoratum.

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184(180’) Head emarginate (with medial concavity at anterior margin) and usually with some laterally protruding part of palps as seen in dorsal view (Fig. 2.329); forefemora with dorsal apical lobe narrower than ventral apical lobe as in Fig. 2.330 ..... Cinygmula, C. subaequalis 184’ Head not emarginate and without laterally exposed mouthparts seen in dorsal view; forefemora with dorsal apical lobe not as above ...................................... Heptagenia, 192 [Note: Although H. elegantula reaches West Virginia, we do not necessarily expect it in the coverage area.] 185(178’) Caudal filaments with interfacing setae developed along inner margins of cerci and both lateral margins of median caudal filament; head width less than or subequal to that of pronotum................................................................................................................ Nixe, 223 185’ Caudal filaments without interfacing setae; head width greater than that of pronotum .................................................................................................................... Leucrocuta, 195 [Note: Larvae in this genus are best resolved to species via rearings to male adults, keeping in mind that species taxonomy, which has been based on adults, is itself in some doubt. Identifying larvae of Leucrocuta remains highly tentative and to a large extent cannot be done with any confidence. Previously published keys are not reliable. Only a partial, tentative key to species of Leucrocuta can be given here.]

Epeorus species

(see also Webb & McCafferty, 2006; Burian et al., 2008) 186(176) Gills 1 not expanded anteriorly and not meeting or nearly meeting ventrally (Fig. 2.331); maxillae galealacinia with one short, distinct seta at base of apical denticles (Fig. 2.334); dorsal apical lobe of femora blunt or sharp .................................................................... 187 [Note: The galealacinia is the main body of the maxilla, and the apex is the inner distal tip of the galealacinia; gills 1 refer to the first pair of gills (originating on abdominal segment 1).] 186’ Gills 1 expanded anteriorly, usually meeting or nearly meeting ventrally (Fig. 2.332), or with slighter expansion about half or less as much as previous (Fig. 2.333) and not nearly meeting ventrally; maxillae with galealacinia with more than one distinct seta at base of apical denticles (Fig. 2.335); dorsal apical lobe of femora bluntly pointed . ..... 190 187(186) Abdomen with outer (dorsal) posterolateral projections distinctly longer than inner (ventral) projections (Fig. 2.336) . ............................................................................. Epeorus vitreus 187’ Abdomen with outer (dorsal) posterolateral projections subequal to, or only somewhat longer than, inner (ventral) posterolateral projections (Fig. 2.337) . .............................. 188

Chapter 2 ~ Key to Mayflies

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Fig. 2.329

Fig. 2.332

Fig. 2.330

Fig. 2.333

Fig. 2.336

Fig. 2.331

Fig. 2.334

Fig. 2.335

Fig. 2.337

Figures 2.329–2.337, Cinygmula subaequalis and Epeorus spp. (Heptageniidae). 2.329–2.230, Cinygmula subaequalis: 2.329, head, dorsal (modified from Jensen, 1972); 2.330, left forefemur apex (dorsal lobe up), anterodorsal. 2.331–2.337, Epeorus spp.: 2.331. E. punctatus, left gill, ventral. 2.332–2.333, left gills, ventral: 2.332, E. pleuralis; 2.333, E. namatus. 2.334–2.335, left maxillary galealaciniae: 2.334, E. punctatus; 2.335, E. namatus. 2.336–2.337, right posterolateral projection of abdominal segment 7, dorsal: 2.336, E. vitreus; 2.337, E. punctatus.

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188(187’) Abdominal terga each with small but distinct medial protuberance at posterior margins, protuberances progressively increasing in size progressively on posterior terga; head much wider anteriorly (Fig. 2.338) and nearly uniform brown ................... Epeorus dispar 188’ Abdominal terga either without posteromedian protuberances or terga 2–7 or 3–7 with small protuberances decreasing in size posteriorly; head subrectangular (Fig. 2.339) and with pale spots . ........................................................................................................ 189 189(188’) Dorsal apical lobe of hindfemora sharply pointed (Fig. 2.340); pronotum often with distinct tubercles; mature body over 8.0 mm .................................................. Epeorus subpallidus 189’ Dorsal apical lobe of hindfemora bluntly pointed (Fig. 2.341); pronotum without distinct tubercles; mature body 6.0–8.0 mm; not known from coverage area . .. Epeorus punctatus [Note: This species is expected to occur in the coverage area because it has been reported from Virginia and West Virginia. Body length does not include caudal filaments. A potentially new species we have not studied may key to this couplet.] 190(186’) Gills 1 meeting or nearly meeting ventrally.......................................................................... 191 190’ Gills 1 not as above; within coverage area known only from Kentucky ....... Epeorus namatus 191(190) Abdomen with at least terga 3–7 having paired submedian dark spots (sometimes faint); mature body length usually over 9.84 mm ............................................. Epeorus pleuralis [Note: Body length never includes caudal filaments.] 191’ Abdomen with terga 3–7 without dark spots as above; mature body length usually less than 9.55 mm ............................................................................................. Epeorus fragilis

Heptagenia species

(see Webb et al., 2007 and Jacobus & Webb, 2013 for more detail) 192(184’) Abdominal terga each with pair of sublateral dark longitudinal bars extending from anterior to posterior tergal margins, bars best developed on terga 3–8, with shape ranging from vertical to oblique to slightly arching medially (e.g., Fig. 2.342) ................................... .............................................................................................. Heptagenia marginalis group [Note: Heptagenia townesi, known from the coverage area but not formally described as a larva, apparently is closely related to H. marginalis. The tentatively associated larva of H. townesi differs from that of H. marginalis by having longer apical spines on the segments of the caudal filaments. These spines are 1/3 the length of the segment for H. townesi compared to 1/8 the length of the segment for H. marginalis. We have associated one larva with H. dolosa, and it, too, is similar to H. marginalis; unfortunately, no reliable morphological characters for species diagnosis have been discovered.] 192’ Abdominal terga without pairs of longitudinal bars as above, but often as in Fig. 2.343 .... 193

Chapter 2 ~ Key to Mayflies

Fig. 2.338

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Fig. 2.339

Fig. 2.340

Fig. 2.341

Figures 2.338–2.341, Epeorus spp. (Heptageniidae). 2.338–2.339, head outline, dorsal: 2.338, E. dispar; 2.339, E. punctatus. 2.340–2.341, left hind femur apex (dorsal lobe up), anterodorsal: 2.340, E. subpallidus; 2.341, E. punctatus.

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193(192’) Mature body length 12 mm or more ..................................................................................... 194 [Note: Body length never includes caudal filaments.] 193’ Mature body length 8–9 mm............................................................................ Heptagenia julia 194(193) Abdominal sterna pale, with dark, transverse band at posterior margin of sternum 9 .......................................................................................................... Heptagenia flavescens 194’ Abdominal sterna pale (sometimes with longitudinal dark stripe medially for length of abdomen), either with no markings at posterior margin of sternum 9 or typically with posterolateral spots on sternum 9 . ................................................................. Heptagenia pulla

Leucrocuta species 195(185’) Gills darkened with black, dark brown, purple brown or purple black; gills without distinct tracheation except occasionally mid-tracheal trunk evident on some gills .................... 196 195’ Gills not darkened, clear to translucent, sometimes appearing grayish, gill with or without distinct tracheation .......................................................................................................... 197 196(195) Gills darkened in upper half above midline of gill, with darkening sometimes extending over apex on both sides; abdominal tergum 9 brown; marginal pale areas on head present or absent ................................................................................................... Leucrocuta thetis [Note: This is the only species that can be associated with these characteristics; however, it is possible that more than one species will key here. The larva of L. thetis originally was described as the larva of L. juno by Traver; thus, previous key concepts of L. thetis and L. juno have been compromised. The larva of L. umbratica may have entirely darkened gills, but it is not yet known from the coverage area.] 196’ Gills darkened along midline on both sides, then becoming infuscated, more so in the upper half of gill; abdominal terga, including 9, with extensive pale areas medially; marginal pale areas not present on head .......................................................................... Leucrocuta sp. 1 197(195’) Gills with indistinct trachea; pale marginal areas absent on head ........... Leucrocuta variety A [Note: This variety cannot be assigned to a species, and it is not known if it is species specific. It was inaccurately keyed as L. thetis by Traver, which apparently has been described as a larva only under the name L. juno.] 197’ Gills with distinct tracheation; pale marginal areas present on head (sometimes not apparent on light individuals) ........................................................................................................ 198

Chapter 2 ~ Key to Mayflies

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Fig. 2.342 Figures 2.342–2.343, Heptagenia spp. 2.342, H. marginalis; 2.343, H. flavescens.

Fig. 2.343 (Heptageniidae).

Abdominal

terga

4–5,

dorsal:

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

198(197’) Head with pair of oblique dark bars between compound eyes; abdominal terga 2–9 (Fig. 2.344) with posterior margins with pair of purple-black transverse lines extending sublaterally to submedially on either side and with considerable gap between them, and with pairs of sublateral dark purple-brown to black longitudinal bars expanding medially at base and attaching to posterior stripes (Fig. 2.344) in well-marked individuals (bars sometimes appearing as simple stripes or diagonal bars, and in some light individuals sometimes evident only on terga 2 and 3) .......................................... Leucrocuta minerva 198’ Not characterized as above ............. Leucrocuta aphrodite, L. hebe, L. maculipennis, L. juno [Note: L. aphrodite and L. hebe may have a pale medial V-shaped area on abdominal terga 7, 8, and most of the medial length of 9, or on tergum 9, three connected or unconnected pale spots (possibly even one medial spot in extreme variants). Sternal and caudal filament markings that have been associated with either of these species vary considerably between and among populations, and other characteristics that may be of diagnostic value are not known with confidence. For example, L. hebe might have an unmarked ventral abdomen or one with pairs of ventral spots in some populations, and also commonly a pair of posterolateral spots on sternum 9, or extensive lateral staining on 9. Abdominal terga of L. maculipennis may be unlike those of L. aphrodite and L. hebe; however, variations in terga 7–9 in L. hebe and L. maculipennis appear to begin to overlap. Especially the medial pale condition of tergum 9 appears to vary among several species. Diagnostics for the L. juno larva are unknown. It is unclear if larvae have been accurately associated with identifiable male adults, and thus the larva only is assumed to be similar to this grouping, and according to previous keys, to L. maculipennis in particular.]

Maccaffertium species

(based on Bednarik & McCafferty, 1979; McCafferty, 1981b, 1990, 2011a; McCafferty & Lenat, 2010; and new research) 199(183) Posterolateral projections well developed on abdominal segments 3–9, 4–9, or 5–9 (e.g., Fig. 2.345) . ............................................................................................................ 200 [Note: Some projections on anterior segments may be quite small and difficult to detect in early or middle instars. Abdominal posterloateral projections are best viewed ventrally with careful lighting, so that gill sockets do not cause confusion. The last visible abdominal sternite (in ventral view) is represented by sternum 9, and the last visible abdominal tergite (in dorsal view) is represented by tergum 10.] 199’ Posterolateral projections well developed on abdominal segments 7–9 (most frequently) or 6–9 ..................................................................................................................................................... 209

Chapter 2 ~ Key to Mayflies

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Fig. 2.344

Fig. 2.345

Figures 2.344–2.345, Leucrocuta minerva and Maccaffertium mediopunctatum (Heptageniidae). 2.344, Leucrocuta minerva, habitus, dorsal. 2.345, Maccaffertium mediopunctatum variant, abdomen, ventral.

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200(199) Maxillary crowns with fewer than 10 (usually 0) hairlike setae (similar to Fig. 2.346) ...... 201 [Note: The maxillary crown is the distal margin of the main body of the maxilla (galealacinia) from the outer curvature to the tip.] 200’ Maxillary crowns with more than 10 (usually many more) hairlike setae (similar to Fig. 2.347) . ..................................................................................................................... 203 201(200) Abdominal sternal pattern with dark bands extending obliquely or transversely (Figs. 2.345, 2.348, 2.349) .............................................. Maccaffertium mediopunctatum 201’ Abdominal sterna without banding pattern as above . .......................................................... 202 202(201’) Maxillary crowns with 6–9 robust setae (non-hairlike setae); galealaciniae with 25–40 setae in submedian row (row lengthwise near but not on the inner edge of body of maxilla); posterolateral processes of abdominal segment 9 subequal to those of segment 8; mature body length 10–12 mm .................................................. Maccaffertium carlsoni [Note: Body length never includes caudal filaments.] 202’ Maxillary crowns with 5 robust setae; galealaciniae with 18–20 setae in submedian row; posterolateral processes of abdominal segment 9 much shorter than those of segment 8; mature body length 8–10 mm; within coverage area known only from North Carolina . .................................................................................... Maccaffertium wudigeum 203(200’) Abdominal sternal pattern with dark bands (sometimes interrupted medially) extending transversely on posterior margins of at least sterna 5–8 (Figs. 2.350, 2.351) ................. ........................................................................................ Maccaffertium vicarium (in part) 203’ Abdominal sterna without dark posterior cross bands on sterna 5–8, other possible markings variable (e.g., Figs. 2.352–2.354) or absent . .................................................................. 204 204(203’) Abdominal sterna with markings near anterior margins on at least sterna 5–8 as in Figs. 2.352, 2.353; foreclaws without minute denticles .................................................. ......................................................................................... Maccaffertium pudicum (in part) [Note: Compound high magnification usually is required to view minute denticles in distal curvature of claw; claw denticles should not be confused with the larger basal prominence on the inner margin of all claws in Maccaffertium.] 204’ Abdominal sterna pale or without markings exactly as above; foreclaws with or without minute denticles .............................................................................................................. 205 205(204’) Abdominal terga 2–7 with distinct cross bands posteriorly, sharply contrasting with pale anterior areas of terga (Fig. 2.355); foreclaws without minute denticles ........................ ........................................................................................ Maccaffertium vicarium (in part) 205’ Abdominal terga not exactly as above, either posterior bands much narrower or absent; foreclaws with or without minute denticles .......................................................................... 206 206(205’) Maxillary crowns with 5–8 robust setae; foreclaws without minute denticles ................ ......................................................................................... Maccaffertium pudicum (in part) [Note: These robust setae are non-hairlike armature, and can be either comblike (pectinate) or spinelike.] 206’ Maxillary crowns with 2–4 robust setae, rarely only one spinelike seta, not including most apical spine at inner edge; foreclaws with or without denticles ..................................... 207

Chapter 2 ~ Key to Mayflies

Fig. 2.346

Fig. 2.351

Page 113

Fig. 2.347

Fig. 2.352

Fig. 2.348

Fig. 2.353

Fig. 2.349

Fig. 2.354

Fig. 2.350

Fig. 2.355

Figures 2.346–2.355, Maccaffertium spp. (Heptageniidae). 2.346–2.347, right maxillary galealaciniae: 2.346, M. puchellum; 2.347, M. modestum. 2.348–2.354, abdomens. ventral: 2.348, M. mediopunctatum variant; 2.349, M. mediopunctatum variant; 2.350, M. vicarium variant; 2.351, M. vicarium variant; 2.352, M pudicum variant; 2.353, M. pudicum variant; 2.354, M. appaloosa. 2.355, M. vicarium, abdomen, dorsal.

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207(206’) Foreclaws with minute denticles; at least several abdominal sterna usually with pair of sublateral spots and/or median spot of some kind near anterior margin ........................ 208 207’ Foreclaws without minute denticles; abdominal sterna generally unmarked or often only abdominal sternum 9 with posterolateral markings ....................... Maccaffertium sinclairi 208(207) Abdominal tergal pattern distinctive, as in Fig. 2.365 ............................. Maccaffertium lenati [Note: The holotype of M. lenati will key here. McCafferty (1990) characterized the abdominal posterolateral projections and claw denticulation differently than what would lead to this point in the key. See couplet 216.] 208’ Abdominal sternal pattern distinctive (Fig. 2.354), most sterna with pair of sublateral spots and median spot near anterior margin (those on 6–9 large and prominent), abdominal sternum 9 also with large pair of posterior marks (Fig. 2.354); known only from Tennessee .......................................................................... Maccaffertium appaloosa [Note: The predominately northeastern and midwestern M. luteum, which is known tentatively from Arkansas and Louisiana based on adults, has minute denticles on the foreclaws and some shared characteristics of mouthpart armature and abdominal posterolateral projections, but differs from M. appaloosa by lacking spotted abdominal sterna or dense outer setae of the cerci; M. luteum instead has abdominal sternal patterns sometimes similar to M. vicarium and cerci with sparse or no outer setae.] 209(199’) Maxillary crowns with fewer than 10 (usually 0) hairlike setae . ......................................... 210 [Note: The maxillary crown is the apical margin of the body of the maxilla (galealacinia) from the outer curvature to the tip.] 209’ Maxillary crowns with more (usually many more) than 10 hairlike setae ........................... 216 210 (209) Head capsule with three highly contrasting pale yellow markings within dark brown anterior area with middle marking crown shaped or diamond shaped (Fig. 2.356); hindfemora relatively short and broad (Fig. 2.356); abdomen generally without strong dark markings, mostly sublateral pairs of brown dots dorsally (Fig. 2.356); mature body length 6–7.5 mm; within coverage area known only from Kentucky ..................................................................................................... Maccaffertium bednariki [Note: This species, known from Missouri and Oklahoma, is expected to be found in Arkansas, and it might occur in Tennessee. A similar larva very recently collected from central Tennessee, with posterior stripes on abdominal terga and other prominent markings, may prove to be this species or a closely related new species; it requires better specimens for further study.] 210’ Head capsule not as above; hindfemora usually not short and broad; abdominal markings variable; mature body length 7–16 mm .......................................................................... 211 [Note: Body length never includes caudal filaments.] 211(210’) Abdominal terga with pale V-shaped areas on terga 5 and 7–9 or 8–9 (Fig. 2.357); mature body length 10–16 mm ...................................................... Maccaffertium meririvulanum 211’ Abdominal terga without pale V-shaped areas; mature body length 7–11 mm .................... 212 212(211’) Mesonotum (dorsum of second thoracic segment) with large, pale spots or transverse pale band at base of wingpads (Fig. 2.358) .......................................... Maccaffertium exiguum 212’ Mesonotum without pale markings as above . ...................................................................... 213

Chapter 2 ~ Key to Mayflies

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Fig. 2.357

Fig. 2.356

Fig. 2.358

Figures 2.356–2.358, Maccaffertium spp. (Heptageniidae). 2.356, M. bednariki, habitus, dorsal. 2.357, M. meririvulanum, abdomen, dorsal. 2.358, M. exiguum, thorax and abdomen, dorsal.

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213(212’) Maxillary crowns with one to eight hairlike setae ........... Maccaffertium terminatum (in part) 213’ Maxillary crowns setae with no hairlike setae . .................................................................... 214 214(213’) Sternum 9 with brown posterior margin (Fig. 2.359) ...... Maccaffertium terminatum (in part) 214’ Sternum 9 with pale posterior margin, at least medially ...................................................... 215 215(214’) Abdominal sterna 4–7 often with pair of brown longitudinal bars subaterally and pair of oblique lines submedially (Fig. 2.360), but sometimes without such markings; abdominal tergum 7 mostly dark (Fig. 2.361) ........... Maccaffertium terminatum (in part) 215’ Abdominal sterna as in Fig. 2.362; abdominal tergum 7 mostly white (Fig. 2.363) .................................................................................................. Maccaffertium pulchellum 216(209’) Abdominal terga 7–9 together with medial V-shaped pale area as in Fig. 2.364; maxillary crowns with 1–3 (very rarely 4) robust (non-hairlike) setae (sometimes only one distinctly comblike) ................................................................. Maccaffertium mexicanum [Note: Some variants of M. lenati, especially early or middle instars, may key here (see also couplet 208). They can be distinguished from M. mexicanum by having abdominal terga 7–9 each pale with darkened lateral areas, posterior margins and variable median longitudinal stripes, and a pair of small submedian dots (Fig. 2.365). Mature body lengths of these species differ, with M. mexicanum being 7–8 mm and M. lenati being 11–14 mm. Body length never includes caudal filaments.] 216’ Abdominal terga not fitting either condition described above; maxillary crowns with 3–7 robust setae (rarely only three, and if three, then abdominal terga 8 and 9 mostly dark) ........................................................................................................................................ 217 217(216’) Abdominal sterna 3–8, 4–8, 5–8, or 6–8 with distinct pattern of oblique (somewhat crescent shaped) crossbars (Figs. 2.366, 2.367) ........................................................................... 218 217’ Abdominal sterna without above patterns; if crescent-shaped crossbars present, then on terga 7 and 8 only . .......................................................................................................... 220 218(217) Foreclaws without denticles; maxillary crowns with 15–35 hairlike setae ........................... ............................................................................................. Maccaffertium ithaca (in part) [Note: Compound high magnification usually is required to view minute claw denticles in distal curvature of claw; denticles should not be confused with the larger basal prominence on the inner margins of all claws in Maccaffertium.] 218’ Foreclaws with denticles; maxillary crowns with 15–50 hairlike setae ............................... 219 219(218’) Maxillary crowns with more than 30 hairlike setae . ......... Maccaffertium modestum (in part) 219’ Maxillary crowns with 30 or fewer hairlike setae .................... Maccaffertium ithaca (in part)

Chapter 2 ~ Key to Mayflies

Fig. 2.359

Page 117

Fig. 2.360

Fig. 2.361

Fig. 2.364

Fig. 2.362

Fig. 2.366

Fig. 2.363

Fig. 2.367

Fig. 2.365 Figures 2.359–2.367, Maccaffertium spp. (Heptageniidae). 2.359–2.361, M. terminatum: 2.359, variant, abdomen, ventral; 2.360, variant, abdomen, ventral; 2.361, abdomen, dorsal. 2.362–2.363, M. pulchellum: 2.362, abdomen, ventral; 2.363, abdomen, dorsal. 2.364, M. mexicanum, abdomen, dorsal. 2.365, M. lenati, habitus, dorsal. 2.366–2.367, abdomens, ventral: 2.366, M. ithaca; 2.367, M. modestum variant.

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220(217’) Abdominal sternum 8 with anteromedial brown marking, sometimes becoming crescent shaped on sternum 8 and sometimes also on sternum 7, as illustrated by Figs. 2.368, 2.369 ....................................................................................... Maccaffertium modestum (in part) 220’ Abdominal sternum 7 and 8 without markings as above, either without markings or with variable median or submedian dots, or with longitudinal lines, or with any combination of dots and lines ........................................................................................................................ 221 221(220’) Abdominal sterna usually with markings as in Fig. 2.370; maxillary crowns with 25–40 (usually more than 30) hairlike setae ............................................. Maccaffertium smithae 221’ Abdominal sterna variable, usually not marked as above; maxillary crowns with 15–50 (usually fewer than 30) hairlike setae .......................... Maccaffertium modestum (in part) [Note: A larval form from South Carolina with median, round, dark spots on abdominal sterna will key here. Associated adults differ from typical M. modestum, and it might be a new species.]

Macdunnoa species

(see Flowers, 1982 for more detail) 222(181’) Vestigial gills 7 with spine on both sides of base; posterior marginal denticles of abdominal terga spatulate (Fig. 2.371) ................................................................ Macdunnoa brunnea 222’ Vestigial gills 7 with spine only on outside of base; posterior marginal denticles of abdominal terga pointed (Fig. 2.372) .............................................................. Macdunnoa persimplex

Nixe species

(based on Webb & McCafferty, 2011) 223(185) Hindtibiae with few robust setae on inner face (Fig. 2.374); gills 6 without fibrillae; dorsal abdominal color pattern as in Fig. 2.373 ................................................. Nixe inconspicua [Note: Patterns must be examined carefully because they often become faint. This name possibly is being used to identify two cryptic species.] 223’ Hindtibiae with numerous robust setae on inner face (Fig. 2.375); gills 6 usually with fibrillae; dorsal abdominal pattern not exactly as above . ............................................... 224 224(223’) Robust setae on anterior (outer) face of femora generally blunt or expanded, peglike or paddlelike (e.g., Figs. 2.376, 2.377); one or two pairs of sublateral pale spots on abdominal terga (not including pale lateral areas) ...........................................................225 224’ Robust setae on anterior (outer) face of femora sharply or bluntly pointed (e.g., Fig. 2.378); one pair of sublateral pale spots on abdominal terga (not including pale lateral areas) (e.g., Fig. 2.379) . ......................................................................................................................226

Chapter 2 ~ Key to Mayflies

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Fig. 2.371

Fig. 2.372

Fig. 2.368

Fig. 2.369

Fig. 2.370 Fig. 2.374

Fig. 2.375

Fig. 2.376

Fig. 2.373

Fig. 2.377

Fig. 2.378

Fig. 2.379

Figures 2.368–2.379, Maccaffertium spp., Macdunnoa brunnea, and Nixe spp. (Heptoageniidae). 2.368–2.370, Maccaffertium spp., ends of abdomens, ventral: 2.368, M. modestum variant; 2.369, M. modestum variant; 2.370, M. smithae. 2.371–2.372, Macdunnoa spp., posterior marginal denticles of abdominal tergum 7 (modified from Flowers, 1982): 2.371, M. brunnea; 2.372, M. sp. 2.373–2.379, Nixe spp.: 2.373, N. inconspicua, habitus. 2.374–2.375, posteroventral (inner) surfaces of right hind tibiae: 2.374, N. inconspicua; 2.375, N. perfida. 2.376–2.378, robust setae on anterodorsal (outer) faces of femora: 2.376, peglike robust setae; 2.377, paddlelike robust setae; 2.378, pointed robust setae. 2.379, N. perfida, abdominal terga 5–10, dorsal.

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225(224) Abdominal terga each with single pair of pale spots or areas sublaterally; maxillary crowns with at least 15 robust setae; known provisionally in coverage area only from Alabama ....................................................................................................................... Nixe rusticalis [Note: Robust maxillary crown setae in Nixe are not hairlike, but rather comblike (pectinate), and found on the distal margin of the galealacinia.] 225’ Abdominal terga each with two pairs of pale spots or areas (Fig. 2.380); maxillary crowns with fewer than 15 robust setae; within coverage area known only from Kentucky ......................................................................................................................... Nixe flowersi [Note: Nixe lucidipennis will also key here but is not yet known from the coverage area (N. flowersi has a dense row of long setae on the posterior margin of the foretibiae, which may be absent in N. lucidipennis).] 226(224’) Forefemora (Fig. 2.381) relatively broad and with long, hairlike setae in medial area of outer margin and relatively profuse robust setae on upper face; outer margin of foretibiae with moderately long, fine setae basally becoming shorter distally, and inner margin with few robust setae ...................................................................................... Nixe spinosa [Note: Our concept of the larva of N. spinosa is based on consistent associations of mature larvae with male adults taken at the same time and locale in North Carolina; they also are the only larvae that do not match any other species of eastern North American Nixe, all of which are known in the larval stage. This species has pale and speckled color forms.] 226’ Forefemora (Fig. 2.382) differing in degree in all characteristics given above, e.g., femur somewhat narrower with hairlike setae somewhat shorter and less numerous and surface robust setae less numerous; foretibiae with only short fine setae along outer margin and more numerous robust setae along inner margin ............................................. Nixe perfida

Rhithrogena species

(based in part on Traver, 1935 and Flowers & Hilsenhoff, 1975) 227(177) Mature body length 9–12 mm . ................................................................... Rhithrogena amica 227’ Mature body length 4.5–8.5 mm . ......................................................................................... 228 [Note: Body length never includes caudal filaments. Based on known adult sizes of these species, the unknown larva of R. anomala (known only from Alabama, and possibly the Great Smoky Mountains, within coverage area) would fall within the small body size range given above, whereas the unknown larva of R. rubicunda (known only from North Carolina within coverage area) may fall in the upper part of the lower range, similar to R. manifesta, or less probably within the lower part of the upper size range.].

Chapter 2 ~ Key to Mayflies

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Fig. 2.381

Fig. 2.380

Fig. 2.382

Figures 2.380–2.382, Nixe spp. (Heptageniidae). 2.380, N. flowersi, abdominal terga 5–10. 2.381–2.382, anterodorsal (outer) surfaces of right forefemora, anterodorsal: 2.381, N. spinosa; 2.382, N. perfida

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

228(227’) Abdominal terga 1 and 2 pale yellow; terga 3–6 dark reddish brown and unmarked except for yellow or yellow-hyaline lateral margins; terga 7–9 yellowish, unmarked; small species usually with mature body length 5 mm or less . ........................ Rhithrogena exilis [Note: Color can fade or be lighter in specimens that have been fluid preserved, and also in less than mature or freshly molted individuals.] 228’ Abdominal terga not as above; mature body length variable, as much as 8.5 mm . ............. 229 229(228’) Gills 2–6 (on abdominal segments 2–6) with distinct lobe on margin (Fig. 2.383) (lobe sometimes folded over margin); abdominal terga brown; sometimes terga 7–9 or posterior of 7 and all of 8 and 9 lighter or opaque, and sometimes with reddish posterior margins in more anterior terga ................................................... Rhithrogena impersonata 229’ Gills without distinct lobe as above; because coloration varies, the following couplets should be used with caution . .......................................................................................... 230 230(229’) Abdominal terga relatively uniform in color, 7–9 or 8–9 not distinctly lighter or yellow, and not sharply contrasting with various shades of brown in preceding terga (terga 7–9 or 7–8 often slightly lighter or more opaque than preceding terga) . .................................. 231 230’ Abdominal terga 7–9 or 8–9 distinctly lighter, often bright yellow, and contrasting with darker shades of brown on preceding terga, or abdominal terga concolorous . .............. 232 231(230) Abdominal terga light yellow brown to light chestnut brown to cinnamon brown; conspicuous dorsal abdominal marking not present in darker forms but some diffuse pairs of markings sometimes evident in lighter-colored forms, including pairs of cloudy patches laterally ..................................................................................... Rhithrogena uhari 231’ Abdominal terga medium to dark brown or orange brown; terga 1 and 2 sometimes pale; terga 4–8 sometimes darker anteriorly and paler posteriorly; terga 7–9 often shade lighter than preceding terga ............................................................... Rhithrogena fasciata 232(230’) Abdominal terga 8–9 and sometimes posterior area of 7 much lighter than usually medium brown preceding terga (sometimes difficult to ascertain in light individuals), or abdominal terga concolorous; mature body length usually 7–8.5 mm ............................ ......................................................................................................... Rhithrogena manifesta 232’ Abdominal terga 1–6 and anterior area of 7 dark reddish brown; small species with mature body length usually 5–5.5 mm . ..................................................... Rhithrogena fuscifrons

Chapter 2 ~ Key to Mayflies

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Fig. 2.383 Figures 2.383, Rhithrogena impersonata (Heptageniidae). Left gill 3, left dorsolateral (Flowers & Hilsenhoff, 1975).

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Stenacron species

(based on Lewis, 1978) 233(182) Maxillae galealacinia with submedian longitudinal row of setae consisting of less than 30 setae (Fig. 2.384) ............................................................................................................ 234 [Note: The galealacinia is the main body of the maxilla, with characteristic armature seen in ventral view; the submedian row of setae is a lengthwise row on the ventral face near the inner margin.] 233’ Maxillae galealacinia with submedian longitudinal row of setae consisting of more than 30 setae ........................................................................................................................... 238 234(233) Maxillary crowns with 7–10 robust setae (Fig. 2.384) ......................................................... 235 [Note: The maxillary crown is the distal margin of the main body of the maxilla, from the outer curvature to the tip. Robust setae on the maxillary crown are not hairlike, and they are comprised of various comblike (pectinate) forms.] 234’ Maxillary crowns with 11–13 robust setae ................................................ Stenacron pallidum 235(234) Abdominal terga uniformly brown, without pale longitudinal markings or stripes; abdominal sterna entirely pale ................................................................................ Stenacron carolina 235’ Abdominal terga with pairs of pale stripes or markings submedially and to some extent sublaterally; usually some distinct markings on posterior abdominal sterna ................. 236 236(235’) Angulate (left) mandible with inner margin of outer incisor having five to seven denticles (Fig. 2.385) ..................................................................................................................... 237 236’ Angulate mandible with inner margin of outer incisor having eight denticles; within coverage area known only from Kentucky ........................................ Stenacron candidum 237(236) Abdominal terga with submedian longitudinal stripes continuous or nearly continuous; maxillae galealacinia with 20–30 setae in submedian row and with 8–10 comblike setae on crown (Fig. 2.384); common, widespread and variable ...... Stenacron interpunctatum [Note: S. interpunctatum might be a cryptic species complex.] 237’ Abdominal terga with submedian longitudinal stripes not apparent or not continuous as above; maxillae galealacinia with 20–25 setae in submedian row and with eight or nine comblike setae on crown; within coverage area known only from Florida . ................... ............................................................................................................ Stenacron floridense [Note: S. floridense might be a variant of S. interpunctatum.] 238(233’) Maxillary crowns with 8–10 robust setae in form of distally comblike setae; abdominal sterna with dark bands or spots laterally; within coverage area known only from Kentucky ......................................................................................... Stenacron minnetonka [Note: The maxillary crown is the distal margin of the main body of the maxilla from the outer curvature to the tip.] 238’ Maxillary crowns with 11–13 robust setae in form of laterally comblike setae; abdominal sterna without dark markings . ....................................................... Stenacron gildersleevei

Chapter 2 ~ Key to Mayflies

Fig. 2.384

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Fig. 2.385

Figures 2.384–2.385, Stenacron interpunctatum (Heptageniidae). 2.384, right maxillary galealacinia. 2.385, angulate (left) mandible (outer incisor left), dorsal (modified from Jensen, 1972).

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ISONYCHIIDAE, Isonychia species

(tentative key based on Kondratieff & Voshell, 1984) 239(9) 239’



Filamentous forecoxal gills each with single filament . ........................................................ 240 Filamentous forecoxal gills each with cluster of filaments . ................................................. 244 [Note: This genus is represented in the coverage area by 11 species, only one of which remains unknown as a larva (I. berneri known from the lower tier of states within the coverage area). Isonychia diversa, known only from winged stages, is presumed to be recently extinct and is not included in the above total.]

240(239) Abdominal sterna 6–8 with markings similar to any of Figs. 2.386–2.389; abdominal terga variable shades of brown and usually with dark medial alate markings distinct through cuticle of mature larvae (with or without lighter mid-longitudinal stripe) ..................... 241 [Note: Caution should be used when relying on color characteristics, which are best realized when numerous mature individuals are studied.] 240’ Abdominal sterna 6–8 not exactly matching any of the above variations; abdominal terga often dark reddish brown with partial or complete lighter-shaded mid-longitudinal stripe (alate markings indistinct in mature larvae) . ........................................................ 242 241(240) Abdominal sterna 6–8 with markings similar to any of Figs. 2.386–2.388; abdominal terga yellowish brown to dark reddish brown (with or without lighter mid-longitudinal stripe) ............................................................................................................... Isonychia georgiae 241’ Abdominal sterna 6–8 with markings similar to Fig. 2.389; abdominal terga light orange brown with indistinct mid-longitudinal stripe (stripe interrupted by reddish-brown posterior margins of terga 2–9) ............................................................................. Isonychia obscura 242(240’) Abdominal terga with wide mid-longitudinal stripe on terga 1–9 ................. Isonychia serrata 242’ Abdominal terga with wide mid-longitudinal stripe usually only on terga 1–4 or 1–5, if stripe present on terga 1–9 then very narrow and dashlike . ........................................... 243 243(242’) Foretibiae usually each with 20–26 short spinelike setae; not currently known from the area of coverage . ......................................................................................... Isonychia hoffmani [Note: This species is known from Virginia and West Virginia and is expected to occur in the coverage area.] 243’ Foretibiae usually each with 30–38 short spinelike setae . ............................. Isonychia similis 244(239’) Abdominal gills with robust marginal spines laterally and distally at least on more posterior gills (similar to Fig. 2.390) ............................................................................................. 245 244’ Abdominal gills without robust marginal spines on distal margin (similar to Fig. 2.391) ....................................................................................................................... Isonychia sayi

Chapter 2 ~ Key to Mayflies

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Fig. 2.386

Fig. 2.387

Fig. 2.388

Fig. 2.389

Fig. 2.390

Fig. 2.391

Figures 2.386–2.391, Isonychia spp. (Isonychiidae). 2.386–2.389, abdominal sterna 7, ventral (modified from Kondratieff & Voshell, 1984): 2.386, I. georgiae variant; 2.387, I. georgiae variant; 2.388, I. georgiae variant; 2.389, I. obscura. 2.390–2.391, right gills 7, right dorsolateral (Kondratieff & Voshell, 1984): 2.390, I. sp.; 2.391, I. sp.

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245(244) Gills 6–7 (gills on abdominal segments 6–7) with row of minute robust setae along entire length of median sclerotized ridge (e.g., Fig. 2.393); three or more layers of minute robust setae in distal area of sclerotized anterior marginal ridge of gills 6–7 (at least small sections of distal area of ridge usually with approximate row near inner margin of ridge, one or more approximate rows medially or submedially on ridge and approximate row near outer edge of ridge) (Fig. 2.394); dorsal color pattern somewhat variable, with common eastern variant shown in Fig. 2.392 ............................................. Isonychia sicca [Note: I. bicolor often has been misidentified as I. sicca, and morphological characteristics for separating it from I. sicca may not be entirely reliable.] 245’ Gills 6–7 without distinct row of minute robust setae on median sclerotized ridge, scattered setae at most; two or fewer layers of minute robust setae in distal area of sclerotized anterior marginal ridge of gills 6–7 (with approximate row near inner edge and/or approximate row near outer edge of ridge) (e.g., Fig. 2.395); color patterns variable . ... 246 246(245’) Foretibiae dark at base and apex, and whitish medially (as alate coloration visible through mature larval cuticle); dorsal abdominal color pattern somewhat similar to Fig. 2.392, at least on the darker terga .......................................................................... Isonychia arida 246’ Foretibiae entirely brown as seen through cuticle, sometimes brown darkening at base and apex; abdominal terga often as in Figs. 2.396–2.398 ..................................................... 247 247(246’) Abdominal terga with wide pale mid-longitudinal stripe bordered by dark brown (Fig. 2.396) ....................................................................................................... Isonychia tusculanensis 247’ Abdominal terga variable regarding mid-longitudinal stripe (e.g., Figs. 2.397, 2.398); if present, usually not bordered by dark brown . ............................ Isonychia bicolor, I. rufa [Note: Larvae of I. bicolor and I. rufa cannot be separated confidently. Isonychia bicolor is widespread in the Southeast, but I. rufa is known only from Alabama, Arkansas, Kentucky, and Mississippi. Rearing larvae to the more reliably identifiable adults is required for the most confident species diagnoses.]

Chapter 2 ~ Key to Mayflies

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Fig. 2.393

Fig. 2.394

Fig. 2.392 Fig. 2.395

Fig. 2.396

Fig. 2.397

Fig. 2.398

Figures 2.392–2.398, Isonychia spp. (Isonychiidae). 2.392, I. sicca, dorsal habitus. 2.393, I. sicca, median ridge detail of left gill 7, dorsal (modified from Kondratieff & Voshell, 1984). 2.394–2.395, anterior marginal ridge (distal edge) detail of left gills 7 (modified from Kondratieff & Voshell, 1984): 2.394, I. sicca; 2.395, I. bicolor. 2.396–2.398, Isonychia spp., abdominal terga 7–9, dorsal (Kondratieff & Voshell, 1984): 2.396, I. tusculanensis; 2.397, I. bicolor variant; 2.398, I. bicolor variant.

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LEPTOHYPHIDAE genera 248(19)



248’



Body depressed and compact, appearing squat; forelegs with femur broad (width about three-fourths or more of length), bordered by many setae and with setal row traversing dorsal face at, or in, basal third (Fig. 2.399); posterolateral projections of abdominal terga 7–8 longer than midlength of respective terga (Fig. 2.400) . ....... Asioplax, A. dolani [Note: Asioplax dolani is known throughout the coverage area except for Kentucky, Mississippi, and Tennessee, but eventually should be found in those states. A second species of this otherwise southwestern and western genus, A. texana, has been reported from Mississippi but is not known as larvae. Asioplax dolani is the only North American species of Asioplax known to have two-segmented labial palps and short-stout robust setae dorsally on the forefemora.] Body not depressed and compact as above; forelegs with femur narrower than above (usually with width half or less length), with femur not completely bordered with setae, and with traversing dorsal setae at about midlength of femur (e.g., Fig. 2.401); posterolateral projections of 7–8 no more than subequal to midlength of respective terga (e.g., Fig. 2.402) . .......................................................................... Tricorythodes, 249 [Note: Further identification of the larvae of the four species of Tricorythodes reported from the coverage area must be considered tentative until a revision of this genus in North America can be undertaken. The integrity of the taxonomy based on adults is in question, and certain larval color patterns are based on the study of reared materials taken outside the coverage area.]

Chapter 2 ~ Key to Mayflies

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Fig. 2.400 Fig. 2.399

Fig. 2.401

Fig. 2.402

Figures 2.399–2.402, Asioplax dolani and Tricorythodes robacki (Leptohyphidae). 2.399–2.400, Asioplax dolani: 2.399, anterodorsal face of right forefemur, anterodorsal. 2.400, abdominal terga 7–10, dorsal (modified from Allen, 1967). 2.401–2.402, Tricorythodes robacki (modified from Allen, 967): 2.401, anterodorsal face of right forefemur, anterodorsal; 2.402, abdominal terga 7–10, dorsal.

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Tricorythodes species 249(248’) Minute maxillary palps (at midlength of outer margin of galealacinia) one segmented; operculate gills (Fig. 2.403) rounded oblongate, not subtriangulate or strongly subobovate; abdominal terga granular brown, sometimes with darker markings such as submedian and sublateral markings on terga 1–8 (markings often not discernible when terga dark brown), and with posterolateral processes on tergum 9 at least in females appressed to tergum 10 as seen in dorsal view (see previous Fig. 2.402 under Leptohyphidae genera); currently known from Georgia and the Carolinas .................... .......................................................................................................... Tricorythodes robacki [Note: Maxillary palp should be examined as a slidemount of a maxilla under compound magnification. An apical seta associated with the maxillary palp is not counted as a segment; T. robacki and other species of Tricorythodes in the coverage area have such an apical seta, which can be broken off, but T. curvatus, which is not known from the coverage area but is known from Arkansas, has a one-segmented maxillary palp that lacks an apical seta. In addition, T. curvatus has subobovate operculate gills (Fig. 2.404). The relative development of the posterolateral projections on abdominal tergum 9 in Tricorythodes is applicable at least to females, but may not be always be distinctive in males. Only nearly brown to dark brown specimens that morphologically fit T. robacki have been examined from the coverage area and other areas, not yellow or light brown specimens as were noted in the original description of the species. Any relatively small, brown, compact Tricorythodes with somewhat rounded operculate gills and minute maxillary palpi with a long apical seta should be T. robacki in the coverage area] 249’ Minute maxillary palps two or three segmented; operculate gills triangulate, greatly broadened proximally, and more or less sharply curved or angulate medioproximally (e.g., Fig. 2.405); abdominal terga mostly blackish or blackish gray on pale background patterned, posterolateral projections of tergum 9 variously developed .......................... 250

Chapter 2 ~ Key to Mayflies

Fig. 2.403

Page 133

Fig. 2.404

Fig. 2.405

Figures 2.403–2.405, Tricorythodes spp. (Leptohyphidae). Left operculate gills 2 outline: 2.403, T. robacki; 2.404, T. curvatus (modified from Allen, 1977); 2.405, T. sp.

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250(249’) Minute maxillary palps three segmented (segment 2 very short); posterolateral projections of abdominal tergum 9 poorly developed and appressed to tergum 10 at least in females (Figs. 2.406,2.407); abdominal terga 5–10 (usually 2–10) usually dark medially and laterally, or dark across entire tergum, except tergum 10 usually pale submedially, terga 7–10 often ranging from that in Fig. 2.406 to Fig. 2.407; dorsal faces of femora often with various dark spotting, similar to Fig. 2.408, with spotting diffuse, patchy or coalesced; within coverage area known only from North Carolina . ............................... ........................................................................................................ Tricorythodes stygiatus [Note: Color patterns attributed to T. stygiatus are based on reared specimens from northern regions of North America, and may not be entirely reliable in the coverage area.] 250’ Minute maxillary palps two segmented; posterolateral projections of tergum 9 relatively well developed (Figs. 2.409, 2.410); abdominal terga 2–9 each often at least partially light medially (generally, or as longitudinal dashes, spots, ellipses, or triangles, or combinations of such), otherwise color pattern variable, as seen for example, in terga 8–10 (Figs. 2.409, 2.410); faces of femora pale to variously darkened or dark patterned ......................................................................................................................................... 251 251(250’) Abdominal terga 8–10 often similar to Fig. 2.409 especially with respect to conspicuous V-shaped markings . .................................................................. Tricorythodes albilineatus [Note: Caution should be used because degree of detail and sharpness of the color pattern can become degraded. Abdominal color pattern can vary considerably, but Figure 2.409 is most typical of populations studied. Some southeastern records of this species may be attributable to T. allectus and vice versa.] 251’ Abdominal terga 8–10 often patterned somewhat as in Fig. 2.410 ....... Tricorythodes allectus [Note: Color patterns attributed to T. allectus are based on reared specimens from northern regions of North America and may not be entirely reliable in the coverage area.]

Chapter 2 ~ Key to Mayflies

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Fig. 2.406

Fig. 2.407

Fig. 2.408

Fig. 2.409

Fig. 2.410

Figures 2.406–2.410, Tricorythodes spp. (Leptohyphidae). 2.406–2.407, abdominal terga 7–10, dorsal: 2.406, T. stygiatus variant; 2.407, T. stygiatus variant; 2.408, T. stygiatus variant, anterodorsal face of right forefemur, anterodorsal. 2.409–2.410, abdominal terga 8–10, dorsal: 2.409, T. albilineatus; 2.410, T. allectus (modified from Burks, 1953).

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LEPTOPHLEBIIDAE genera 252(3) 252’

Gills 2–7 (attached to abdominal segments 2–7) narrow-elongate, forked from base to near midlength into pair of lanceolate or narrow forks (e.g., Figs. 2.411–2.413), narrow pair of forks of some each branching distally into cluster of outspread filamentous branches (Fig. 2.414) ..................................................................................................................... 253 Gills 2–7 broad-elongate or short and broad; if broad and elongate, then gills with distinctly narrowed apical extension (e.g., Fig. 2.415); if short and broad, then gills with extensive filaments marginally (Fig. 2.416) ................................................................................... 255

253(252) Gills 2–7 with each fork branching distally into cluster of about six filaments as in Fig. 2.414 .................................................................................................... Habrophlebia, H. vibrans 253’ Gills 2–7 not as above . ......................................................................................................... 254 254(253’) Labrum with deep, triangular median emargination; gills 2–7 with distinct lateral tracheation, forked near midlength of gill; posterior marginal spines absent on abdominal terga 1–5 (Fig. 2.417) ...................................................................... Habrophlebiodes, 257 [Note: The three species known from the coverage area are keyed here. A fourth species, H. annulata, known in the Southeast only from Arkansas, is not expected to occur in the coverage area.] 254’ Labrum with broad and shallow median emargination; gills 2–7 with or without distinct lateral tracheation, forked at various distances from base; posterior marginal spines present on abdominal terga 1–10 (Fig. 2.418), though sometimes difficult to discern .......................................................................................................... Paraleptophlebia, 263 255(252’) Gills 2–7 with broad and elongate lamellae (main leaflike portions) terminating medioapically in variously developed extension (e.g., Fig. 2.415); labrum not as broad as head capsule . .............................................................................................................. 256 255’ Gills relatively small, short, and broad with extensive filaments over entire margins (Fig. 2.416); labrum as broad or slightly broader than head capsule (Fig. 2.419); within coverage area, known only from Kentucky ......................................... Traverella, T. lewisi [Note: This Midwestern species has been taken only from the Ohio River on the border between Kentucky and Indiana and elsewhere in North Dakota. It may be extirpated from its historic locales.] 256(255) Gills 2–7 each with both gill lamellae ending in narrow, lanceolate medioapical extension (Fig. 2.420) ............................................................................................. Leptophlebia, 259 256’ Gills 2–7 (e.g., Fig. 2.421) each with dorsal (anterior) lamella with somewhat spatulate or spindle-shaped medioapical extension, and with ventral (posterior) lamella with continually attenuate medioapical extension (no medial broadening along length of extension) .............................................................................................................. Choroterpes, C. basalis

Chapter 2 ~ Key to Mayflies

Fig. 2.411 Fig. 2.412 Fig. 2.413

Fig. 2.417

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Fig. 2.414

Fig. 2.415

Fig. 2.416

Fig.2.418 Fig. 2.419

Fig. 2.420

Fig. 2.421

Figures 2.411–2.421, Leptophlebiidae genn. spp. 2.411–2.416, left gills 4, left dorsolateral: 2.411, Paraleptophlebia debilis (Traver, 1935); 2.412, P. adoptiva (Traver, 1935); 2.413, P. guttata (Traver, 1935); 2.414, Habrophlebia vibrans (Edmunds et al., 1963); 2.415, Leptophlebia sp. (Burks, 1953); 2.416, Traverella lewisi (Allen, 1973). 2.417–2.418, left halves of posterior margins of abdominal terga 4, dorsal: 2.417, Habrophlebiodes sp.; 2.418, Paraleptophlebia sp. 2.419, Traverella lewisi, head, frontal (modified from Allen, 1973). 2.420, Leptophlebia sp., habitus, dorsal. 2.421, Choroterpes basalis, left gill 4, left dorsolateral (Burks, 1953).

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Habrophlebiodes species

(see Berner, 1975 for more details) 257(254) Posterior marginal row of spines on terga 6–10 ................... Habrophlebiodes brunneipennis 257’ Abdomen with posterior marginal row of spines on terga 7–10 . ......................................... 258 258(257’) Dorsum of head and body chestnut brown, without light mid-longitudinal stripe on abdomen; abdominal sterna white to light yellow becoming tan on more posterior sterna; mature body length 5–6 mm ...................................... Habrophlebiodes americana [Note: Body length never includes caudal filaments (tails).] 258’ Head and mesonotum (dorsum of second thoracic segment) pale with some darker lines; pronotum and abdominal terga dusky yellowish, with abdominal terga paler at anterior margins and with pale mid-longitudinal stripe for length of dorsal abdomen; abdominal sterna 1–3 heavily pigmented and abdominal sterna 3–9 with pigmentation at posterolateral margins; mature body less than 4 mm . .......... Habrophlebiodes celeteria [Note: Possible variability in color has not been determined for this species.]

Leptophlebia species (based on Burian, 2001)

259(256) Abdominal terga with numerous spinelike setae on lateral margins (Fig. 2.422) ................ 260 259’ Abdominal terga with very few or no spinelike setae on lateral margins . ........................... 262 260(259) Forelegs with broad tibial and tarsal banding as in Fig. 2.423; inner margin of forefemur with considerable palmate setae (Fig. 2.424) .............................. Leptophlebia intermedia 260’ Forelegs without broad tibial and tarsal banding (if banding present, then much narrower or much fainter); inner margin of forefemur with numerous serrate setae with small serrations on one or both sides of setae (Fig. 2.425) ...................................................... 261 261(260’) Labrum typically with dorsal surface setae as in Fig. 2.426 . ................... Leptophlebia cupida [Note: It may not always be possible to differentiate the larvae of L. cupida and L. nebulosa. Rearing to obtain the more easily differentiated adults is advisable.] 261’ Labrum typically with dorsal surface setae as in Fig. 2.427 . ............... Leptophlebia nebulosa 262(259’) Gills 2–7 with apicolateral modifications (short-pointed shoulders) on one or both sides of each lamella, in addition to well-developed medioapical extension (e.g., similar to Fig. 2.428); foreclaws with double row of denticles distally (Fig. 2.430) . ..................... .......................................................................................................... Leptophlebia johnsoni 262’ Gills 2–7 with each lamella with medioapical extension only (Fig. 2.429), with no apicolateral modifications; foreclaws with single row of denticles distally (Fig. 2.431) ........................................................................................................... Leptophlebia bradleyi

Chapter 2 ~ Key to Mayflies

Page 139

Fig. 2.423

Fig. 2.422

Fig. 2.424

Fig. 2.425

Fig. 2.426

Fig. 2.428

Fig. 2.427

Fig. 2.429

Fig. 2.430

Fig. 2.431

Figures 2.422–2.431, Leptophlebia spp. (Leptophlebiidae). 2.422, L. sp., left half abdominal tergum 4, dorsal. 2.423, L. intermedia, right foreleg, anterodorsal. 2.424–2.425, typical inner marginal robust setae of forefemora: 2.424, L. intermedia; 2.425, L. cupida. 2.426–2.427, labra, anterodorsal: 3.426, L. cupida; 2.427, L. nebulosa. 2.428–2.429, right gills 4, right dorsolateral: 2.428, L. sp.; 2.429, L. bradleyi. 2.430–2.431, right foreclaws, anterior: 2.430, L. johnsoni; 2.431, L. bradleyi.

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Paraleptophlebia species

(based in part on Traver 1935, Jacobus & McCafferty 2004b and new research; see Tiunova & Kluge 2016 for details about Neoleptophlebia) 263(254’) Gills 2–7 forked at one-fourth or more length from base (e.g., Fig. 2.432); gill trachea with distinctly pigmented branches at least in unforked region ............................................. 264 [Note: During final production of this book, species keyed in couplets 264–266 have been moved to the genus Neoleptophlebia (Kluge 1997) by Tiunova & Kluge (2016).] 263’ Gills 2–7 forked near base, usually not more than one-sixth length from base (e.g., Fig. 2.433); gill trachea without distinctly pigmented lateral branches, but often with very short and faint lateral branches ............................................................................... 267 [Note: Middle gills (3–5) are most reliable for gill characterization in Paraleptophlebia. Unfortunately, gills often are dislodged from the body of these fragile mayflies when specimens are handled.] 264(263) Mandibles relatively elongate, with about half length of angulate (left) mandible beyond angulate shelf (Fig. 2.434) ................................................... Paraleptophlebia swannanoa 264’ Mandibles not elongated as above, with less (usually much less) than half length of angulate mandible beyond angulate shelf (Figs. 2.435, 2.436) ..................................................... 265 265(264’) Posterolateral projections present on abdominal segments 8 and 9 similar to Fig. 2.437 ................................................................................. Paraleptophlebia assimilis [Note: Projections on segment 8 are not well formed in young larvae.] 265’ Posterolateral projections present on abdominal segment 9 only . ....................................... 266 266(265’) Gills 3–5 forked about midlength or slightly less from base .......... Paraleptophlebia adoptiva 266’ Gills 3–5 forked at one-fourth to one-third length from base . ........... Parlaeptophlebia mollis 267(263’) Combined length of distal two maxillary palp segments about 1.5× or more length of proximal segment . .......................................................................................................... 268 267’ Combined length of distal two maxillary palp segments about 1.3× or less length of proximal segment . .......................................................................................................... 273 268(267) Posterolateral projections present on abdominal segments 8 and 9 . .................................... 269 [Note: Projections on tergum 8 may not be well formed in young larvae.] 268’ Posterolateral projections present on abdominal segment 9 only . ....................................... 272 269(268) Legs with contrastingly dark area at femur-tibia joint (Fig. 2.438); otherwise with only diffuse shading not forming distinct bands on femur and tibia . ..................................... 277 269’ Legs not as above, with distinct dark banding, including for example, dark areas more centrally on femur and elsewhere on tibia and tarsus ..................................................... 270

Chapter 2 ~ Key to Mayflies

Fig. 2.432

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Fig. 2.433 Fig. 2.434

Fig. 2.435

Fig. 2.436

Fig. 2.438 Fig. 2.437

Figures 2.432–2.438, Paraleptophlebia spp. (Leptophlebiidae). 2.432–2.433, right gills 4, right dorsolateral: 2.432, P. mollis (Traver, 1935); 2.433, P. jeanae. 2.434–2.436, angulate (left) mandibles, dorsal: 2.434, P. swannanoa (Traver, 1935); 2.435, P. assimilis; 2.436, P. mollis. 2.437, P. assimilis, abdominal terga 8–10 outline, dorsal. 2.438, P. ontario, right forefemur and base of tibia, anterior.

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270(269’) Femora with two dark bands; abdominal terga without pair of dark saggital streaks (lacking longitudinal streaks juxtaposed at either side of midline); not currently known from coverage area . ........................................................................ Paraleptophlebia sticta [Note: Larvae have been associated with adults by rearing. The species occurs in Indiana and Illinois (possibly Wisconsin) and may eventually be found in the coverage area, especially Kentucky.] 270’ Femora with single dark band; abdominal terga sometimes with pair of dark, saggital streaks ................................................................................................................ 271 271(270’) Femora with dark band situated distally; abdominal terga without distinctive saggital streaks ........................................................................ Paraleptophlebia georgiana [Note: Larvae were recently associated with P. georgiana by rearing specimens from the type locale; it remains known only from this location in northern Georgia.] 271’ Femora with dark band situated medially; each abdominal tergum usually with distinctive pair of short to long saggital streaks (e.g., Fig. 2.439) ............ Paraleptophlebia calcarica [Note: Paraleptophlebia calcarica is currently known from Kansas and Arkansas and eventually may be found in the coverage area. Tennessee and Arkansas, for example, have many species of mayflies in common.] 272(268’) Abdominal terga 2–6 with short transverse blackish streak along posterior margin in lateral fourths of terga along with diffuse blackish spot in posterolateral corners; not known from coverage area . .................................................................... Paraleptophlebia strigula [Note: This species is known from the extreme southeastern corner of Kansas, southeastern Ohio, and southern Indiana, and because there is technically a Southeast record from the Ozarks of far eastern Oklahoma, along with a non-specific report of it from Virginia, it eventually may be found in the coverage area.] 272’ Abdominal terga 2–6 laterally without blackish bands as above, but often with diffuse blackish spot in posterolateral corners . ....................................... Paraleptophlebia guttata 273(267’) Legs pale with dark banding ................................................................................................. 274 273’ Legs brown without banding or with only faint, diffuse banding . ....................................... 275 274(273) Maxillary palps with setae of outer margin of segment 1 not confined to single approximate longitudinal row (e.g., Fig. 2.440), often with two or more layers of setae across or around outer face, and sometimes appearing as distinct patch of setae; segment 1 of labial palps subequal in length to segments 2 and 3 combined . ..............................Paraleptophlebia debilis 274’ Maxillary palps with outer margin of segment 1 with more-or-less single longitudinal row of setae, never appearing as a patch of setae or layered across or around the outer face (Fig. 2.441); segment 1 of labial palps shorter than segments 2 and 3 combined ...................................................................................................... Paraleptophlebia jeanae

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Fig. 2.439

Fig. 2.440

Fig. 2.441

Figures 2.439–2.441, Paraleptophlebia spp. (Leptophlebiidae). 2.439, P. calcarica, abdominal tergum 5, dorsal. 2.440–2.441, basal segments of left maxillary palps (base down, outer margin right), ventral: 2.440, P. debilis; 2.441, P. jeanae.

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275(273’) Gills 3–5 of mature or nearly mature larvae bordered with sparse, fine hairlike setae (Fig. 2.442) ................................................................................ Paraleptophlebia volitans 275’ Gills 3–5 of mature or nearly mature larvae with very few or no marginal setae ................ 276 276(275’) Abdominal sterna dark brown laterally and lighter brown medially (Fig. 2.443); mature body length usually slightly less than 7 mm . ........................ Paraleptophlebia praepedita [Note: The lateral sternal shading may be severely degraded in older fluidpreserved material. Body length never includes caudal filaments (tails).] 276’ Abdominal sterna not colored as above; mature body length usually greater than 7 mm ................................................................................................... Paraleptophlebia moerens 277(269) Abdominal sterna 2–8 or at least 6–7 with large, medial, orange spot in anterior half of segments; sternum 7 with additional, reddish spot posteriorly, sometimes coalescing with anterior spot ..................................................................... Paraleptophlebia kirchneri [Note: Spots are best seen in fresh material. Because the extent of variation for this set of characters is unknown, this couplet should be used with caution. Paraleptophlebia kirchneri is known only from Tennessee, and association of larvae is based on collection from the type locality at precisely the correct time.] 277’ Abdominal sterna not exactly as above ............................................ Paraleptophlebia ontario

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Fig. 2.442

Fig. 2443

Figures 2.442–2.443, Paraleptophlebia spp. (Leptophlebiidae). 2.442, P. volitans, left gill 4, left dorsolateral (Ide, 1930). 2.443, P. praepedita, abdominal sterna 6–8, ventral.

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METRETOPODIDAE, Siphloplecton species (see Berner, 1978 for more detail)

278(10)

278’



Gills 1–3 with ventral flap folded under main lamella (Fig. 2.444) ..................................... 279 [Note: The gill flap in Fig. 2.444 is unnaturally depicted with the flap unfolded from its natural position—folded under the main lamella. The gill flap may not be present on early and middle instar larvae. In the following couplets, the sternal patterns should work much of the time. The key must be based on known distribution to some degree and should be regarded as provisional.] Gills 1–3 without ventral flap (Fig. 2.445) ............................................. Siphloplecton fuscum [Note: This is the only species from the coverage area (Florida only) known as larvae that lacks ventral flaps on gills 1–3. Siphloplecton costalense is known from Virginia and North Carolina; its currently unknown larva may key here. Siphloplecton interlineatum is a primarily northern and central species lacking the ventral flap, and it has been reported from nearby Arkansas; it probably does not occur in the coverage area.]

279(278) Ventral abdomen with only partial dark mid-longitudinal stripe or with no mid-longitudinal stripe .......................................................................................... Siphloplecton basale (in part) [Note: The larva of S. simile Berner, which remains unknown, probably will key here; it likely is smaller than S. basale. The two species might be synonymous.] 279’ Ventral abdomen with entire dark mid-longitudinal stripe ................................................... 280 [Note: It becomes apparent from the following couplets that while S. basale may be differentiated in some instances, other species may not be exclusively differentiated as larvae.] 280(279’) Ventral abdomen also with pair of lateral longitudinal stripes ..............................................281 280’ Ventral abdomen either with pair of lateral lines of dark spots (generally two spots on either side of each sternum), or without stripes or spots laterally .................................................. 282 281(280) Known only from Florida .................................................................. Siphloplecton brunneum 281’ Widespread in coverage area ..................................................... Siphloplecton basale (in part) 282(280’) Known from Florida, Georgia, and Alabama .................................... Siphloplecton speciosum 282’ Known from above states and Carolinas ................................... Siphloplecton basale (in part)

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Fig. 2.444

Fig. 2.445

Figures 2.444–2.445. Siphloplecton spp. (Metretopodidae) (modified from Berner, 1978 photos): 2.444, S. basale, right gill 2 with ventral flap unfolded from beneath, right dorsolateral. 2.445, S. fuscum, right gill 2, right dorsolateral.

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NEOEPHEMERIDAE, Neoephemera species

(based in part on Berner, 1956 and Bae & McCafferty, 1998) 283(20’) Pronotum with highly developed anterolateral processes (Fig. 2.446); mature body length 14–17 mm ..................................................................................... Neoephemera purpurea 283’ Pronotum with poorly developed anterolateral processes (Fig. 2.447–450); mature body length 8–11 mm .............................................................................................................. 284 284(283’) Mesonotum (dorsum of second thoracic segment) with bluntly pointed anterolateral processes (Fig. 2.447, 2.449, 2.450); pronotum with pair of conical tubercles submedially near anterior margin (Fig. 2.447, 2.449, 2.450) .............................................................. 285 284’ Mesonotum with broadly rounded anterolateral processes (Fig. 2.448); pronotum with, at most, a pair of poorly developed, low-rounded, rudimentary tubercles submedially, near the anterior margin (Fig. 2.448) ................................................ Neoephemera youngi [Note: Neoephemera sp. B, a new species from North Carolina, may key to this couplet. A manuscript about the species has been submitted to Zootaxa for review. The most prominent characteristic of the unnamed species is its short, but very stout, legs.] 285(284) Pronotal tubercles very close (Fig. 2.447, 2.449) ............................. Neoephemera compressa 285’ Pronotal tubercles widely separated (Fig. 2.450) ................................... Neoephemera bicolor [Note: Neoephemera bicolor is documented from Missouri, and its range may extend further into the study area and include the Carolinas.]

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Fig. 2.447

Fig. 2.446

Fig. 2.449

Fig. 2.448

Fig. 2.450

Figures 2.446–2.450, Neoephemera spp. (Neoephemeridae). 2.446, N. purpurea, habitus, dorsal (Edmunds et al., 1963). 2.447–2.448, outlines of pronota and mesonota with wingpads, dorsal (modified from Berner, 1956): 2.447, N. compressa; 2.448, N. youngi. 2.449–2.450, details of pronota and mesonota, dorsal (Berner, 1956): 2.449, N. compressa; 2.450, N. bicolor.

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OLIGONEURIIDAE, Homoeoneuria species (see Pescador & Peters, 1980 for more detail)

286(9’) 286’

Abdominal terga with distinct cross bands of shading near posterior margins of terga; distal portion of foretibiae (most distal segment of highly modified forelegs) slightly concave and relatively long distad of tarsal nodule as in Fig. 2.452; forecoxae (Fig. 2.451, Cx) with dark basal mark . .............................................................. Homoeoneuria cahabensis Abdominal terga without shaded cross bands as above, either without markings or, at most, with slight transverse penciling at posterior margins of terga; distal portion of foretibiae straight or very slightly convex and relatively short distad of tarsal nodule as in Fig. 2.453; outer face of forecoxae without dark basal mark .......... Homoeoneuria dolani

POLYMITARCYIDAE genera 287(16) 287’



Tusks with scattered spines dorsally (Fig. 2.454); narrow frontal process present (Fig. 2.454); foretibiae not overlapping claw with anterodistal extension (Fig. 2.454) ........................................................................................................................ Ephoron, 288 Tusks with only lateral series of spines along with a large, subdistal spine on inner margin (Fig. 2.455); frons more or less a straight shelf, with no medial process (Fig. 2.455); foretibiae overlapping claw with anterolateral extension (Fig. 2.455) ............................................................................................................... Tortopsis, T. puella [Note: Although the mainly central plains species T. primus is reportedly known from a few records in Arkansas and Louisiana, it may or may not occur in the coverage area (e.g., in the Mississippi River). Larvae of T. primus putatively can be differentiated from T. puella by the comparative development of the lateral horns of the frontal shelf of the head, with the horns extending slightly beyond the antennal scapes in T. puella and extending not quite as far as the antennal scapes in T. primus (McCafferty, 1975). Taxonomy of the North American species needs further study.]

Ephoron species 288(287) Gills 2–7 with distinct extensive lateral tracheation (Fig. 2.456) .................... Ephoron leukon 288’ Gills 2–7 without apparent lateral tracheation (Fig. 2.457); within the coverage area known only from Kentucky ................................................................................... Ephoron album [Note: Number of tusk spines (as used, for example, by McCafferty, 1975) is not a reliable character for separating the larvae in Ephoron. In rare instances in streams where E. leukon and E. album co-occur, they are subject to hybridization, with gill tracheation in some hybrid larvae partially developed in one lamella of the gill or only portions of lamellae. Adults of the two species are symmorphic; the most reliable differentiating character for species is sculpturing of the egg chorion.]

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Fig. 2.452

Fig. 2.451

Fig. 2.453

Fig. 2.454

Fig. 2.455

Fig. 2.456

Fig. 2.457

Figures 2.451–2.457, Homoeoneuria spp. (Oligoneuriidae) and Polymitarcyidae genn. spp. 2.451–2.453, Homoeoneuria spp. (modified from Pescador and Peters, 1980): 2.451, H. cahabensis, head and thorax, left lateral [Cx = coxa, Tr = trochanter, Fe = femur, Tb = tibia]. 2.452–2.453, left foretibiae with preapical nodule up, left lateral: 2.452, H. cahabensis; 2.453, H. dolani. 2.454–2.457, Polymitarcyidae genn. spp.: 2.454–2.455, head, pronotum, and right foreleg, dorsal: 2.454, Ephoron sp.; 2.455, Tortopsis sp. 2.456–2.457, Ephoron spp., left gills 4, left dorsolateral: 2.456, E. leukon; 2.457, E. album.

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POTAMANTHIDAE, Anthopotamus species (modified from Bae & McCafferty, 1991)

289(14) 289’



Tusks narrowing abruptly along outer margin basad of midlength of tusk (Fig. 2.458); elongate narrow spine produced apically on inner margin of foretibiae (Fig. 2.459); mature body typically relatively short, 9–11 mm ........................................................... 291 [Note: Body length never includes caudal filaments (tails).] Tusks not as above, gradually and evenly attenuating to apex, with unwavering outer margin (Fig. 2.460, 2.461) or with only slight waver of continuous line of outer margin slightly basad of midlength (Fig. 2.462); inner margin of foretibiae produced into acute process (Figs. 2.460, 2.461) but never into long sharp spine as above; mature body typically relatively long, 13–18 mm ............................................................................... 290 [Note: If it is unclear which type of tusk a specimen has, then the presence or absence of the elongate spine of the foretibia (not to be confused with an acute process) may be considered reliable.]

290(289’) Upper (forward or outward) face of femora without hairlike setae (Fig. 2.460); pronotum with well-developed lateral flange and with anterolateral corner produced into short spine (Fig. 2.460) .............................................................................. Anthopotamus myops 290’ Upper (forward or outward) face of femora with scattered hairlike setae (Fig. 2.461), with degree of hairiness varying: pronotum usually with only moderately developed lateral flange (sometimes more developed) and with anterolateral corner not produced into short spine (Fig. 2.461), although corner itself often acute ......... Anthopotamus distinctus 291(289) Dorsal abdominal pale markings minimal (e.g., Fig. 2.463) or lacking, with pairs of small pale spots sometimes present on some terga, and narrow mid-longitudinal stripe often present and sometimes faint; tusks (outer straight-line length visible in dorsal view of head) about as long as, or slightly longer than, midlength of head .......................................................................................................... Anthopotamus verticis [Note: Keep in mind that although a mid-longitudinal light stripe is shown for both A. verticis and A. neglectus in Figs. 2.463 and 2.464, these stripes sometimes are absent in both species.] 291’ Dorsal abdominal pale marking more extensive (e.g., Fig. 2.464), including large submedian paired spotting and lightening at anterior and posterior margins of most terga, mid-longitudinal stripe present or absent; tusks (outer straight-line length visible in dorsal view of head) usually slightly shorter than head .......... Anthopotamus neglectus

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Fig. 2.461

Fig. 2.458

Fig. 2.459

Fig. 2.462

Fig. 2.463

Fig. 2.464

Fig. 2.460 Figures 2.458–2.464, Anthopotamus spp. (Potamanthidae). 2.458–2.459, A. verticis: 2.458, left half of head and left tusk, dorsal; 2.459, apex of left foretibia and base of tarsus (up), anterodorsal. 2.460, A. myops, habitus, dorsal. 2.461–2.462, A. distinctus: 2.461, head, pronotum, and left foreleg, dorsal; 2.462, left half of head and left tusk variation, dorsal. 2.463–2.464, abdominal terga 6–8, dorsal: 2.463, A. verticis; 2.464, A. neglectus.

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SIPHLONURIDAE, Siphlonurus species

(based in part on Traver, 1935 and Provonsha & McCafferty, 1982) 292(12) 292’



Abdominal gills 1–6 or 1–7 with dorsal recurved flap ........................ Siphlonurus alternatus [Note: This species has not been reported from the coverage area, but it is present in Indiana and Missouri and eventually may be found in Kentucky.] Only abdominal gills 1 and 2 with dorsal recurved flap ...................................................... 293 [Note: Dorsal flaps of gills may make them appear double. Among species known from the coverage area, only S. decorus remains unknown as a larva. Siphlonurus luridipennis is presumed to be recently extinct.]

293(292’) Caudal filaments (tails) not dark banded near middle; legs without dark banding; head with pair of dark oblique bands below antennae (Fig. 2.465); abdomen often well patterned with dark and reddish-brown markings ................................................. Siphlonurus mirus 293’ Caudal filaments with dark band near middle, usually just distad of middle; legs variously with some brown or dark banding; head and body patterning variable [Note: Banding of caudal filaments is subject to fading.] ........................................................................ 294 294(293’) Abdominal sterna light tan or light brown, each with pale gray or pale yellow partially conical area medially, best developed on sterna 2–6 as in Fig. 2.466, sterna sometimes diffuse brown far laterally; otherwise any staining or patches faint or diffuse .................................................................................................... Siphlonurus quebecensis 294’ Ventral abdomen pale cream to yellow or light brown, not patterned as above but with at least some distinctive brown or dark stripes ................................................................... 295 295(294’) Abdominal sterna 3–9 with mid-longitudinal dark stripe (partial on sternum 2); mature body length (excluding caudal filaments) 9–10 mm . .................. Siphlonurus marginatus 295’ Abdominal sterna without mid-longitudinal stripe; mature length 11–16 mm . ................... 296 296(295’) Hindwingpads with adult wing brown; abdominal sterna with lateral stripes and four submedian dots ............................................................................... Siphlonurus marshalli [Note: This species has not been reported from the coverage area, but it has been reported from West Virginia and Arkansas. Coloration of wingpads is obscured in ultimate instar larvae that have darkened fore and hindwingpads as a natural pre-emergent condition; otherwise wingpad characterization may be reliable only in nearly mature larvae. Ventral markings may not be discernible in some females and immature larvae.] 296’ Hindwingpads not darkened as above (except for fully mature, pre-emergent larvae); abdominal sterna, at least in more mature males, with lateral stripes but no submedian markings (Fig. 2.468) or with lateral stripes and submedian dots, dashes, and diagonal bars (Fig. 2.467) . ............................................................................................................ 297 297(296’) Abdominal sterna as shown in Fig. 2.467, but stripes and bars reduced to short dashes in some females and often in immature larvae ....................................... Siphlonurus minnoi 297’ Abdominal sterna as shown in Fig. 2.468, but stripes indistinct and sometimes with diffuse staining in some females and immature larvae .................................... Siphlonurus typicus

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Fig. 2.465

Fig. 2.467

Fig. 2.466

Fig. 2.468

Figures 2.465–2.468, Siphlonurus spp. (Siphlonuridae). 2.465, S. mirus, head, frontal. 2.466–2.468, abdominal sterna 2–6, ventral: 2.466, S. quebecensis; 2.467, S. minnoi; 2.468, S. typicus.

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References Cited Allen, R.K. 1967. New species of New World Leptohyphinae (Ephemeroptera: Tricorythidae). Canadian Entomologist 99: 349–375. Allen, R.K. 1973. Generic revision of mayfly nymphs. 1. Traverella in North and Central America (Leptophlebiidae). Annals of the American Entomological Society 66: 1287–1295. Allen, R.K. 1977. New species of Tricorythodes with notes (Ephemeroptera: Tricorythidae). Journal of the Kansas Entomological Society 50: 431–435. Allen, R.K., & G.F. Edmunds, Jr. 1961. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). III. The subgenus Attenuatella. Journal of the Kansas Entomological Society 34: 161–173. Allen, R.K., & G.F. Edmunds, Jr. 1962. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). V. The subgenus Drunella in North America. Miscellaneous Publications of the Entomological Society of America 3: 147–179. Allen, R.K., & G.F. Edmunds, Jr. 1963a. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). VII. The subgenus Eurylophella. Canadian Entomologist 95: 597–623. Allen, R.K., & G.F. Edmunds, Jr. 1963b. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). VI. The subgenus Serratella in North America. Annals of the Entomological Society of America 56: 583–600. Allen, R.K., & G.F. Edmunds, Jr. 1965. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). VIII. The subgenus Ephemerella in North America. Miscellaneous Publications of the Entomological Society of America 4: 243–282. Bae, Y.J., & W.P. McCafferty. 1991. Phylogenetic systematics of the Potamanthidae (Ephemeroptera). Transactions of the Entomological Society of America. 117: 1–143. Bae, Y.J., & W.P. McCafferty. 1998. Phylogenetic systematics and biogeography of the Neoephemeridae (Ephemeroptera: Pannota). Aquatic Insects 20: 35–68. Barber-James, H.M., J.-L. Gattolliat, M. Sartori, & M.D. Hubbard. 2008. Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater. Hydrobiologia 595: 339–350. Barber-James, H.M., M. Sartori, J.-L. Gattolliat, & J. Webb. 2013. World checklist of freshwater Ephemeroptera species. Available from http://fada.biodiversity.be/group/show/35 (Accessed 18 December 2013). Bednarik, A.F., & W.P. McCafferty. 1979. A biosystematic revision of the genus Stenonema (Ephemeroptera: Heptageniidae). Canadian Bulletin of Fisheries and Aquatic Sciences 201: i–v + 1–73. Berner, L. 1956. The genus Neoephemera in North America (Ephemeroptera: Neoephemeridae). Annals of the Entomological Society of America 49: 33–42. Berner, L. 1975. The mayfly family Leptophlebiidae in the southeastern United States. Florida Entomologist 58: 137–156. Berner, L. 1978. A review of the mayfly family Metretopodidae. Transactions of the American Entomological Society 104: 91–137. Burian, S.K. 2001. A revision of the genus Leptophlebia Westwood in North America (Ephemeroptera: Leptophlebiidae: Leptophlebiinae). Ohio Biological Survey, New Series 13: 1–80. Burian, S.K., B.I. Swartz, & P.C. Wick. 2008. Taxonomy of Epeorus frisoni (Burks) and a key to New England species of Epeorus. In: Hauer, F.R., J.A. Stanford, & R.L. Newell (Eds.), International Advances in the Ecology, Zoogeography and Systematics of Mayflies and Stoneflies. University of California Publications in Entomology, Vol. 128, pp. 277–294. Burks, B.D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bulletin of the Illinois Natural History Survey 26: 1–216.

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Check, G.R. 1982. A revision of the North American species of Callibaetis (Ephmeroptera: Baetidae). Unpublished Masters thesis, University of Minnesota, Minneapolis, Minnesota, USA. 155 pp. Day, W.C. 1955. New genera of mayflies from California. Pan-Pacific Entomologist 31: 121–137. Edmunds, G.F., Jr., L. Berner, & J.R. Traver. 1958. North American mayflies of the family Oligoneuriidae. Annals of the Entomological Society of America 51: 375–382. Edmunds, G.F. Jr., R.K. Allen, & W.L. Peters. 1963. An annotated key to the nymphs of the families and subfamilies of mayflies (Ephemeroptera). University of Utah Biological Series 13 (1): 1–49. Flowers, R.W. 1982. Review of the genus Macdunnoa (Ephemeroptera: Heptageniidae) with description of a new species from Florida. Great Lakes Entomologist 15: 25–30. Flowers, R.W., & W. L. Hilsenhoff. 1975. Heptageniidae (Ephemeroptera) of Wisconsin. Great Lakes Entomologist 8: 201–218. Funk, D.H., & B.W. Sweeney. 1994. The larvae of eastern North American Eurylophella Tiensuu (Ephemeroptera: Ephemerellidae). Transactions of the American Entomological Society 120: 209–286. Funk, D.H., J.K. Jackson, & B.W. Sweeney. 2006. Taxonomy and genetics of the parthenogenetic mayfly Centroptilum triangulifer and its sexual sister Centroptilum alamance (Ephemeroptera: Baetidae). Journal of the North American Benthological Society 25: 417–429. Funk, D.H., J.K. Jackson, & B.W. Sweeney. 2008a. A new parthenogenetic mayfly (Ephemeroptera: Ephemerellidae: Eurylophella Tiensuu) oviposits by abdominal bursting in the subimago. Journal of the North American Benthological Society 27: 269–279. Funk, D.H., B.W. Sweeney, & J.K. Jackson. 2008b. A taxonomic reassessment of the Drunella lata (Morgan) species complex (Ephemeroptera: Ephemerellidae) in northeastern North America. Journal of the North American Benthological Society 27: 647–663. Ide, F.P. 1930. Contribution to the biology of Ontario mayflies with descriptions of new species. Canadian Entomologist 62: 204–213, 218–231. Ide, F.P. 1937. Descriptions of eastern North American species of baetine mayflies with particular reference to the nymphal stages. Canadian Entomologist 69: 219–231, 235–243. Jacobus, L.M. 2006. Phylogeny and classification of the mayfly subfamily Ephemerellinae (Ephemeroptera: Ephemerellidae). Unpublished PhD dissertation, Purdue Univeristy, West Lafayette, Indiana, USA. 156 pp. Jacobus, L.M. 2013. South Carolina mayflies (Insecta: Ephemeroptera) of conservation concern. Journal of the South Carolina Academy of Science 11: 23–26. Jacobus, L. M., & E.D. Fleek. 2010. Insecta, Ephemeroptera, Ephemerellidae, Attenella margarita (Needham, 1927): Southeastern range extension to North Carolina, USA. Check List 6: 311–313. Jacobus, L.M., & W.P. McCafferty. 2000. Variability in the larvae of Serratella serrata (Ephemeroptera: Ephemerellidae). Entomological News 111: 39–44. Jacobus, L.M., & W.P. McCafferty. 2004a. Contribution to the morphology and descriptive biology of Caurinella idahoensis (Ephemeroptera: Ephemerellidae). Western North American Naturalist 64: 101–108. Jacobus, L.M., & W.P. McCafferty. 2004b. First larval description of Paraleptophlebia calcarica Rowbotham and Allen (Ephemeroptera: Leptophlebiidae). Journal of the Kansas Entomological Society 77: 110–115. Jacobus, L.M., & W.P. McCafferty. 2006. A new species of Acentrella Bengtsson (Ephemeroptera: Baetidae) from Great Smoky Mountains National Park, USA. Aquatic Insects 28: 101–111. Jacobus, L.M., & W.P. McCafferty. 2008. Revision of Ephemerellidae genera (Ephemeroptera). Transactions of the American Entomological Society 134: 185–274. Jacobus, L.M., & J.M. Webb. 2013. A new junior synonym for Raptoheptagenia cruentata (Walsh, 1863) and remarks about Nearctic Heptagenia Walsh, 1863 (Insecta: Ephemeroptera: Heptageniidae). Proceedings of the Indiana Academy of Science 121: 143–146.

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Jacobus, L.M., & N.A. Wiersema. 2014. The genera Anafroptilum Kluge, 2011 and Neocloeon Traver, 1932, reinstated status, in North America, with remarks about the global composition of Centroptilum Eaton, 1869 (Ephemeroptera: Baetidae). Zootaxa 3814: 385–391. Jensen, S.L. 1972. A generic revision of the Heptageniidae of the world (Ephemeroptera). Unpublished PhD dissertation, University of Utah, Salt Lake City, Utah, USA. 266 pp. Keffermüller, M. 1960. Badania nad fauna jetek (Ephemeroptera) Wielkopolski. Prace Komisji Biologicznej Poznanskie Towarzystwo Przyjaciól Nauk, Wydzial-Matematyczno-Przyrodniczy 19(8): 1–57, pl. 1–11. Kluge, N.J. 1997. New subgenera of Holarctic mayflies (Ephemeroptera: Heptageniidae, Leptophlebiidae, Ephemerellidae). Zoosystematica Rossica (1996) 5(2): 233–235. Kluge, N.J. 2015. Ephemeroptera of the world. Available from http://www.insecta.bio.spbu.ru/z/Eph-spp/ Contents.htm (Accessed 22 March 2015). Kondratieff, B.C., & J.R. Voshell. 1984. The North and Central American species of Isonychia (Ephemeroptera: Oligoneuriidae). Transactions of the American Entomological Society 110: 129–244. Lewis, P.A. 1978. Taxonomy and ecology of Stenonema mayflies (Heptageniidae: Ephemeroptera). EPA670/4-74-006, emended 1978 reprint. USEPA National Environmental Research Center, Cincinnati, Ohio, USA. Lugo-Ortiz, C.R., & W.P. McCafferty. 1998a. A new North American genus of Baetidae (Ephemeroptera) and key to Baetis complex genera. Entomological News 109: 345–353. Lugo-Ortiz, C.R., & W.P. McCafferty. 1998b. New larval variants and distributional records for Plauditus cestus (Ephemeroptera: Baetidae). Great Lakes Entomologist 31: 201–204. McCafferty, W.P. 1971. The systematics of the mayfly superfamily Ephemeroidea (Ephemeroptera). Unpublished PhD dissertation, University of Georgia, Athens, Georgia, USA. 205 pp. McCafferty, W.P. 1975. The burrowing mayflies (Ephemeroptera: Ephemeroidea) of the United States. Transactions of the Entomological Society of America 101: 447–504. McCafferty, W.P. 1977a. Biosystematics of Dannella and related subgenera of Ephemerella (Ephemeroptera: Ephemerellidae). Annals of the Entomological Society of America 70: 881–889. McCafferty, W.P. 1977b. Newly associated larvae of three species of Heptagenia (Ephemeroptera: Heptageniidae). Journal of the Georgia Entomological Society 12: 350–358. McCafferty, W.P. 1981a. Aquatic Entomology. Jones & Bartlett, Boston, Massachusetts, USA. 448 pp. McCafferty, W.P. 1981b. A distinctive new species of Stenonema (Ephemeroptera: Heptageniidae) from Kentucky and Missouri. Proceedings of the Entomological Society of Wahsington 83: 512–515. McCafferty, W.P. 1985. New spiny crawlers from headwaters of the Savannah River (Ephemeroptera: Ephemerellidae). Proceedings of the Entomological Society of Washington 87: 421–425. McCafferty, W.P. 1990. A new species of Stenonema (Ephemeroptera: Heptageniidae). Proceedings of the Entomological Society of Washington 92: 760–764. McCafferty, W.P. 1991. Comparison of Old and New World Acanthametropus (Ephemeroptera: Acanthametropodidae) and other psammophilous mayflies. Entomological News 102: 205–214. McCafferty, W.P. 1996. The mayflies of North America online. Entomological News 107: 61–63. McCafferty, W.P. 2001. Status of some historically unfamiliar American mayflies (Ephemeroptera). PanPacific Entomologist 77: 210–218. McCafferty, W.P. 2010. A new species of Maccaffertium Bednarik (Ephemeroptera: Heptageniidae) from the southeastern U.S.A. Transactions of the Entomological Society of America 136: 217–219. McCafferty, W.P. 2011a. A new genus and species of small minnow mayflies (Ephemeroptera: Baetidae) from far northern North America. Transactions of the American Entomological Society 137: 11–14. McCafferty, W.P. 2011b. Notable new North and Central American records of Ephemeroptera species. Transactions of the American Entomological Society 137: 1–10. McCafferty, W.P., & L.M. Jacobus. 2015. Mayfly Central. Available from http://www.entm.purdue.edu/ mayfly/index.php (Accessed 8 July 2015).

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McCafferty, W.P., & D.R. Lenat. 2003. A new Nearctic Paracloeodes (Ephemeroptera: Baetidae). Entomological News 114: 33–36. McCafferty, W.P., & D.R. Lenat. 2010. Maccaffertium wudigeum, a new species of Ephemeroptera (Heptageniidae) from North Carolina. Transactions of the American Entomological Society 136: 185–187. McCafferty, W.P., & A.V. Provonsha. 1986. Comparative mouthpart morphology and evolution of the carnivorous Heptageniidae (Ephemeroptera). Aquatic Insects 8: 83–89. McCafferty, W.P., & R.D. Waltz. 1995. Labiobaetis (Ephemeroptera: Baetidae): new status, new North American species, and related new genus. Entomological News 106: 19–28. McCafferty, W.P., & R.D. Waltz. 1998. A new species of the small minnow mayfly genus Plauditus (Ephemeroptera: Baetidae) from South Carolina. Entomological News 109: 354–356. McCafferty, W.P., R.D. Waltz, J.M. Webb, & L.M. Jacobus. 2005. Revision of Heterocloeon McDunnough (Ephemeroptera: Baetidae). Journal of Insect Science 5(35): 1–11. McCafferty, W.P., R.D. Waltz, & J.M. Webb. 2009. Acentrella nadineae, a new species of small minnow mayflies (Ephemeroptera: Baetidae). Proceedings of the Entomological Society of Washington 111: 12–17. McCafferty, W. P., D.R. Lenat, L.M. Jacobus, & M.D. Meyer. 2010. The mayflies (Ephemeroptera) of the southeastern United States. Transactions of the Entomological Society of America 136: 221–233. McDunnough, J. 1926. Notes on North American Ephemeroptera with descriptions of new species. Canadian Entomologist 58: 184–196. Morihara, D.K., & W.P. McCafferty. 1979a. The Baetis larvae of North America (Ephemeroptera: Baetidae). Transactions of the American Entomological Society 105: 139–221. Morihara, D.K., & W.P. McCafferty. 1979b. Systematics of the propinquus group of Baetis species (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 72: 130–135. Morihara, D.K., & W.P. McCafferty. 1979c. The evolution of Heterocloeon, with the first larval description of Heterocloeon frivolus comb. n. (Ephemeroptera: Baetidae). Aquatic Insects 1: 225–231. Müller-Liebenau, I. 1974. Rheobaetis: a new genus from Georgia (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 67: 555–567. Nieto, C. 2016. The Baetodes complex (Ephemeroptera: Baetidae), phylogeny, biogeography, and new species of Mayobaetis. Freshwater Science 35(1), in press. DOI: 10.1086/684859. Pescador, M.L., & L. Berner. 1981. The mayfly family Baetiscidae (Ephemeroptera). Part II. Biosystematics of the genus Baetisca. Transactions of the American Entomological Society 107: 163–228. Pescador, M.L., & W.L. Peters. 1980. A revision of the genus Homoeoneuria (Ephemeroptera: Oligoneuriidae). Transactions of the American Entomological Society 106: 357–393. Pescador, M.L., & B.A. Richard. 2006. A new species of Caenis (Ephemeroptera: Caenidae) from Florida, USA. Zootaxa 1355: 61–68. Provonsha, A.V. 1990. A revision of the genus Caenis in North America (Ephemeroptera: Caenidae). Transactions of the American Entomological Society 116: 801–884. Provonsha, A.V., & W.P. McCafferty. 1982. New species and previously undescribed larvae of North American Ephemeroptera. Journal of the Kansas Entomological Society 55: 23–33. Provonsha, A.V., & W.P. McCafferty. 2006. A second species of the North American mayfly genus Amercaenis Provonsha and McCafferty (Ephemeroptera: Caenidae). Journal of Insect Science 6(10): 1–6. Randolph, R.P., & W.P. McCafferty. 1996. First larval descriptions of two species of Paraleptophlebia (Ephemeroptera: Leptophlebiidae). Entomological News 107: 225–229. Salles, F.F., J.-L. Gattolliat, & M. Sartori. 2015. Phylogenetic analyses of Cloeodes Traver and related genera (Ephemeroptera: Baetidae). Systematic Entomology. Available from onlinelibrary.wiley.com/ doi/10.1111/syen.12144/full (Accessed 24 November 2015).

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Soldán, T. 1986. A revision of the Caenidae with ocellar turbercles in the nymphal stage (Ephemeroptera). Acta Univeritatis Carolinae – Biologica 1982–1984: 289–362. Sun, L., & W.P. McCafferty. 2008. Cladistics, classification and identification of the bracycercine mayflies (Insecta: Ephemeroptera: Caenidae). Zootaxa 1801: 1–239. Tiunova, T.M., & N. Kluge. 2016. Redescription of Paraleptophlebia falcula Traver 1934 with notes on status and composition of Paraleptophlebia Lestage 1917 and Neoleptophlebia Kluge 1997 (Ephemeroptera: Leptophlebiidae). Zootaxa 4098 (2): 369–382. Traver, J.R. 1934. New North American species of mayflies (Ephemerida). Journal of the Elisha Mitchell Scientific Society 50(1–2): 189–254, pl. 16. Traver, J.R. 1935. Part II: Systematic. In: Needham, J.G., J.R. Traver, & H.-C. Hsu (Eds.), The Biology of Mayflies. Comstock Publishing, Ithaca, New York, USA. Pp. 237–739. Waltz, R.D., & W.P. McCafferty. 1989. New species, redescriptions, and cladistics of the genus Pseudocentroptiloides (Ephemeroptera: Baetidae). Journal of the New York Entomological Society 97: 151–158. Wang, T.-Q., & W.P. McCafferty. 1996. New diagnostic characters for the mayfly family Baetidae (Ephemeroptera). Entomological News 107: 207–212. Webb, J.M., & W.P. McCafferty. 2006. Contribution to the taxonomy of eastern North American Epeorus (Ephemeroptera: Heptageniidae). Zootaxa 1128: 57–64. Webb, J.M., & W.P. McCafferty. 2008. Heptageniidae of the world. Part II: Key to the genera. Canadian Journal of Arthropod Identification 7: 1–55. Webb, J.M., & W.P. McCafferty. 2011. Contributions to the larvae of North American Nixe (Ephemeroptera: Heptageniidae), with description of N. dorothae sp. nov. from southern Indiana. Zootaxa 3065: 27–37. Webb, J.M., L. Sun, W.P. McCafferty, & V.R. Ferris. 2007. A new species and new synonym in Heptagenia Walsh (Ephemeroptera: Heptageniidae: Heptageniinae) based on molecular and morphological evidence. Journal of Insect Science 7(63): 1–16. Webb, J.M., L.M. Jacobus, D.H. Funk, X. Zhou, B. Kondratieff, C.J. Geraci, R.E. DeWalt, D. Baird, B. Richard, I. Phillips, & P.D.N. Hebert. 2012. A DNA barcode library for North American Ephemeroptera: Progress and prospects. PLoS ONE 7(5): e38063. Available from http://journals.plos.org/plosone/ article?id=10.1371/journal.pone.0038063 (Accessed 23 November 2015). Wiersema, N.A. 1999. Two new species of Procloeon (Ephemeroptera: Baetidae) from Texas. Entomological News 110: 27–35. Wiersema, N.A. 2000. A new combination for two North American small minnow mayflies (Ephemeroptera: Baetidae). Entomological News 111: 140–142. Wiersema, N.A., & L.S. Long. 2000. Plauditus grandis (Ephemeroptera: Baetidae), a new small minnow mayfly from Tennessee. Entomological News 111: 45–48. Wiersema, N.A., & W.P. McCafferty. 2000. Generic revision of the North and Central American Leptohyphidae (Ephemeroptera: Pannota). Transactions of the American Entomological Society 126: 337–371. Wiersema, N.A., & W.P. McCafferty. 2005. Contribution to the taxonomy of Asioplax (Ephemeroptera: Leptohyphidae: Tricorythodinae). Entomological News 116: 147–158. Wiersema, N.A., C.R. Nelson, & K.F. Kuehnl. 2004. A new small minnow mayfly (Ephemeroptera: Baetidae) from Utah, U.S.A. Entomological News 115: 139–145.

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Chapter 3

PLECOPTERA B.P. Stark

The order Plecoptera, a small group of hemimetabolous insects, includes approximately 3,500 extant world species of stoneflies placed in 16 families (DeWalt et al., 2015; Fochetti & Tierno de Figueroa, 2008; Stark et al., 2009). Two suborders, Arctoperlaria and Anarctoperlaria, are currently recognized and the Arctoperlaria, which includes all Nearctic stoneflies, are further classified in group Systellognatha or Euholognatha (Zwick, 1973). In North America nine families, all of which occur in the southeastern region, and about 715 species are known (DeWalt et al., 2015; Stark et al., 2012; Szczytko & Kondratieff, 2015). Table 3.1 presents a systematic list of 294 species currently recognized from, or adjacent to, the southeastern region. Although reliable keys exist for identification of adult and larval stoneflies to family and genus level (e.g. Stewart & Stark, 2002, 2008), in most areas of North America, including the eight southeastern states that comprise USEPA Region 4, species identification requires extensive use of the primary literature. Many of the classic publications (e.g. Frison, 1942; Needham & Claassen, 1925; Ricker, 1952; Ross & Ricker, 1971) are still essential, and some, more recent works, (e.g., Stark & Armitage, 2000, 2004) provide species keys to adults for entire families from eastern North America. Unfortunately, larvae were not included in the Stark & Armitage (2000, 2004) series and the studies of only five families (Chloroperlidae, Peltoperlidae, Perlidae, Pteronarcyidae, Taeniopterygidae) and subfamily Perlodinae were published (Kondratieff, 2004; Nelson, 2000; Stark, 2000, 2004; Stewart, 2000; Surdick, 2004). Recently, Szczytko & Kondratieff (2015) completed a monograph of eastern Nearctic Isoperlinae in which 22 new Isoperla species were described. All but one of these is known from USEPA Region 4 and larval descriptions are lacking for each of these new species. Literature for species identification of most North American stonefly larvae does not exist and larval stages for the majority of species are still unassociated or undescribed. The stonefly key presented below includes 49 genera and 162 species; the latter figure represents about 55% of the species recorded from, or adjacent to states in USEPA Region 4 (Table 3.1). Despite these problems, several studies assist in recognizing larval stonefly species on a regional basis (Harper & Hynes, 1971a, 1971b, 1971c, 1971d; Stark & Lacey, 2005), and in a few cases (e.g., Earle & Stewart, 2008; Fullington & Stewart, 1980; Kondratieff et al., 1988; Myers & Kondratieff, 2009; Sandberg & Stewart, 2005; Stark & Szczytko, 1981) all, or most larvae of the known species for a single genus, have been described and keyed in a single publication. These studies and others in which even single species have been described (e.g., Brown & Stark, 1995; Kondratieff & Kirchner, 1988; Nelson, 1997) provide the basis for the keys that follow. Most supporting figures are original to this key, but some are borrowed from Stewart & Stark (2002) or redrawn from various other sources. Nearctic stonefly larvae are distinguished from those of similar aquatic orders by having external wingpads similar in size and shape, mouthparts of the chewing variety, three tarsal segments, two tarsal claws, paired cerci with no median caudal filament, and by lacking paired gills on the dorsum of the abdomen. One California perlodid species, Oroperla barbara Needham, has paired, finger-like, ventrolateral gills on abdominal segments 1-7 (Stewart & Stark, 2002), but this is the nearest exception to that character among Nearctic stoneflies. Among aquatic insects, stonefly larvae are superficially similar to those of Ephemeroptera, but mayfly larvae have the anterior wingpads much larger than hind wingpads (if present), a single tarsal segment and claw, paired dorsal or dorsolateral abdominal gills and often a median caudal filament in addition to paired cerci. Figs. 3.1–3.2 show dorsal and ventral morphological features for Eccoptura xanthenes (Newman), a typical stonefly larva. Figs. 3.1, 3.3–3.10 show the larval habitus for each of the nine stonefly families known for North America. Variations found on the following structures are used extensively in the keys below.

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Gills: Many stonefly species lack gills, but among those with gills, position and structural form may vary (Shepard & Stewart, 1983, Stewart & Stark, 2002). The gills consist of single, finger-like structures (some Perlodidae), segmented or telescoping structures (Taeniopteryx), long, single or double conical structures (some Peltoperlidae) or highly branched structures (Amphinemura, Perlidae, Pteronarcyidae). Positions occupied by gills of southeastern stoneflies include the submentum (some Perlodidae); neck or cervical area (some Nemouridae); coxae (Taeniopteryx); on the thorax, above, in front of, or behind the coxae (Peltoperlidae, Perlidae, Pteronarcyidae); ventrally on the first two abdominal segments (Pteronarcyidae); or on the paraprocts (some Perlidae, Peltoperlidae). In Fig. 3.2, highly branched thoracic and anal gills (located on paraprocts) are shown, and additional gill variations are shown in Figs. 3.11–3.14. Mouthparts: Stonefly larvae have well-developed mandibles and maxillae, a simple labrum, and multilobed labium (Stewart & Stark, 2002). Species with primarily carnivorous food habits have the maxillae, mandibles, and labium modified for prey detection, capture, and processing, whereas those with primarily herbivorous or detritivorous food habits have these same structures modified for shredding, scraping, or sweeping periphyton or leaf litter. In the former case, the maxillary laciniae have 1 or 2 relatively long spine-like teeth, the galeae lack a terminal brush or comb-like structure and instead, bear one or more long, slender sensory setae, the mandibles lack a mola and the glossae are much smaller than the paraglossae. Herbivorous-detritivorous species have maxillary laciniae with 2 or 3 short apical teeth grouped into a scoop-like structure, the galea may display a terminal brush or comb-like structure, the mandibles have a prominent mola, and the glossae and paraglossae of the labium extend forward about the same distance. Setation: The body and appendages of stonefly larvae are covered by several types of setae: long, thin fringe setae of the legs and sometimes cerci; shorter (but of variable length), thicker setae of the cercal and body segmental whorls or fringes; fine, decumbent clothing hairs scattered on the body surface; and short peg-like setae (or sensilla) often found on the occiput. Short, thick setae found on the surface of a segment, independently of those in the posterior whorl of the same segment, are called intercalary setae; those found behind the compound eye often occur in a semicircular row called the postocular fringe. Fine fringe setae usually occur on the outer margins of legs, but in some groups the inner tibial margin may bear at least a few of these hairs. Unfortunately they are easily lost through handling. Many of these setal types are shown in Fig. 3.1. Wingpads: Reduction of wings in brachypterous or micropterous species is relatively rare among stoneflies in the southeastern region. Only among Capniidae has this been noted in the region, and only in a few species of Allocapnia and Paracapnia is wing reduction so extreme that wingpads are eliminated in mature larvae. The primary use of wingpad morphology in the key involves the relative divergence of these structures from the body axis. In some groups (e.g,. Nemouridae, Fig. 3.5) the wingpads diverge strongly from the central body axis, whereas in other groups (e.g., Leuctridae, Fig. 3.4) they lie parallel to the body axis. In addition to these basic features, the setal arrangements, particularly along outer margins of the wingpads, can be useful. Abdomen: Stonefly larvae have 10 distinct abdominal segments and the remnants of segment 11, which provides a morphological platform for the cerci. Abdominal segments may consist of completely sclerotized rings with the terga and sterna fused laterally, or one or more segments may be divided laterally by a pleural membrane; in some groups the first abdominal sternum is fused to the metathorax. Variations associated with the number of completely fused segments are often used in larval stonefly keys, particularly to separate Capniidae (segments 1–9 divided laterally) from Leuctridae (segments 1–4, 1–5, 1–6 or 1–7 divided laterally). These characters, though difficult to see, are retained in the key that follows (see couplet 8, for example), but their use in separating these families is minimized through addition of other characters including the relative size of the mentum (Zwick, 2004, 2006). In some groups (e.g., Pteronarcyidae, Capniidae males, etc.) tergum 10 projects strongly as a caudal spine-like process between the cercal bases

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(Fig. 3.15), whereas, in most groups, the apex of tergum 10 is evenly rounded (Fig. 3.16). Among capniid males, the relative length and shape of this developing epiproct is used to assist in separation of larval species (Stark & Lacey, 2005). Cercal and tergal setation patterns are also useful characters for recognition of selected genera and species. Known species of genus Alloperla, for example, have a vertical fringe of fine cercal setae not found in most other chloroperlid genera (Fig. 3.39), and species of genus Allocapnia are distinguished, in some cases, by the distribution of tergal setae observed in lateral aspect. The key presented below is based on characters found primarily in late instars, and in some cases, preemergent larvae. The latter situation applies especially for species of Allocapnia and certain Taeniopteryx where developing adult features in pharate males are required. In both these genera, and in stoneflies in general, female specimens are recognized by presence of a small notch on abdominal sternum 8. Developing males of Allocapnia, Taeniopteryx, and other genera with epiprocts may be recognized by a projection on tergum 10 to accommodate the developing epiproct. Males with or without epiprocts may be recognized by absence of the notch on the 8th abdominal sternum.

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Figure 3.1, Eccoptura xanthenes (Perlidae), habitus, dorsal, with structures labeled. © Stewart & Stark (2002).

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Figure 3.2, Eccoptura xanthenes (Perlidae), habitus, ventral, with structures labeled. © Stewart & Stark (2002).

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Fig. 3.3

Fig. 3.4

Fig. 3.5

Fig. 3.6

Figures 3.3–3.6, Plecoptera famm., habiti, dorsal. 3.3, Paracapnia angulata (Capniidae). 3.4, Leuctra sibleyi (Leuctridae). 3.5, Prostoia completa (Nemouridae). 3.6, Taeniopteryx maura (Taeniopterygidae). © Stewart & Stark (2002). Scale lines = 2 mm.

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Fig. 3.7

Fig. 3.8

Fig. 3.9

Fig. 3.10

Figures 3.7–3.10, Plecoptera famm., habiti, dorsal. 3.7, Alloperla imbecilla (Chloroperlidae). 3.8, Tallaperla maria (Peltoperlidae). 3.9, Malirekus hastatus (Perlodidae). 3.10. Pteronarcys dorsata (Pteronarcyidae). © Stewart & Stark (2002). Scale lines = 2 mm.

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Table 3.1. Systematic list of the stoneflies (Order Plecoptera) known from the southeastern United States (AL, FL, GA, KY, MS, NC, SC, TN). List modified from Stewart & Stark (2002), with unpublished records from North Carolina and Georgia provided by S. Beaty, D. Lenat, and B. Kondratieff. # = larval species described and included in the accompanying key, * = species occurring in one or more southeastern states, + = species known from states adjacent to the southeastern region. Species names are followed by the name of the original author(s) and the date of publication. Where the author(s) and date of publication are enclosed in parentheses, the species name has been transferred to a genus different from the one in which it was originally described: the original genus name is indicated following the author and date. Two-letter postal codes are used to indicate state and provincial distributions in the United States and Canada. Indented species names are considered synonyms of the immediately preceding unindented species name; the author and date of publication of the synonymy is indicated in parenthesis for each synonym. EPA Region IV state codes are in bold.

Capniidae

# * Allocapnia aurora Ricker, 1952—AL, DC, GA, MD, MS, NC, PA, SC, TN, VA. * Allocapnia brooksi Ross, 1964—TN. * Allocapnia cunninghami Ross & Ricker, 1971—KY, TN. * Allocapnia curiosa Frison, 1942—KY, MD, NY, PA, VA, WV. * Allocapnia forbesi Frison, 1929—IL, IN, KY, OH, TN, WV. * Allocapnia frisoni Ross & Ricker, 1964— KY, NY, OH, PA, TN, VA, WI, WV. + Allocapnia frumi Kirchner, 1982—WV. * Allocapnia fumosa Ross, 1964—NC, TN, VA. # * Allocapnia granulata (Claassen, 1924), Capnella—AL, AR, DC, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN,MO, MS, NJ, NY, OH, OK, ON, PA, QC, TN, TX, VA, WI, WV. + Allocapnia harperi Kirchner, 1980—PA, VA, WV. # + Allocapnia illinoensis Frison, 1935—IL, IN, ME, MN, NY, OH, ON, QC, VA, WI, WV. * Allocapnia indianae Ricker, 1952—IN, KY, NY, OH. * Allocapnia loshada Ricker, 1952—TN, VA, WV. + Allocapnia maria Hanson, 1942—CT, MA, MD, ME, NB, NH, NS, NY, PA, QC, VA, VT, WV. * Allocapnia menawa Grubbs & Sheldon, 2008—AL.

Capniidae continued

* Allocapnia muskogee Grubbs & Sheldon, 2008—AL. # * Allocapnia mystica Frison, 1929—AL, AR, GA, IL, IN, KY, MO, MS, OH, TN, VA, WV. # * Allocapnia nivicola (Fitch, 1847), Perla— AL, CT, DC, DE, IL, IN, KY, MA, MD, NB, NC, NJ, NS, NY, OH, PA, QC, RI, TN, VA, VT, WI, WV. * Allocapnia ohioensis Ross & Ricker, 1964—IN, KY, NY, OH, WV. * Allocapnia perplexa Ross & Ricker, 1971— TN. * Allocapnia polemistis Ross & Ricker, 1971—AL, MS. # * Allocapnia pygmaea (Burmeister, 1839), Semblis—CT, DC, IA, KY, MA, MD, ME, MI, MN, MO, NB, ND, NH, NS, NY, OH, ON, PA, QC, RI, TN, VA, VT, WI, WV. Allocapnia torotonensis Ricker, 1935 (Frison, 1942). # * Allocapnia recta (Claassen, 1924), Capnella—AL, CT, DC, DE, GA, IL, IN, KY, MA, MD, ME, NC, NH, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. # * Allocapnia rickeri Frison, 1929—AL, AR, DC, DE, GA, IA, IN, KS, KY, MD, MN, MO, MS, NC, NJ, NY, OH, OK, ON, PA, TN, VA, WI, WV. * Allocapnia sano Grubbs, 2006—AL. * Allocapnia sequatchie Kondratieff & Kirchner, 2000—TN.

Chapter 3 ~ Key to Stoneflies

Capniidae continued

* Allocapnia smithi Ross & Ricker, 1971— AL, IL, IN, KY, OH. * Allocapnia stannardi Ross, 1964—NC, TN, VA. + Allocapnia simmonsi Kondratieff & Voshell, 1980—PA, VA. # * Allocapnia starki Kondratieff & Kirchner, 2000—FL, LA, MS. * Allocapnia tennessa Ross & Ricker, 1964— AL, TN. * Allocapnia tsalagi Grubbs, 2008—AL. * Allocapnia unzickeri Ross & Yamamoto, 1966—GA, TN. # * Allocapnia virginiana Frison, 1942—AL, DE, GA, LA, MS, NC, SC, VA. # * Allocapnia vivipara (Claassen, 1924), Capnella—AR, DC, IA, IL, IN, KS, KY, MD, MI, MN, MO, NE, NY, OH, OK, ON, PA, QC, TN, VA, WI, WV. * Allocapnia wrayi Ross, 1964—DC, DE, GA, MD, NC, PA, SC, TN, VA. * Allocapnia zola Ricker, 1952—CT, KY, ME, NB, NY, OH, PA, TN, VA, WV. # * Nemocapnia carolina Banks, 1938—AL, AR, FL, IL, IN, MS, NC, QC, SC, VA. # * Paracapnia angulata Hanson, 1942—AR, CO, CT, DE, IL, KY, MA, MB, MD, ME, MI, MO, NC, NH, NL, NY, OH, OK, ON, PA, QC, SD, SK, TN, VA, WI, WV, WY.

Leuctridae

* Leuctra alabama James, 1974—AL. * Leuctra alexanderi Hanson, 1941—KY, NC, PA, TN, VA, WV. * Leuctra alta James, 1974—AL, IL, IN, KY. * Leuctra biloba Claassen, 1923—AL, GA, NC, ON, TN, VA. * Leuctra carolinensis Claassen, 1923—MD, MS, NC, NJ, TN, VA. # * Leuctra colemanorum Harrison & Stark, 2010—MS.

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Leuctridae continued

* Leuctra cottaquilla James, 1974—AL, FL. * Leuctra crossi James, 1976—AL. + Leuctra duplicata Claassen, 1923—CT, MD, ME, NB, NS, NY, ON, PA, PE, QC, VA, WV. # * Leuctra ferruginea (Walker, 1852), Nemoura—AL, CT, DE, FL, KY, LA, MA, MD, ME, MI, MN, MS, NB, NC, NJ, NL, NS, NY, OH, PA, QC, SC, SK, VA, WI, WV. Leuctra decepta Claassen, 1923 (Ricker, i.l.). * Leuctra grandis Banks, 1906—MA, ME, NB, NC, PA, WV. * Leuctra hicksi Harrison & Stark, 2010— MS. + Leuctra maria Hanson, 1941—CT, ME, NH, ON, PA, QC, VT, WV. * Leuctra mitchellensis Hanson, 1941— NC, VA. * Leuctra moha Ricker, 1952—GA, SC. * Leuctra monticola Hanson, 1941—NC, TN, VA. * Leuctra nephophila Hanson, 1941— NC, TN. * Leuctra pinhoti Grubbs & Sheldon, 2009— AL. * Leuctra rickeri James, 1976—AL, IA, IL, IN, KY, MI, MS, OH, WV. # * Leuctra sibleyi Claassen, 1923—CT, DE, IL, IN, KY, MA, MD, ME, NB, NC, NY, OH, ON, PA, QC, TN, VA, WI, WV. # * Leuctra tenella Provancher, 1878—CT, MA, MD, ME, MN, MS, NB, NJ, NL, NS, NY, ON, PA, QC, WI, WV. Leuctra hamula Claassen, 1923 (Ricker, 1952). # * Leuctra tenuis (Pictet, 1841), Nemoura— AL, AR, CT, DE, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, MS, NB, NC, NJ, NS, NY, OH, OK, PA, QC, VA, WI, WV. * Leuctra triloba Claassen, 1923—AL, FL, NC, NY, QC, SC, VA, WV. Leuctra crosbyi Claassen, 1937 (Ricker i.l.).

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Leuctridae continued

# + Leuctra truncata Claassen, 1923—CT, ME, NL, NY, PA, QC, VA, WV. * Leuctra usdi Grubbs, 2010—AL, TN. * Leuctra variabilis Hanson, 1941—MA, MD, ME, NC, NH, PA, VA, VT. # + Megaleuctra flinti Baumann, 1973—MD, PA, VA, WV. # * Megaleuctra williamsae Hanson, 1941— NC, SC, TN, VA.

* Paraleuctra sara (Claassen, 1937), Leuctra—AL, CT, DE, KY, MA, MD, ME, NB, NC, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, WV. # * Zealeuctra claasseni (Frison, 1929), Leuctra—AL, AR, IL, IN, KS, KY, MO, OK, TN, TX, WV. * Zealeuctra fraxina Ricker & Ross, 1969— IL, IN, KY, OH, PA, TN, WV. * Zealeuctra talladega Grubbs, 2005—AL. * Zealeuctra ukayodi Grubbs, 2013 in Grubbs et al., 2013—AL, TN.

Nemouridae

# * Amphinemura alabama Baumann, 1996— AL, KY, MS. * Amphinemura appalachia Baumann, 1996—GA, NC, PA, SC, TN, VA. # * Amphinemura delosa (Ricker, 1952), Nemoura (Amphinemura)—AL, AR, GA, IA, IL, IN, KS, KY, MI, MO, NC, OH, OK, ON, PA, QC, TN, VA, WI, WV. # * Amphinemura mockfordi (Ricker, 1952), Nemoura (Amphinemura)—TN. # * Amphinemura nigritta (Provancher, 1876), Nemoura—AL, AR, CT, DE, FL, GA, IL, IN, KY, MD, ME, MI, MO, MS, NB, NC, NL, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, WV. Nemoura styllata Banks, 1920 (Ricker, 1952); Nemoura venosa Banks, 1897 (Ricker, 1952).

Nemouridae continued

* Amphinemura varshava (Ricker, 1952), Nemoura (Amphinemura)—IA, IL, IN, KY, WI. # * Amphinemura wui (Claassen, 1936), Nemoura—CT, DE, GA, KY, MA, MD, ME, NB, NC, NJ, NS, NY, PA, PE, QC, SC, TN, VA, WV. # * Ostrocerca albidipennis (Walker, 1852), Nemoura—CT, MA, MD, ME, MI, NC, NH, NS, NY, ON, PA, QC, VA, WV. Nemoura serrata Claassen, 1923 (Ricker, 1952). + Ostrocerca complexa (Claassen, 1937), Nemoura—CT, MA, ME, NB, NS, NY, ON, PA, QC, VA, WV. + Ostrocerca prolongata (Claassen, 1923), Nemoura—DE, ME, NB, NH, NY, PA, QC, VA, WV. # * Ostrocerca truncata (Claassen, 1923), Nemoura—CT, IN, KY, MA, MD, ME, NC, NY, OH, ON, PA, QC, VA, WV. # * Paranemoura perfecta (Walker, 1852), Nemoura—CT, MA, MD, ME, MI, NB, NC, NH, NS, NY, ON, PA, PE, QC, TN, VA, VT, WV. Nemoura punctipennis Claassen, 1923 (Ricker, 1938) # * Prostoia completa (Walker, 1852), Nemoura—AL, DE, IA, IN, KY, MA, MD, ME, MI, MN, MS, NB, NC, NL, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, WI, WV. Nemoura glabra Claassen, 1923 (Ricker, 1938). # * Prostoia hallasi Kondratieff & Kirchner, 1984—CT, GA, IL, MA, NC, VA. # * Prostoia similis (Hagen, 1861), Taeniopteryx—CT, DE, IL, IN, KY, MA, MD, ME, MI, MN, MO, NY, OH, ON, PA, QC, SC, VA, WI, WV. Nemoura divergens Claassen, 1923 (Illies, 1966).

Chapter 3 ~ Key to Stoneflies

Nemouridae continued # * Shipsa

rotunda (Claassen, 1923), Nemoura—AB, AK, AL, AR, IL, MB, MD, ME, MI, MN, MS, NB, NC, NT, ON, QC, SC, SK, VA, WI.

* Soyedina alexandria Grubbs, 2006—AL, MS, TN. * Soyedina calcarea Grubbs, 2006—KY. # * Soyedina carolinensis (Claassen, 1923), Nemoura—DE, MD, NC, PA, TN, VA, WV. * Soyedina kondratieffi Baumann & Grubbs, 1996—NC, TN. # * Soyedina vallicularia (Wu, 1923), Nemoura—CT, IA, IL, IN, KY, MA, MD, ME, MI, NS, NY, OH, ON, PA, QC, TN, VA, WI, WV. * Soyedina washingtoni (Claassen, 1923), Nemoura—CT, ME, NB, NC, NH, PA, WV. # * Zapada chila (Ricker, 1952), Nemoura (Zapada)—TN. * Zapada species A Baumann & Grubbs, 2015-NC, TN, VA.

Taeniopterygidae

# * Bolotoperla rossi (Frison, 1942), Brachyptera—MA, ME, NC, NH, NY, PA, QC, SC, TN, VA, WV. # * Oemopteryx contorta (Needham & Claassen, 1925), Taeniopteryx— CT, KY, MA, MD, ME, NC, NH, PA, TN, VA, WV. + Oemopteryx glacialis (Newport, 1848), Nemoura (Brachyptera)—CT, MI, MN, NY, ON, QC, WI, WV. Perla chicoutimiensis Provancher, 1878 (Ricker, 1952); Taeniopteryx (Oemopteryx) alex Hanson, 1938 (Frison, 1942). # * Strophopteryx appalachia Ricker & Ross, 1975—KY, NC, NY, PA, SC, TN, VA, WV.

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Taeniopterygidae continued

# * Strophopteryx fasciata (Burmeister, 1839), Semblis—AL, AR, CT, DE, IA, IL, IN, KS, KY, MB, ME, MI, MN, MO, MS, NC, ND, OH, OK, ON, PA, QC, SC, VA, WI, WV. Taeniopteryx frigida Hagen, 1861 (Needham & Claassen, 1925). # * Strophopteryx limata (Frison, 1942), Brachyptera—NC, TN, VA. # * Taenionema atlanticum Ricker & Ross, 1975—CT, KY, MA, MD, ME, NB, NC, NH, NL, NS, NY, OH, PA, QC, SC, TN, VA, VT, WV.

# * Taeniopteryx burksi Ricker & Ross, 1968— AL, AR, CO, CT, DE, FL, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, MS, NC, NE, NY, OH, OK, ON, PA, QC, SD, TN, TX, VA, WI, WV. # * Taeniopteryx lita Frison, 1942—AL, AR, FL, IL, IN, KY, LA, MS, NC, NJ, OK, SC, TX, VA, WV. # * Taeniopteryx lonicera Ricker & Ross, 1968—AL, AR, FL, GA, LA, MD, MS, NC, SC, TN, TX, VA. # * Taeniopteryx maura (Pictet, 1841), Nemoura (Taeniopteryx)—AL, AR, CT, DE, GA, IN, KY, MA, MD, ME, MI, MN, MS, NC, NY, OH, PA, SC, TN, TX, VA, WV. # * Taeniopteryx metequi Ricker & Ross, 1968—AL, AR, IL, IN, KS, KY, MO, NC, NY, OH, OK, ON, PA, VA, WV. # * Taeniopteryx nelsoni Kondratieff & Kirchner, 1982—NC, VA. # * Taeniopteryx parvula Banks, 1918—AB, AR, CO, CT, GA, IL, IN, KY, MB, ME, MI, MN, MO, MS, NC, NM, NY, OH, ON, PA, QC, SC, TN, VA, WI, WV, WY. Taeniopteryx pecos Baumann & Jacobi, 1984 (Kondratieff & Baumann, 1988).

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Taeniopterygidae continued

# * Taeniopteryx robinae Kondratieff & Kirchner, 1984—SC. # * Taeniopteryx ugola Ricker & Ross, 1968— AL, GA, KY, PA, TN, VA.

Chloroperlidae

+ Alloperla aracoma Harper & Kirchner, 1978—PA, VA, WV. # * Alloperla atlantica Baumann, 1974—AL, CT, GA, MA, MD, ME, MI, MN, NB, NC, NH, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, VT. + Alloperla banksi Frison, 1942—ME, MI, NB, NS, NY, ON, QC, VA. + Alloperla biserrata Nelson & Kondratieff, 1980—MD, PA, VA, WV. * Alloperla caudata Frison, 1934—AL, AR, IL, KY, MO, MS, OK. * Alloperla chloris Frison, 1934—CT, GA, KY, MA, MD, ME, NB, NC, NH, NS, NY, OH, PA, QC, SC, TN, VA, WV. Chloroperla milnei Ricker, 1935 (Frison, 1942). # + Alloperla concolor Ricker, 1935—CT, MA, ME, NB, NH, NL, NS, NY, ON, PA, QC, VT, WV. # * Alloperla furcula Surdick, 1981—SC. * Alloperla hamata Surdick, 1981—AL, AR, IN, KY, MO. * Alloperla idei (Ricker, 1935), Chloroperla—AL, GA, KY, ME, OH, ON, QC, VA, WV. # * Alloperla imbecilla (Say, 1823), Sialis— KY, MD, NY, OH, PA, VA, WV. * Alloperla lenati Kondratieff & Kirchner, 2004—NC. * Alloperla nanina Banks, 1911—GA, NC, NY, TN, VA. Alloperla lodgei Frison, 1935 (Frison, 1942) # * Alloperla natchez Surdick & Stark, 1980— MS. # * Alloperla neglecta Frison, 1935—NC, TN, VA.

Chloroperlidae continued

# * Alloperla petasata Surdick, 2004—GA, MA, ME, NB, NC, NH, NL, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WV. # * Alloperla prognoides Surdick & Stark, 2004 in Surdick, 2004—AL, FL. + Alloperla stipitata Surdick, 2004—VA. # * Alloperla usa Ricker, 1952—AL, GA, KY, MD, NC, ON, PA, SC, TN, VA, WV.

# * Haploperla brevis (Banks, 1895), Chloroperla—AL, AR, BC, CT, DE, GA, IL, IN, KY, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NH, NJ, NS, NY, OH, OK, ON, PA, PE, QC, SC, SK, TN, VA, VT, WI, WV. Isopteryx oculata Klapálek, 1923 (Ricker, 1938); Hastaperla calcarea Ricker, 1935 (Frison, 1937). # * Haploperla chukcho (Surdick & Stark, 1980), Hastaperla—MS. * Haploperla fleeki Kondratieff, Kirchner & Lenat, 2005—NC. * Haploperla parkeri Kirchner & Kondratieff, 2005 in Kondratieff et al., 2005—NC. # * Rasvena terna (Frison, 1942), Chloroperla—ME, NC, NH, NY, PA, TN, VA, VT, WV. # * Suwallia marginata (Banks, 1897), Chloroperla—GA, MA, MD, ME, MI, NB, NC, NH, NL, NY, ON, PA, QC, TN, VA, VT, WI, WV. # * Sweltsa hoffmani Kondratieff & Kirchner, 2009—AL, IN, KY, NY, OH, PA, TN, WV. * Sweltsa holstonensis Kondratieff & Kirchner, 1998—NC, VA. # * Sweltsa lateralis (Banks, 1911), Alloperla— CT, GA, MA, MD, ME, NB, NC, NH, NY, OH, PA, QC, SC, TN, VA, VT, WV.

Chapter 3 ~ Key to Stoneflies

Chloroperlidae continued

# * Sweltsa mediana (Banks, 1911), Alloperla—AL, NC, SC, TN, VA. # + Sweltsa naica (Provancher, 1876), Perla— ME, NB, NH, NL, NS, NY, PA, PE, QC, VA, VT, WV. Alloperla novascotiana Needham & Claassen, 1925 (Ricker, 1952). # * Sweltsa onkos (Ricker, 1935), Alloperla— CT, DE, KY, MD, ME, NB, NH, NL, NS, NY, OH, ON, PA, PE, QC, VA, VT, WV. # + Sweltsa palearata Surdick, 2004—MD, PA, VA, WV. # + Sweltsa pocahontas Kirchner & Kondratieff, 1988—MD, WV. # * Sweltsa urticae (Ricker, 1952), Alloperla (Sweltsa)—NC, VA. * Sweltsa voshelli Kondratieff & Kirchner, 1991—GA, NC, VA.

Peltoperlidae

* Peltoperla arcuata Needham, 1905—KY, NY, OH, PA, QC, TN, VA, WV. * Peltoperla tarteri Stark & Kondratieff, 1987—NC, VA, WV.

* Tallaperla anna (Needham & Smith, 1916), Peltoperla—GA, NC, SC, VA. * Tallaperla cornelia (Needham & Smith, 1916), Peltoperla—AL, FL, GA, NC, SC. * Tallaperla elisa Stark, 1983—NC, TN. * Tallaperla laurie (Ricker, 1952), Peltoperla—AL, GA, NC, SC, TN. + Tallaperla lobata Stark, 1983—VA. * Tallaperla maiyae Kondratieff, Kirchner & Zuellig, 2007—NC. * Tallaperla maria (Needham & Smith, 1916), Peltoperla—AL, CT, DE, GA, KY, MA, MD, ME, NC, NH, NY, PA, SC, TN, VA, WV. Peltoperla dorothea Needham & Smith, 1916 (Ricker, 1952). #*Viehoperla ada (Needham & Smith, 1916), Peltoperla—GA, NC, SC, TN. Peltoperla zipha Frison, 1942 (Stark & Stewart, 1981).

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Perlidae

# * Acroneuria abnormis (Newman, 1838), Perla—AB, AL, CO, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MS, MT, NB, NC, ND, NE, NL, NM, NY, OH, ON, PA, QC, SC, SD, SK, TN, UT, VA, WI, WV, WY. Perla sonans Newport, 1851 (Hagen, 1861); Acroneuria eidmanni Samal, 1933 (Ricker, 1938); Acroneuria depressa Needham & Claassen, 1922 (Stark & Gaufin, 1976). # * Acroneuria arenosa (Pictet, 1841), Perla— AL, CT, DC, DE, FL, GA, LA, MA, MD, ME, MS, NC, NJ, NY, PA, QC, SC, TX, VA, WV. Perla pennsylvanica Rambur, 1842 (Ricker, 1938); Perla trijuncta Walker, 1852 (Ricker, 1938). * Acroneuria arida (Hagen, 1861), Perla— GA, NC, NJ, PA, TN. Perla valida Banks, 1906 (Needham & Claassen, 1925). # * Acroneuria carolinensis (Banks, 1905), Perla—AL, CT, GA, KY, MA, MB, MD, ME, MS, NC, NH, NJ, NY, OH, ON, PA, QC, SC, TN, VA, WV. Acroneuria cuestae Ricker, 1935 (Frison, 1942). * Acroneuria covelli Grubbs & Stark, 2004— IN, KY, TN. # * Acroneuria evoluta Klapálek, 1909—AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, NC, OH, OK, PA, TN, TX. Acroneuria prolonga Claassen, 1937 (Stark & Brown, 1991); Acroneuria mela Frison, 1942 (Stark & Brown, 1991). # * Acroneuria filicis Frison, 1942—AL, AR, GA, IL, IN, KY, MD, MO, NC, OH, PA, SC, TN, VA, WV. + Acroneuria flinti Stark & Gaufin, 1976— VA.

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Perlidae continued

# * Acroneuria frisoni Stark & Brown, 1991— AL, AR, IL, IN, KS, KY, MA, MD, MI, MO, NC, OH, OK, ON, PA, TN, VA, WV. # * Acroneuria hitchcocki Kondratieff & Kirchner, 1988—KY. # * Acroneuria internata (Walker, 1852), Perla—AR, IL, IN, KY, MI, MN, MO, OK, VA, WI, WV. # * Acroneuria kirchneri Stark & Kondratieff, 2004—KY, PA, VA, WV. + Acroneuria kosztarabi Kondratieff & Kirchner, 1993—VA. # * Acroneuria lycorias (Newman, 1839), Perla—CT, KY, IA, MB, ME, MI, MN, NB, NC, ND, NJ, NS, NY, OH, ON, PA, QC, SK, TN, VA, WI, WV. Perla navalis Provancher, 1876 (Klapálek, 1909); Perla riparia Provancher, 1876 (Klapálek, 1917); Perla excavata Banks, 1908 (Klapálek, 1917); Acroneuria perbranchiata Neave, 1933 (Frison, 1942). # * Acroneuria perplexa Frison, 1937—AL, AR, DC, GA, IL, IN, KY, MO, OH, OK, PA, TN, WV. * Acroneuria petersi Stark & Gaufin, 1976— AL, GA, TN. + Acroneuria yuchi Stark & Kondratieff, 2004—VA. # * Agnetina annulipes (Hagen, 1861), Perla— AL, DC, FL, IN, LA, MD, MS, NC, PA, SC, VA, WV. # * Agnetina capitata (Pictet, 1841), Perla— AR, CT, IA, KY, MA, MD, ME, MI, MN, MO, NB, NC, NH, NS, NY, OH, ON, PA, PE, QC, VA, WI, WV. Perla tristis Hagen, 1861 (Klapálek, 1923); Perla hieroglyphica Provancher, 1876 (Needham & Claassen, 1925);

Perlidae continued

Perla marginata Provancher, 1876 (Klapálek, 1923); Perla americana Banks, 1900 (Klapálek, 1923); Perla illustris Banks, 1908 (Needham & Claassen, 1925); Perla inota Banks, 1918 (Needham & Claassen, 1925); Harrisiola nigriscens Banks, 1948 (Illies, 1966); Harrisiola abbreviata Banks, 1948 (Hitchcock, 1974); Harrisiola modesta Banks, 1948 (Illies, 1966). # * Agnetina flavescens (Walsh, 1862), Perla— AL, AR, GA, IL, IN, KY, MD, MI, MN, MO, NC, NY, OH, OK, PA, SC, TN, VA, WV. Harrisiola klapaleki Banks, 1948 (Stark, 1986). # * Attaneuria ruralis (Hagen, 1861), Perla— AL, AR, DC, FL, GA, IA, IL, IN, KS, MB, MD, MI, MN, MO, MS, NC, NE, NY, OH, PA, SC, TN, VA, WI. Perla rupinsulensis Walsh, 1862 (Needham & Claassen, 1922); Perla quebecensis Provancher, 1876 (Needham & Claassen, 1922); Perla attenuata Banks, 1905 (Needham & Claassen, 1922). # * Beloneuria georgiana (Banks, 1914), Perla—GA, NC. # * Beloneuria jamesae Stark & Szczytko, 1976—AL. # * Beloneuria stewarti Stark & Szczytko, 1976—GA, NC, SC, TN. # * Eccoptura xanthenes (Newman, 1838), Perla—AL, CT, DE, FL, GA, KY, MD, MS, NC, NY, OH, PA, SC, TN, VA, WV. Acroneuria brevicauda Klapálek, 1909 (Needham & Claassen, 1922).

Chapter 3 ~ Key to Stoneflies

Perlidae continued

# * Hansonoperla appalachia Nelson, 1979— MA, NC, NH, PA, SC, TN, VA, WV. # * Hansonoperla cheaha Kondratieff & Kirchner, 1996—AL. * Hansonoperla hokolesqua Kondratieff & Kirchner, 1996—KY, WV.

* Neoperla carlsoni Stark & Baumann, 1978—AL, AR, FL, LA, MO, MS, SC, TX, VA. # * Neoperla catharae Stark & Baumann, 1978—AL, AR, IL, IN, KY, MI, MO, NC, OH, OK, PA, TN, VA. # * Neoperla choctaw Stark & Baumann, 1978—AR, KY, MO, OK, PA, WV. # * Neoperla clymene (Newman, 1839), Chloroperla—AL, FL, GA, IA, IL, IN, KY, LA, MS, NC, OK, PA, SC, TX, VA, WV. * Neoperla coosa Smith & Stark, 1998—AL, NC, TN. # * Neoperla coxi Stark, 1995—MS. * Neoperla gaufini Stark & Baumann, 1978—IN, KY, OH. * Neoperla harrisi Stark & Lentz, 1988— AL, TN. * Neoperla occipitalis (Pictet, 1841), Perla— AL, IL, IN, KY, ME, MS, NC, NS, NY, OH, ON, PA, SC, TN, VA, WI. Neoperla freytagi Stark & Baumann, 1978 (Stark, 1990). # * Neoperla robisoni Poulton & Stewart, 1986—AR, IL, MO, MS, OK, PA, TN, WI. * Neoperla stewarti Stark & Baumann, 1978—AL, IL, IN, KY, MA, MD, ME, MI, MN, MS, NC, OH, PA, TN, VA, WI, WV. # * Paragnetina fumosa (Banks, 1902), Perla—AL, DC, FL, GA, LA, MS, NC, PA, SC, TX, VA. # * Paragnetina ichusa Stark & Szczytko, 1981—NC, SC, TN, VA.

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Perlidae continued

# * Paragnetina immarginata (Say, 1823), Sialis—CT, DE, GA, KY, MA, ME, NC, NH, NY, PA, QC, SC, TN, VA, WV. Perla lurida Hagen, 1861 (Ricker, 1949); Acroneuria fumosa Ricker, 1935 (Frison, 1937). # * Paragnetina kansensis (Banks, 1902), Perla—AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, NC, SC. Paragnetina fattigi Ricker, 1949 (Stark & Szczytko, 1981). # * Paragnetina media (Walker, 1852), Perla— AR, CT, DE, IA, IL, IN, KY, MB, MD, ME, MI, MN, MO, NB, NH, NY, OH, ON, PA, QC, SK, VA, WI, WV. Acroneuria salvelini Ricker, 1935 (Frison, 1937). # * Perlesta adena Stark, 1989—IN, KY, OH, TN. * Perlesta beatyi Kondratieff, Zuellig & Lenat, 2011—NC. * Perlesta bjostadi Kondratieff & Lenat, 2006—NC. # * Perlesta cinctipes (Banks, 1905), Perlinella—AR, IA, IL, KS, KY, MO, NE, OH, OK, WV. + Perlesta cranshawi Kondratieff & Kirchner, 2006—VA. # * Perlesta decipiens (Walsh, 1862), Perla— AR, CO, IA, IL, IN, KY, MI, MO, NC, ND, NE, NM, OH, OK, PA, SD, TX, VA, WI, WV, WY. Perla brunnipennis Walsh, 1862 (Stark, 1989); Isoperla texana Banks, 1914 (Stark, 1989); Perlesta costalis Klapálek, 1921 (Stark, 1989). * Perlesta durfeei Kondratieff, Zuellig & Kirchner, 2008-NC, VA. * Perlesta etnieri Kondratieff & Kirchner, 2002—TN.

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Perlidae continued

# * Perlesta frisoni Banks, 1948—NC, SC, TN, VA, WV. * Perlesta georgiae Kondratieff, Zuellig & Lenat, 2008-NC. * Perlesta lagoi Stark, 1989—IA, IL, KY, MI, MS, TN. * Perlesta leathermani Kondratieff & Zuellig, 2006—NC. * Perlesta nelsoni Stark, 1989—NC, PA, SC, TN, WV. * Perlesta nitida Banks, 1948—CT, IN, KY, MA, OH, PA. * Perlesta placida (Hagen, 1861), Perla— AL, DC, FL, GA, LA, MD, ME, MS, NC, PA, TX, VA. Chloroperla virginica Banks, 1898 (Needham & Claassen, 1925); Perlesta virginica immaculata Klapálek, 1921 (Needham & Claassen, 1925). * Perlesta puttmanni Kondratieff & Kirchner, 2003—NC, VA. * Perlesta roblei Kondratieff & Kirchner, 2003—NC, VA. * Perlesta shawnee Grubbs, 2005—AL, IL, IN, NC, VA. # * Perlesta shubuta Stark, 1989—AR, IA, IL, KY, MI, MO, MS, OK. # * Perlesta teaysia Kirchner & Kondratieff, 1997—IL, IN, KY, MD, OH, PA, TN, VA, WV.

# * Perlinella drymo (Newman, 1839), Isogenus—AL, AR, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK, PA, QC, SC, TN, TX, VA, WI, WV. Perla elongata Walsh, 1862 (Needham & Claassen, 1925); Perla trivittata Banks, 1895 (Needham & Claassen, 1925).

Perlidae continued

# * Perlinella ephyre (Newman, 1839), Chloroperla—AL, AR, CT, FL, GA, IA, IN, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NJ, NY, OH, OK, PA, RI, SC, TN, VA, WI, WV. Perla producta Walsh, 1962 (Needham & Claassen, 1925); Perla fumipennis Walsh, 1862 (Kondratieff et al., 1988); Atoperla consors Banks, 1948 (Illies, 1966). # * Perlinella zwicki Kondratieff, Kirchner & Stewart, 1988—AL, FL, MS, SC, NC.

Perlodidae

# * Clioperla clio (Newman, 1839), Isogenus— AL, AR, CT, DE, FL, GA, IA, IL, IN, KY, MA, MD, MI, MO, MS, NC, OH, OK, ON, PA, SC, TN, VA, WI, WV. Isoperla confusa Frison, 1935 (Ricker, 1952). # * Cultus decisus isolatus (Banks, 1920), Isoperla—GA, NC, VA. # * Cultus verticalis (Banks, 1920), Perla— NC, NH, PA, QC, TN, VA, WV. # * Diploperla duplicata (Banks, 1920), Perla—AL, DE, GA, MD, MS, NC, PA, SC, TN, VA, WV. # * Diploperla kanawholensis Kirchner & Kondratieff, 1984—KY, NC, VA, WV. # * Diploperla morgani Kondratieff & Voshell, 1979—NC, VA. # * Diploperla robusta Stark & Gaufin, 1974— AL, CT, IN, KY, OH, PA, TN, VA, WV. # * Helopicus bogaloosa Stark & Ray, 1983— AL, FL, GA, LA, MS, NC, SC. # * Helopicus subvarians (Banks, 1920), Perla—AL, CT, FL, KY, ME, NC, ON, PA, QC, SC, TN, VA, WV.

Chapter 3 ~ Key to Stoneflies

Perlodidae continued

# * Hydroperla fugitans (Needham & Claassen, 1925), Perla—AR, IA, IL, IN, KS, MO, MS, TN, TX. Hydroperla harti Frison, 1935 (Ricker, 1952). # * Hydroperla phormidia Ray & Stark, 1981—FL, NC, SC. # * Hydroperla rickeri (Stark, 1984), Helopicus—TN. # * Isogenoides hansoni (Ricker, 1952), Isogenus (Isogenoides)—CT, MA, MD, ME, NB, NC, NS, NY, PA, QC, VA, WV. # * Isogenoides varians (Walsh, 1862), Perla— IA, IL, IN, KS, MI, MN, MS, NC, SC, TN, VA. * Isoperla bellona Banks, 1911—NC, SC. # * Isoperla bilineata (Say, 1823), Sialis—IA, IL, IN, KS, MB, MI, MN, MO, MS, ND, NE, OH, SK, WI. # * Isoperla burksi Frison, 1942—AL, AR, IL, IN, KY, MD, MO, NC, OH, OK, SC, VA, WV. * Isoperla catawba Szczytko & Kondratieff, 2015—SC. * Isoperla cherokee Szczytko & Kondratieff, 2015—NC, TN. * Isoperla chickamauga Szczytko & Kondratieff, 2015—GA. # * Isoperla davisi James, 1974—AL, AR, DE, FL, LA, MS, NC, OK, SC, TX, VA. Isoperla coushatta Szczytko & Stewart, 1976 (Szczytko & Kondratieff, 2015). # * Isoperla decepta Frison, 1935—AL, IL, IN, KY, MI, MO, OH, ON, TN. # * Isoperla dicala Frison, 1942—AL, CT, FL, GA, IA, IN, KY, MA, MB, ME, MI, MN, MO, MS, NB, NC, NY, OH, ON, PA, QC, SC, TN, WI, VA, WV. * Isoperla distincta Nelson, 1976—TN. * Isoperla fauschi Szczytko & Kondratieff, 2015—NC, VA.

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Perlodidae continued

# * Isoperla frisoni Illies, 1966—CT, DE, GA, IN, MB, ME, MI, MN, NB, NC, NS, NY, ON, PA, PE, QC, SC, TN, VA, WI. Isoperla truncata Frison, 1937 (Illies, 1966, Homonym of Navas, 1918). # * Isoperla holochlora (Klapálek, 1923), Chloroperla—AL, CT, DE, KY, MD, ME, NC, NS, NY, OH, PA, QC, SC, TN, VA, WV. * Isoperla jamesae Grubbs & Szczytko, 2010—AL, MS. * Isoperla kircherni Szczytko & Kondratieff, 2015—NC, NY, PA, TN, VA, WV. # * Isoperla lata Frison, 1942—ME, MI, MN, NB, NC, NL, NS, NY, ON, QC, TN, WI, WV. * Isoperla lenati Szczytko & Kondratieff, 2015—FL, GA, NC, SC. # * Isoperla major Nelson, 1983—VA. # * Isoperla marlynia Needham & Claassen, 1925—CO, IA, IL, IN, KY, MB, ME, MI, MN, NB, NE, NJ, NS, NY, ON, PA, QC, VA, WI, WV. * Isoperla montana (Banks, 1898), Chloroperla—CT, DE, IN, KY, MD, ME, MN, NC, NH, NJ, NS, NY, OH, ON, PA, QC, SC, VA, WV. # * Isoperla nana (Walsh, 1962), Chloroperla— IL, IN, KY, MI, NY, OH, ON, PA, QC, WI. Chloroperla minuta Banks, 1900 (Ricker in Illies, 1966). * Isoperla nelsoni Szczytko & Kondratieff, 2015—AL, GA, KY, NC, TN, VA. # * Isoperla orata Frison, 1942—NC, PA, TN, VA. * Isoperla pauli Szczytko & Kondratieff, 2015—NC. * Isoperla poffi Szczytko & Kondratieff, 2015—FL, NC, SC. + Isoperla powhatan Szczytko & Kondratieff, 2015—PA, VA. * Isoperla pseudolata Szczytko & Kondratieff, 2015—NC, VA, WV.

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Perlodidae continued

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Perlodidae continued

* Isoperla pseudosimilis Szczytko & # * Oconoperla innubila (Needham & Claassen, Kondratieff, 2015—CT, MA, ME, 1925), Perla—NC, SC, TN. NC, NH, NY, PA, TN, VA, VT, WV. Oconoperla weaveri Stark & Stewart, * Isoperla reesi Szczytko & Kondratieff, 1982 (Stark, 1985). 2015—NC, VA. * Isoperla sandbergi Szczytko & Kondratieff, # * Remenus bilobatus (Needham & Claassen, 2015—AL. 1925), Perla—AL, CT, DE, KY, # + Isoperla signata (Banks, 1902), Perlinella— MD, NC, NY, PA, SC, TN, VA, WV. AR, CT, IA, MB, ME, MI, MN, MO, # * Remenus duffieldi Nelson & Kondratieff in NB, NS, NY, OH, OK, ON, PA, VA, Kondratieff & Nelson, 1995—GA. WI, WV. # * Remenus kirchneri Kondratieff & Nelson, Pictetia bimaculata Banks, 1948 (Ricker 1995—NC, VA. in Illies, 1966). # + Isoperla similis (Hagen, 1861), Perla—CT, # * Yugus arinus (Frison, 1942), Diploperla— MD, NH, NJ, NY, PA, VA. GA, NC, SC, TN, VA. * Isoperla siouan Szczytko & Kondratieff, # * Yugus bulbosus (Frison, 1942), Diploperla— 2015—NC. GA, NC, TN. # * Isoperla slossonae (Banks, 1911), Perla— # * Yugus kirchneri Nelson, 2001—KY, PA, IA, ME, MI, MN, NB, NC, NH, NS, VA, WV. NY, PA, PE, QC, VA, WI, WV. # * Yugus kondratieffi Nelson, 2001—NC, VA. Clioperla annecta Needham & Claassen, 1925 (Frison, 1942). Pteronarcyidae + Isoperla smithi Szczytko & Kondratieff, # * Pteronarcys biloba Newman, 1838—AL, 2015—VA, WV. CT, GA, MA, MD, ME, NB, NC, * Isoperla starki Szczytko & Kondratieff, NH, NS, NY, PA, PE, QC, SC, 2015—NC. VA, WV. * Isoperla stewarti Szczytko & Kondratieff, Pteronarcys bicarinatus Provancher, 2015—NC, VA. 1876 (Needham & Claassen, 1925). # * Isoperla transmarina (Newman, 1838), # * Pteronarcys comstocki Smith, 1917—KY, Chloroperla—BC, CT, DE, IA, KY, ME, NB, NC, NH, NS, NY, PA, MB, ME, MI, MN, NB, NC, NE, VA, WV. NH, NL, NS, NY, OH, ON, PA, PE, # * Pteronarcys dorsata (Say, 1823), Sialis— QC, SD, SK, WI, WV. AB, AK, AL, BC, CO, FL, GA, IL, Isoperla ventralis Banks, 1908 IN, KS, KY, LA, MB, ME, MI, MN, (Frison, 1942); MS, MT, NB, NC, NL, NT, NY, OH, Isoperla fumosa Neave, 1933 (Ricker, ON, PA, QC, SC, SK, TN, VA, WI, 1946). WV, WY. * Isoperla tutelo Szczytko & Kondratieff, Pteronarcys regalis Newman, 1838 2015—NC. (Smith, 1917); + Isoperla yuchi Szczytko & Kondratieff, Kollaria insignis Pictet, 1841 2015—VA. (Gerstaecker, 1873); * Isoperla zuelligi Szczytko & Kondratieff, Pteronarcys nobilis Hagen, 1861 2015—NC. (Frison, 1942); Pteronarcys frigida Gerstaecker, 1873 # * Malirekus hastatus (Banks, 1920), (Frison, 1942); Isogenus—GA, KY, NC, OH, PA, SC, TN, VA, WV.

Chapter 3 ~ Key to Stoneflies

Pteronarcyidae continued

Pteronarcys rectus Provancher, 1876 (Smith, 1917); Pteronarcys flavicornis Provancher, 1876 (Smith, 1917); Pteronarcys labradoriensis Samal, 1933 (Brink, 1958); Pteronarcys shellfordi Frison, 1934 (Ricker, 1938). # * Pteronarcys pictetii Hagen, 1873—AR, CT, IL, IN, KS, KY, MB, MI, MN, MO, ND, NE, OH, PA, TN, WI. # * Pteronarcys proteus Newman, 1838—KY, MA, MD, ME, NC, NJ, NH, NY, ON, PA, QC, VA, VT, WV. # * Pteronarcys scotti Ricker, 1952—GA, NC, PA, SC, TN, VA.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

KEY TO PLECOPTERA LARVAE

(Region IV state codes are placed in bold for species distributions)

FAMILIES

(After Stewart & Stark, 2002, 2008) 1 1’



Highly branched filamentous gills present around leg bases of all thoracic segments (Fig. 3.11) . .......................................................................................................................... 2 Gills present or absent; if present around leg bases, not consisting of highly branched filamentous type (Fig. 3.12) ................................................................................................ 3

2(1)

Some gills originating from abdominal sterna 1-2 (Fig. 3.13); abdomen circular in cross section and terminating in an apical spine-like structure (Fig. 3.15); most species with lateral knob-like projections on abdominal segments (Fig. 3.15) . .................................. ........................................................................... PTERONARCYIDAE, Pteronarcys, 154 2’ Gills absent from abdominal sterna but may be present on paraprocts (Fig. 3.14); abdomen flattened and without apical spine-like structure (Fig. 3.16); lateral knob-like projections absent ......................................................................................................... PERLIDAE, 27 3(1’) 3’

Each coxa bearing a single, segmented, ventrally directed gill (Fig. 3.17); mid tarsal segment at least half as long as apical segment (Fig. 3.18) ............................................. ............................................................................... TAENIOPTERYGIDAE (in part), 49 Coxae without segmented gills; mid tarsal segment at least half as long as apical segment (Fig. 3.18), or much smaller and wedge shaped (Fig. 3.19) ............................................... 4

4(3’) 4’

Thoracic sterna overlapping base of succeeding segment (Fig. 3.21); coxae bearing small dorsal lobe (Fig. 3.20); thoracic segments 2–3 with long, apically pointed lateral gills (Fig. 3.22) ..................................................................................... PELTOPERLIDAE, 25 Thoracic sterna not overlapping base of succeeding segment; coxae without small dorsal lobe; thoracic segments 2–3 without gills ................................................................................ 5

5(4’) 5’

Abdomen terminating in a large, ventral, shield-shaped plate (Fig. 3.23); mid tarsal segment at least half as long as apical segment (Fig. 3.18) ........................................................... ............................................................................... TAENIOPTERYGIDAE (in part), 49 Abdomen without terminal shield-shaped plate; mid tarsal segment small and wedge shaped (Fig. 3.19) ............................................................................................................... 6

Chapter 3 ~ Key to Stoneflies

Page 181

Fig. 3.11

Fig. 3.12

Fig. 3.13

Fig. 3.14

Fig. 3.17 Fig. 3.15

Fig. 3.16

Fig. 3.19

Fig. 3.18

Fig. 3.20

Fig. 3.21

Fig. 3.22

Fig. 3.23

Figures 3.11–3.23, Perlidae, Peltoperlidae, Pteronarcyidae and Taeniopterygidae genn. spp. 3.11, Perlinella drymo (Perlidae), left mesothoracic supracoxal gill, left lateral. 3.12, Tallaperla anna (Peltoperlidae), left mesothoracic supracoxal gill, left lateral. 3.13, Pteronarcys dorsata (Pteronarcyidae), right half of mesosternum and abdominal sterna 1–2 with gills, ventral. 3.14, Agnetina capitata (Perlidae), anal gills, ventral. 3.15, P. scotti (Pteronarcyidae), abdominal terga 1–4 and 8–10, dorsal. 3.16, Acroneuria carolinensis (Perlidae), abdominal tergum 10, dorsal. 3.17, Taeniopteryx maura (Taeniopterygidae), left forecoxa with gill, mesoventral. 3.18, T. maura, right hind tarsus, posterior. 3.19, Tallaperla anna (Peltoperlidae), right hind tarsus, posterior. 3.20, T. anna, base of left hind leg with coxal lobe, left lateral. 3.21, T. anna, thoracic sterna, ventral. 3.22, T. anna, left mesothoracic supracoxal gill, left lateral. 3.23, Strophopteryx appalachia, abdominal sterna 9–10 with projecting sternum 9, ventral. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

6(5’) 6’

Labial palps usually short, scarcely extending beyond anterior margin of paraglossae (Fig. 3.24); paraglossae and glossae subequal in length (Fig. 3.24) . ................................. 7 Labial palps project well beyond anterior margin of paraglossae (Fig. 3.25); paraglossae much larger than glossae (Fig. 3.25) . ................................................................................. 9

7(6) 7’

Hind wingpads diverging strongly from body axis (Fig. 3.26); extended hindlegs reach near abdominal apex; body robust; some species with ventral gills on neck (Fig. 3.27) ............................................................................................................ NEMOURIDAE, 19 Hind wingpads lying almost parallel to body axis (Fig. 3.28); extended hindlegs reach far short of abdominal apex; body slender; gills absent . ......................................................... 8

8(7’) 8’

Mentum small, not extending over base of maxilla; membranous folds occur ventrolaterally on abdominal segments 1-9; projecting flap of hind wingpads usually about as long as wide (Fig. 3.28) but may be absent or reduced . ...................................... CAPNIIDAE, 10 Mentum extends forward partially covering inner margins of maxillary base (Fig. 3.29); membranous folds occur ventrolaterally, at most on abdominal segments 1-7, absent on segment 8; hind wingpads usually longer than wide ........................... LEUCTRIDAE, 16

9(6’) 9’

Apical maxillary palpal segment small and set asymmetrically on penultimate segment (Fig. 3.30); thoracic nota usually without conspicuous pigment pattern; cerci about three fourths abdomen length .................................................. CHLOROPERLIDAE, 12 Apical maxillary palpal segment subequal to penultimate segment (Fig. 3.31); thoracic nota usually with conspicuous pigment pattern; cerci about as long as, or longer than abdomen (Fig. 3.32) . ........................................................................... PERLODIDAE, 37

CAPNIIDAE genera

(After Stewart & Stark, 2002, 2008.) 10(8) 10’

Body clothed with numerous long setae (Fig. 3.33); likely shredders/detritivores in leaf litter or hyporhea of small mountain streams; AR, CO, CT, DE, IL, KY, MA, MB, MD, ME, MI, MO, NC, NH, NL, NY, OH, OK, ON, PA, QC, SD, SK, TN, VA, WI, WV, WY ............................................................................................................ Paracapnia angulata Body clothed primarily with short setae (Fig. 3.34) ............................................................... 11

11(10’) 11’

Apical cercal segments with a vertical setal fringe (Fig. 3.36); likely shredders/detritivores in hyporhea of small to medium-sized sandy, Coastal Plain streams; AL, AR, FL, IL, IN, MS, NC, QC, SC, VA . ............................................................... Nemocapnia carolina Cercal segments without vertical setal fringe (Fig. 3.35) ................................... Allocapnia, 53

Chapter 3 ~ Key to Stoneflies

Fig. 3.24

Fig. 3.27

Fig. 3.30

Fig. 3.33

Page 183

Fig. 3.25

Fig. 3.26

Fig. 3.28

Fig. 3.29

Fig. 3.31

Fig. 3.34

Fig. 3.32

Fig. 3.35

Fig. 3.36

Figures 3.24–3.36, Nemouridae, Chloroperlidae, Perlodidae, Capniidae, Leuctridae genn. spp. 3.24, Amphinemura wui (Nemouridae), labium, ventral. 3.25, Sweltsa palearata (Chloroperlidae), labium, ventral. 3.26, A. wui, right halves of thoracic nota with wingpads, dorsal. 3.27, A. wui, cervical gills, ventral. 3.28, Allocapnia granulata (Capniidae), right halves of meso- and metathoracic nota with wingpads, dorsal. 3.29, Zealeuctra warreni (Leuctridae), head with mentum, ventral. 3.30, Sweltsa palearata (Chloroperlidae), left maxilla, ventral. 3.31, Diploperla duplicata (Perlodidae), left maxilla, ventral. 3.32, D. duplicata, left half of abdomen and left cercus, dorsal. 3.33, Paracapnia angulata (Capniidae), pronotum, dorsal. 3.34, Allocapnia virginiana (Capniidae), pronotum, dorsal. 3.35, A. virginiana, apical and basal cercal segments, dorsal. 3.36, Nemocapnia carolina (Capniidae), apical and basal cercal segments, dorsal. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

CHLOROPERLIDAE genera (After Stewart & Stark, 2002, 2008.)

12(9) 12’

Apical cercal segments usually with a vertical setal fringe (Fig. 3.37) ................ Alloperla, 63 Apical cercal segments without vertical setal fringe .............................................................. 13

13(12’) 13’

Thoracic mesosternum bearing anterolateral patches of pale brown to dark brown or black clothing hairs (Fig. 3.38) ................................................................................... Sweltsa, 72 Thoracic mesosternum without anterolateral patches of pigmented clothing hairs . .............. 14

14(13’) 14’

Longest pronotal fringe hairs about 0.3-0.4 times as long as pronotal width (Fig. 3.40) and subequal to abdominal tergum 9; inner margins of wingpads nearly parallel with body axis . ......................................................................................................................... 15 Longest pronotal fringe hairs about 0.2 times as long as pronotal width (Fig. 3.39) and shorter than abdominal tergum 9; inner margins of wingpads divergent from body axis (Fig. 3.39); likely facultative detritivores/ omnivores/carnivores; clingers in mountain streams; GA, MA, MD, ME, MI, NB, NC, NH, NL, NY, ON, PA, QC, TN, VA, VT, WI, WV .............................................................................................................. Suwallia marginata

15(14) 15’

Abdomen with longitudinal rows of dark pigment spots (Fig. 3.41), sometimes obscure in immature specimens; anterior and posterior margins of pronotum bearing numerous long setae, lateral margins with shorter setae (Fig. 3.40); rarely collected, larval biology unknown; ME, NC, NH, NY, PA, TN, VA, VT, WV . .................................. Rasvena terna Abdomen without pattern of dark pigment spots (Fig. 3.42); long hairs sparse on anterior pronotal margin, but abundant on posterior margin (Fig. 3.43); widespread and often common .............................................................................................. Haploperla, 71

LEUCTRIDAE genera

(After Stewart & Stark 2002, 2008.) 16(8’) 16’

Body form robust (Fig. 3.44); tergum 10 and fused paraprocts forming triangular projection extending caudally beyond basal 2 cercal segments in pre-emergent males (Fig. 3.45), or with a small triangular projection on sternum 8 in pre-emergent females (Fig. 3.46); pronotum completely fringed laterally with short, thick setae (Fig. 3.47); found in small spring seeps . .............................................................................................. Megaleuctra, 86 Body form slender (Fig. 3.50); tergum 10, paraprocts or sternum 8 not forming slender triangular projections extending beyond basal 2 cercal segments (Fig. 3.49); pronotum without complete lateral fringe of thick setae . ................................................................. 17

17(16’) 17’

Pronotum without long setae on corners (Fig. 3.51) . ......................................... Zealeuctra, 87 Pronotum with cluster of long setae on corners (Fig. 3.48) . .................................................. 18

Chapter 3 ~ Key to Stoneflies

Page 185

Fig. 3.39 Fig. 3.37

Fig. 3.38

Fig. 3.41 Fig. 3.47 Fig. 3.40

Fig. 3.43

Fig. 3.42

Fig. 3.45 Fig. 3.48

Fig. 3.46

Fig. 3.44

Fig. 3.50

Fig. 3.49

Fig. 3.51

Figures 3.37–3.51, Chloroperlidae and Leuctridae genn. spp. 3.37, Alloperla concolor (Chloroperlidae), apical and basal cercal segments, lateral. 3.38, Sweltsa pocahontas (Chloroperlidae), mesosternum, ventral. 3.39, Suwallia marginata (Chloroperlidae), right halves of thoracic nota, dorsal. 3.40, Rasvena terna (Chloroperlidae), pronotum, dorsal. 3.41, R. terna, abdominal terga 7–10, dorsal. 3.42, Haploperla chukcho (Chloroperlidae), abdominal terga 7–10, dorsal. 3.43, H. brevis, pronotum, dorsal. 3.44. Megaleuctra kincaidi (Leuctridae), body outline, dorsal. 3.45, M. kincaidi, male abdominal sterna 8–10, ventral. 3.46, M. kincaidi, female abdominal sterna 8–10, ventral. 3.47, M. kincaidi, pronotum, dorsal. 3.48, Paraleuctra sara (Leuctridae), pronotum, dorsal. 3.49, Zealeuctra claasseni (Leuctridae), abdominal sterna 8–10, ventral. 3.50, Z. claasseni, body outline, dorsal. 3.51, Z. claasseni, pronotum, dorsal. Figures 3.45-3.46, 3.49 redrawn from Stewart & Stark (2002); all other figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

18(17’)

Abdominal terga without posterior setal fringes, or short setae on surface, although one or more long setae may occur laterally (Fig. 3.52); mesosternal Y-arms with double stem and median dark band (Fig. 3.53); abdominal segments 1–6 divided by ventrolateral membrane; labial palps barely reaching beyond tips of paraglossae (Fig. 3.54); larval biology unknown; AL, CT, DE, KY, MA, MD, ME, NB, NC, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA,WV . ....................................................................... Paraleuctra sara Abdominal terga with posterior setal fringes and short surface setae (Fig. 3.55); mesosternal Y-arm stem single, median band absent (Fig. 3.56); abdominal segments 1–4 divided by ventrolateral membrane; labial palps extending well beyond tips of paraglossae ............................................................................................................................ Leuctra, 79

18’

NEMOURIDAE genera

(After Baumann, 1975; Stewart & Stark, 2002, 2008.) 19(7) 19’

Gills present in neck region (Figs. 3.57-3.58) ........................................................................ 20 Gills absent from neck region ................................................................................................. 21

20(19) 20’

Femora bearing a transverse row of long, stout setae (Fig. 3.59) . .......................... Zapada, 96 Femora without transverse row of setae ....................................................... Amphinemura, 88

21(19’) 21’

Pronotum usually with small lateral notch (Fig. 3.60); marginal pronotal setal fringe composed of close-set, relatively uniform, short thick setae (Fig. 3.60) ........ Soyedina, 95 Pronotum without lateral notch (Fig. 3.61); pronotal setal fringe may vary in spacing or setal length and sometimes is poorly developed . ............................................................. 22

22(21’) 22’

Outer fore tibial fine setal fringe well developed (Fig. 3.63) ................................................. 23 Outer fore tibial fine setal fringe sparse (Fig. 3.62) . .............................................................. 24

23(22)

Fore tibiae with obscure, unpigmented thick marginal setae along base of fine setal fringe, and usually with a few long silky setae along inner tibial margin (Fig. 3.63); legs obscurely banded; marginal pronotal setae inconspicuous (3.64); shredders/detritivores/ scrapers; sprawlers/ clingers in medium-sized to large creeks and rivers; AB, AK, AL, AR, IL, MB, MD, ME, MI, MN, MS, NB, NC, NT, ON, QC, SC, SK, VA, WI ..................................................................................................................... Shipsa rotunda Fore tibiae with conspicuous short, thick outer marginal setae (Fig. 3.65); inner marginal long silky setae absent from tibiae; legs not banded; marginal pronotal setae conspicuous but irregularly spaced (Fig. 3.66) . .................................................................... Prostoia, 93

23’

Chapter 3 ~ Key to Stoneflies

Page 187

Fig. 3.53

Fig. 3.54

Fig. 3.52

Fig. 3.56

Fig. 3.57

Fig. 3.55 Fig. 3.58 Fig. 3.62

Fig. 3.59 Fig. 3.60

Fig. 3.61

Fig. 3.64

Fig 3.66

Fig. 3.65

Fig. 3.63

Figures 3.52–3.66, Leuctridae and Nemouridae genn. spp. 3.52, Paraleuctra sara (Leuctridae), abdominal terga 8–10, dorsal. 3.53, P. sara, thoracic mesosternum, ventral. 3.54, P. sara, labium, ventral. 3.55, Leuctra sibleyi (Leuctridae), abdominal terga 8–10, dorsal. 3.56, L. sibleyi, mesosternum, ventral. 3.57, Zapada chila (Nemouridae), cervical gills and prosternum, ventral. 3.58, Amphinemura wui (Nemouridae), cervical gills and prosternum, ventral. 3.59, Zapada chila (Nemouridae), left foreleg, anterior. 3.60, Soyedina carolinensis (Nemouridae), left half of pronotum, dorsal. 3.61, Ostrocerca truncata (Nemouridae), left half of pronotum, dorsal. 3.62, O. truncata, right foreleg, anterior. 3.63, Shipsa rotunda (Nemouridae), right foreleg, anterior. 3.64, S. rotunda, left half of pronotum, dorsal. 3.65, Prostoia completa (Nemouridae), right foreleg, anterior. 3.66, P. completa, left half of pronotum, dorsal. Figures 3.53, 3.55, 3.56 redrawn from Stewart & Stark (2002); all other figures original.

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24



24’

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Marginal pronotal setae short and irregularly spaced (Fig. 3.67); fore tibiae with submarginal row of thick setae fairly complete (Fig. 3.68); known primarily from springs and small first-order rheocrenes, uncommon, or absent in much of southeastern U.S. ....................................................................................................................... Ostrocerca, 92 Marginal pronotal setae inconspicuous; sub-marginal thick setae of fore tibiae scattered, not organized into a distinct row (Fig. 3.69); uncommon, likely shredders/detritivores; clingers; CT, MA, MD, ME, MI, NB, NC, NH, NS, NY, ON, PA, PE, QC, TN, VA, VT, WV ......................................................................................................... Paranemoura perfecta

PELTOPERLIDAE genera

(After Stark & Stewart, 1982; Stewart & Stark, 2002, 2008.) 25(4) 25’

Single gills located above coxae on meso- and metathorax (Fig. 3.70); long fringe setae on hind tibiae reduced to a few hairs (Fig. 3.71); shredders/detritivores; sprawlers/ clingers in small mountain streams and spring seeps; GA, NC, SC, TN ...... Viehoperla ada Double gills located above coxae on meso- and metathorax (Fig. 3.72); long setal fringe on hind tibiae relatively well developed ................................................................................ 26

26(25’) 26’

Hind femora with sparse fringe of long setae (Fig. 3.73); meso- and metanota often with a pair of dark pigment spots (Fig. 3.75); larvae of two known species not separable, but P. tarteri rare in Region IV .................................................................................. Peltoperla Hind femora with relatively well developed long setal fringe (Fig. 3.74); meso- and metanota often without pair of dark pigment spots; larvae of seven known species not separable at present .............................................................................................. Tallaperla

PERLIDAE genera

(After Stark & Gaufin, 1976; Stewart & Stark, 2002, 2008.) 27(2) 27’

Two ocelli (Fig. 3.76) ............................................................................................................. 28 Three ocelli (Fig. 3.77) ........................................................................................................... 29

28(27) 28’

Occiput with low transverse ridge extending across head behind compound eyes (Fig. 3.76); abdominal terga conspicuously pigmented with transverse dark and pale bands ........................................................................................................................ Neoperla, 112 Occiput without transverse ridge behind compound eyes (Fig. 3.78); abdominal terga without transverse pigment bands . ............................................... Perlinella (in part), 125

Chapter 3 ~ Key to Stoneflies

Page 189

Fig. 3.67

Fig. 3.69

Fig. 3.68

Fig. 3.70

Fig. 3.71 Fig. 3.72

Fig. 3.75

Fig. 3.74

Fig. 3.76

Fig. 3.73

Fig. 3.77

Fig. 3.78

Figures 3.67–3.78, Nemouridae, Peltoperlidae, Peltoperlidae and Perlidae genn. spp. 3.67, Ostrocerca truncata (Nemouridae), left half of pronotum, dorsal. 3.68, O. truncata, apex of left foretibia, anterior. 3.69, Paranemoura perfecta (Nemouridae), apex of left foretibia, anterior. 3.70, Viehoperla ada (Peltoperlidae), left mesothoracic supracoxal gill, left lateral. 3.71, V. ada, left hind leg, anterior. 3.72, Tallaperla anna (Peltoperlidae), left mesothoracic supracoxal gill, left lateral. 3.73, Peltoperla arcuata (Peltoperlidae), left hind femur, anterior. 3.74, Tallaperla anna (Peltoperlidae), left hind femur, anterior. 3.75, P. arcuata, left halves of meso- and metathoracic nota, dorsal. 3.76, Neoperla clymene (Perlidae), head, dorsal. 3.77, Perlesta sp. (Perlidae), head, dorsal. 3.78, Perlinella drymo (Perlidae), head, dorsal. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

29(27’) 29’

Occiput with a close-set, complete, relatively straight, transverse row of short peg-like setae extending across head behind compound eyes (Fig. 3.79) ...................................... 30 Occiput with peg setae separated by irregular gaps (Fig. 3.80), or clustered 2 or 3 deep in areas, or present only near postocular fringe (Fig. 3.81), or row sinuate ......................... 31

30(29) 30’

Basal cercal segments with inner fringe of long, silky setae (Fig. 3.82); anal gills usually absent, sometimes a few filaments present; posterior setal fringe on abdominal sternum 7 interrupted mesally ............................................................................... Paragnetina, 115 Basal cercal segments without inner fringe of long, silky setae (Fig. 3.83); anal gills well developed; posterior fringe on abdominal sternum 7 complete mesally (Fig. 3.84) ....................................................................................................................... Agnetina, 107

31(29’)



Pronotum fringed along lateral and posterior margins with long, thin setae (Fig. 3.85); occipital row of peg setae almost complete and only slightly irregular; pronotum about 2.5 times as wide as median length; anal gills absent; predators/engulfers; sprawlers/ clingers in medium-sized to large streams, often associated with organic substrate in deeper water; AL, AR, DC, FL, GA, IA. IL, IN, KS, MB, MD, MI, MN, MO, MS, NC, NE, NY, PA, OH, SC, TN, VA, WI ........................................................ Attaneuria ruralis Lateral and posterior pronotal fringe consists of short, thick setae interspersed with a few long setae (Fig. 3.86); occipital setal row variable, but with conspicuous irregularities, or reduced to a few setae near postocular fringe (Fig. 3.90); pronotum about 1.5 to 2.2 times as wide as median length; anal gills present or absent . .......................................... 32

32(31’) 32’

Postocular fringe consisting of a few, often 1 or 2, long setae (Fig. 3.89); pronotal fringe consisting of about 2 to 5 long setae at posterolateral and anterolateral corners (Fig. 3.87) ......................................................................................................................... 33 Postocular fringe consisting of a curved row of several thick setae (Fig. 3.88); pronotal fringe consisting of variable length thick setae in a close-set row, often interrupted on lateral margins (Fig. 3.86) . ............................................................................................... 34

31’

Chapter 3 ~ Key to Stoneflies

Fig. 3.79

Page 191

Fig. 3.80

Fig. 3.81

Fig. 3.84 Fig. 3.82

Fig. 3.83

Fig. 3.85

Fig. 3.86

Fig. 3.88

Fig. 3.89

Fig. 3.87

Fig. 3.90

Figures 3.79–3.90, Perlidae genn. spp. 3.79, Paragnetina fumosa, head, dorsal. 3.80, Perlesta sp., head, dorsal. 3.81, Acroneuria sp., head, dorsal. 3.82, Paragnetina immarginata, abdominal terga 8–10, dorsal. 3.83, Agnetina flavescens, abdominal terga 8–10, dorsal. 3.84, A. flavescens, apex of abdominal sternum 7, ventral. 3.85, Attaneuria ruralis, left half of pronotum, dorsal. 3.86, Acroneuria sp., left half of pronotum, dorsal. 3.87, Perlinella drymo, left half of pronotum, dorsal. 3.88, Perlesta sp., head, dorsal. 3.89, Perlinella drymo, right half of head with postocular fringe, dorsal. 3.90, Acroneuria sp., head, dorsal. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

33(32) 33’

Anal gills present; femora and tibiae with dorsal and ventral fringes of long, silky setae (Fig. 3.91); widespread and common ........................................... Perlinella (in part), 125 Anal gills absent; femora and tibiae without ventral fringes of long, silky setae (Fig. 3.92); uncommon and restricted in distribution............................................... Hansonoperla, 111

34(32’) 34’

Occiput usually with partial row of short, thick setae extending to near stem of ecdysial suture (Fig. 3.93); abdominal terga often yellow-brown with darker, freckle-like pigment spots around setal bases (Fig. 3.94) ................................................................ Perlesta, 120 Occipital setal row reduced to a few thick setae located near postocular fringe (Fig. 3.95); abdominal terga without dark pigment spots around setal bases ...................................... 35

35(34’)

Anterior region of frons anterior to median ocellus mostly yellow with dark lateral margins (Fig. 3.96); predators/engulfers; sprawlers/clingers usually in small streams of Coastal Plain, Piedmont and Appalachians; AL, CT, DE, FL, GA, KY, MD, MS, NC, NY, OH, PA, SC, TN, VA, WV ........................................................................ Eccoptura xanthenes

35’

Head with extensive brown pigment anterior to median ocellus, often patterned with a pale M-line (Fig. 3.95) ............................................................................................................. 36

36(35’) 36’

Cerci without basal fringe of long silky setae (Fig. 3.97); pronotal flange narrow throughout (Fig. 3.99); anal gills present; uncommon except in small Appalachian or Piedmont streams and seeps . ..................................................................................... Beloneuria, 109 Cerci with basal fringe of long silky setae (Fig. 3.98), but sometimes reduced; pronotal flange wider at posterior angles than along lateral margins (Fig. 3.100); anal gills present or absent; common in small, medium-sized, and large streams throughout the southeastern region ...................................................................................... Acroneuria, 97



PERLODIDAE genera

(After Stark & Stewart, 2002, 2008.) 37(9’) 37’

Femora and tibiae without long silky setal fringe (Fig. 3.101); pronotum excavated on posterolateral margins (Fig. 3.102); stem of mesosternal Y poorly developed, arms diverging from base; larval biology unknown; found in splash zones under large rocks in first-order rheocrenes and springs; NC, SC, TN ........................... Oconoperla innubila Femora and tibiae with long silky setal fringe; pronotum without posterolateral notch; mesosternal Y-arms usually diverging from well-developed stem ................................... 38

Chapter 3 ~ Key to Stoneflies

Fig. 3.91

Page 193

Fig. 3.92 Fig. 3.93

Fig. 3.94

Fig. 3.97

Fig. 3.95

Fig. 3.96

Fig. 3.98 Fig. 3.99

Fig. 3.101 Fig. 3.100

Fig. 3.102

Figures 3.91–3.102, Perlidae and Perlodidae genn. spp. 3.91, Perlinella drymo (Perlidae), left foreleg, anterior. 3.92, Hansonoperla appalachia (Perlidae), left foreleg, anterior. 3.93, Perlesta sp. (Perlidae), head, dorsal. 3.94, Perlesta sp., abdominal terga 8–10, dorsal. 3.95, Acroneuria sp. (Perlidae), head, dorsal. 3.96, Eccoptura xanthenes (Perlidae), head, dorsal. 3.97, Beloneuria georgiana (Perlidae), abdominal terga 8–10, dorsal. 3.98, Acroneuria sp. (Perlidae), abdominal terga, 8–10, dorsal. 3.99, B. georgiana (Perlidae), left half of pronotum, dorsal. 3.100, Acroneuria sp. (Perlidae), left half of pronotum, dorsal. 3.101, Oconoperla innubila (Perlodidae), left foreleg, anterior. 3.102, O. innubila, left half of pronotum, dorsal. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

38(37’) 38’

Abdomen with dark longitudinal stripes (Fig. 3.103) . .......................... Isoperla (in part), 135 Abdomen without dark longitudinal stripes . .......................................................................... 39

39(38’) 39’

Lacinia with a single tooth, broad basally and narrowed into a long spine-like structure (Fig. 3.104) ................................................................................................... Remenus, 150 Lacinia with two teeth (Fig. 3.105) . ....................................................................................... 40

40(39’) 40’

Anterolateral pronotal fringe reduced to a few long setae, short, thick setae absent (Fig. 3.106) ....................................................................................................................... 41 Anterolateral pronotal fringe consisting of row of short, thick setae, sometimes interspersed with longer setae (Fig. 3.108) ........................................................................................... 42

41(40) 41’

Dorsum of cerci without fringe of long setae; mesosternal Y-arms without common stem and fork, arms diverging from base (may be difficult to observe) (Fig. 3.107) ..................................................................................................................... Diploperla, 128 Cerci with dorsal fringe of long, silky setae, most conspicuous in lateral aspect (Fig. 3.109); mesosternal Y-arms diverging from a common stem (Fig. 3.110) . ................... Cultus, 127

42(40’) 42’

Submental gills conspicuous, projecting portion usually 2 or more times as long as basal diameter (Fig. 3.111) . ....................................................................................................... 43 Submental gills absent, or barely projecting beyond margin of submentum . ........................ 46

43(42) 43’

Mesosternum with median longitudinal suture connecting fork of Y-arms to anterior transverse suture (Fig. 3.112)..................................................................... Isogenoides, 134 Mesosternum without median longitudinal suture (Fig. 3.113) . ............................................ 44

44(43’)

Occipital spinules mostly consisting of a single row; ventral lacinial surface with basal patch of about 50 dark clothing hairs (Fig. 3.114); likely predators/engulfers; sprawlers/ clingers in small first-order to third-order Appalachian streams; GA, KY, NC, OH, PA, SC, TN, VA, WV ................................................................... Malirekus hastatus (in part)

Chapter 3 ~ Key to Stoneflies

Page 195

Fig. 3.103 Fig. 3.104

Fig. 3.106

Fig. 3.105

Fig. 3.107 Fig. 3.108

Fig. 3.109

Fig. 3.112

Fig. 3.110

Fig. 3.111

Fig. 3.113 Fig. 3.114

Figures 3.103–3.114, Perlodidae genn. spp. 3.103, Isoperla sp., abdominal terga 7–10, dorsal. 3.104, Remenus bilobatus, left lacinia, ventral. 3.105, Diploperla duplicata, left lacinia, ventral. 3.106, D. duplicata, left half of pronotum, dorsal. 3.107, D. duplicata, mesosternum, ventral. 3.108, Isogenoides varians, left half of pronotum, dorsal. 3.109, Cultus verticalis, right sides of abdominal segments 8–10 and base of right cercus, right lateral. 3.110, C. verticalis, mesosternum, ventral. 3.111, Isogenoides varians, labium with submental gills, ventral. 3.112, I. varians, mesosernum, ventral. 3.113, Helopicus bogaloosa, mesosternum, ventral. 3.114, Malirekus hastatus, left lacinia, ventral. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

44’

Occipital spinules comprising rows 2-3 deep; lacinial surface without patch of brown clothing hairs, but a patch of about 10 setae may be present (Fig. 3.115) . ...................... 45

45(44’)

Transverse dark pigment band crossing frons without interruption by enclosed yellow pigment spots (Fig. 3.116); ventral lacinial surface without outer patch of setae; inner margin of largest tooth of right mandible with fine serrations (Fig. 3.117) ...................................................................................................................... Helopicus, 131 Transverse dark pigment band of frons interrupted by enclosed yellow areas near anterior ocellus (Fig. 3.118); ventral lacinial surface with outer patch of about 10 setae (Fig. 3.115); inner margins of teeth on right mandible without serrations (Fig. 3.119) .................................................................................................................... Hydroperla, 132

45’ 46(42’)

46’

Ocellar region crossed by dark pigment band almost connecting lateral ocelli and compound eyes, but not covering anterior ocellus (Fig. 3.120); mature individuals predators/ engulfers, early instars shift from herbivory/detritivory and omnivory; clingers to leaf packs and stones in medium-sized to large creeks; AL, AR, CT, DE, FL, GA, IA, IL, IN, KY, MA, MD, MI, MO, MS, NC, OH, OK, ON, PA, SC, TN, VA, WI, WV ........................................................................................................................ Clioperla clio Dorsal pigment pattern on head variable, but not as above; ocellar area often covered with dark pigment ..................................................................................................................... 47

47(46’) 47’

Lacinial margin without knob below subapical tooth (Fig. 3.121) . ...... Isoperla (in part), 135 Lacinial margin with low knob below subapical tooth (Fig. 3.122) ....................................... 48

48(47’)

Occipital spinules mostly consisting of a single row; ventral lacinial surface with basal patch of about 50 dark clothing hairs (Fig. 3.122); likely predators/engulfers; sprawlers/ clingers in small first-order to third-order Appalachian streams; GA, KY, NC, OH, PA, SC, TN, VA, WV ................................................................... Malirekus hastatus (in part) Occipital spinules mostly consisting of an irregular double row; ventral lacinial surface without basal patch of dark clothing hairs (Fig. 3.123) ..................................... Yugus, 151

48’

TAENIOPTERYGIDAE genera (After Stark & Stewart, 2002, 2008.)

49(3, 5) Each coxa bearing a single, segmented, ventrally directed gill (Fig. 3.124); apex of abdomen without a projecting, ventral triangular plate on segment 9 (Fig. 3.125) ........................ ................................................................................................................. Taeniopteryx, 161 49’ Coxae without gills; apex of abdomen with a ventral, projecting, triangular plate on segment 9 (Fig. 3.126) ...................................................................................................... 50

Chapter 3 ~ Key to Stoneflies

Page 197

Fig. 3.115

Fig. 3.116

Fig. 3.117

Fig. 3.118

Fig. 3.119

Fig. 3.120

Fig. 3.124 Fig. 3.121

Fig. 3.122

Fig. 3.125

Fig. 3.123

Fig. 3.126

Figures 3.115–3.126, Perlodidae and Taeniopterygidae genn. spp. 3.115, Hydroperla sp. (Perlodidae), left lacinia, ventral. 3.116, Helopicus bogaloosa (Perlodidae), head, dorsal. 3.117, H. bogaloosa, left mandible, dorsal. 3.118, Hydroperla sp., head, dorsal. 3.119, Hydroperla sp., left mandible, dorsal. 3.120. Clioperla clio (Perlodidae), head, dorsal. 3.121, Isoperla sp. (Perlodidae), left lacinia, ventral. 3.122, Malirekus hastatus (Perlodidae), left lacinia, ventral. 3.123, Yugus sp. (Perlodidae), left lacinia, ventral. 3.124, Taeniopteryx maura (Taeniopterygidae), left forecoxa with gill, mesoventral. 3.125, T. maura, abdominal sterna 8–10, ventral. 3.126, Strophopteryx appalachia (Taeniopterygidae), abdominal sterna 9–10, ventral. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

50(49’) 50’

Basal 2-3 antennal segments bearing an obscure dorsal grouping of fine setae (Fig. 3.127); abdominal terga usually dark brown (Fig. 3.128); uncommon, likely scrapers or facultative shredders; sprawlers/clingers in rocky second-order to fourth-order Appalachian streams; MA, ME, NC, NH, NY, PA, QC, SC, TN, VA, WV ............................... Bolotoperla rossi Basal antennal segments without groupings of fine setae; abdominal terga variable, often pale brown, banded, or with rows of dark spots ............................................................... 51

51(50’) 51’

Basal 3–6 cercal segments each usually bearing a single, short fine seta on dorsum (Fig. 3.129); if basal fine setae absent, tibiae bearing a distinct outer and sparse inner setal fringe (Fig. 3.130) ........................................................................ Strophopteryx, 159 Basal cercal segments without fine setae (3.131); tibiae without inner setal fringe................ 52

52(51’)

Body and legs brown with indistinct pattern; pronotum usually wider on posterior margin than on anterior margin (Fig. 3.132); likely shredders/ scrapers/detritivores; clingers in second-order to fourth-order Appalachian streams; CT, KY, MA, MD, ME, NB, NC, NH, NL, NS, NY, OH, PA, QC, SC, TN, VA, VT, WV ............... Taenionema atlanticum Body and legs pale with distinct pattern; pronotum about as wide on anterior margin as on posterior margin; likely scrapers/ detritivores; clingers in medium-sized to large Appalachian streams; CT, KY, MA, MD, ME, NC, NH, PA, TN, VA, WV ............................................................................................................ Oemopteryx contorta

52’

SPECIES CAPNIIDAE, Allocapnia

(After Harper & Hynes, 1971b; Stark & Lacey, 2005. Suitable characters are vailable for only 11 of the 38 regional species.) 53(11’) 53’

Galea in lateral aspect bearing an outer marginal fringe of short setae for at least one third of length (Fig. 3.133); shredders/detritivores; clingers in leaf packs, early instars in hyporhea; CT, DC, IA, KY, MA, MD, ME, MI, MN, MO, NB, ND, NH, NS, NY, OH, ON, PA, QC, RI, TN,VA, VT, WI, WV ...................................... Allocapnia pygmaea Galea bearing a few short, thick sensilla near tip, but fringe absent or reduced to a few setae (Fig. 3.134) ....................................................................................................................... 54

54(53’) 54’

Lateral aspect of abdominal terga usually bearing one or more erect setae near tergal midlength (Fig. 3.135) ...................................................................................................... 55 Erect setae seen in lateral aspect restricted to posterior margins of most abdominal terga (Fig. 3.136) ....................................................................................................................... 59

Chapter 3 ~ Key to Stoneflies

Page 199

Fig. 3.127 Fig. 3.129 Fig. 3.128

Fig. 3.130 Fig. 3.131

Fig. 3.132

Fig. 3.135

Fig. 3.133

Fig. 3.134

Fig. 3.136

Figures 3.127–3.136, Taeniopterygidae genn. spp. and Allocapnia spp. (Capniidae). 3.127, Bolotoperla rossi (Taeniopterygidae), right half of head and right antennal base, dorsal. 3.128, B. rossi, abdominal terga 9–10, dorsal. 3.129, Strophopteryx appalachia (Taeniopterygidae), right sides of abdominal segments 8–10 and base of right cercus, right lateral. 3.130, S. appalachia, left foreleg, anterior. 3.131, Taenionema atlanticum (Taeniopterygidae), right sides of abdominal segments 8–10 and base of right cercus, right lateral. 3.132, T. atlanticum, head and pronotum, dorsal. 3.133, Allocapnia pygmaea (Capniidae), left maxilla with base of palp, ventral. 3.134, A. aurora, left maxilla with base of palp, ventral. 3.135, A. virginiana, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.136, A. aurora, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. Figure 3.133 redrawn from Harper & Hynes (1971b); all other figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

55(54) 55’

Erect setae scattered along entire tergal length (Fig. 3.137); shredders/ detritivores; clingers in leaf packs, early instars in hyporhea; not known to occur in Region IV, rare in adjacent states; IL, IN, ME, MN, NY, OH, ON, QC, VA, WI, WV................. Allocapnia illinoensis Erect setae restricted to posterior margin and midlength of abdominal terga (Fig. 3.140)..... 56

56(55’) 56’

Erect setae on posterior margins of terga about half as long as mid-dorsal tergal length ...... 57 Erect setae on posterior margins of terga about one third, or less, as long as middorsal tergal length (Fig. 3.139) . ...................................................................................... 58

57(56)

Lateral aspect of developing male 10th tergum bent upward near tip (Fig. 3.138); shredders/ detritivores; clingers in leaf packs, early instars in hyporhea; AL, AR, DC, DE, GA, IA, IN, KS, KY, MD, MN, MO, MS, NC, NJ, NY, OH, OK, ON, PA, TN, VA, WI, WV ....................................................................................................................... Allocapnia rickeri Lateral aspect of developing male 10th tergum slanted slightly upward, but not bent (Fig. 3.140); shredders/detritivores; clingers in leaf packs, early instars in hyporhea; AL, DE, GA, LA, MS, NC, SC, VA ............................................... Allocapnia virginiana

57’ 58(56’)



Developing epiproct of mature male about twice as long as remainder of tergum 10 (Fig. 3.141); mature developing male without wingpads; head and pronotum hairy in lateral aspect; shredders/ detritivores; clingers in leaf packs, early instars in hyporhea; AR, DC, IA, IL, IN, KS, KY, MD, MI, MN, MO, NE, NY, OH, OK, ON, PA, QC, TN, VA, WI, WV . ....................................................................................... Allocapnia vivipara Developing epiproct of mature male about as long as remainder of 10th tergum (Fig. 3.142); developing male with short wingpads; head and pronotum with few hairs in lateral aspect; shredders/detritivores; clingers in leaf packs, early instars in hyporhea; AL, AR, DC, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, NJ, NY, OH, OK, ON, PA, QC, TN, TX, VA, WI, WV . ....................................................... Allocapnia granulata

59(54’) 59’

Developing epiproct of mature male subequal to, or slightly longer than, remainder of tergum 10 (Fig. 3.143) ...................................................................................................... 60 Developing epiproct of mature male about 1.5 to twice as long as remainder of tergum 10 (Fig. 3.144) ....................................................................................................................... 61

60(59) 60’

Widely distributed outside Mississippi; shredders/detritivores; clingers in leaf packs, early instars in hyporhea; AL, CT, DC, DE, GA, IL, IN, KY, MA, MD, ME, NC, NH, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV ................................... Allocapnia recta Known only from Mississippi, the Florida Parishes of Louisiana and the Florida Panhandle; shredders/detritivores; clingers in leaf packs, early instars in hyporhea of small streams; FL, LA, MS ............................................................................................. Allocapnia starki

58’

Chapter 3 ~ Key to Stoneflies

Page 201

Fig. 3.137

Fig. 3.139

Fig. 3.138

Fig. 3.140

Fig. 3.141 Fig. 3. 142

Fig. 3.143

Fig. 3.144

Figures 3.137–3.144, Allocapnia spp. (Capniidae). 3.137, Allocapnia illinoensis, right sides of abdominal segments 7–10 and base of right cercus, developing male nymph, right lateral. 3.138, A. rickeri, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.139, A. granulata, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.140, A. virginiana, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.141, A. vivipara, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.142, A. granulata, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.143, A. starki, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.144, A. aurora, right sides of abdominal segments 8–10 and right cercus, developing male nymph, right lateral. Figures 3.137 and 3.141 redrawn from Harper & Hynes (1971b); all other figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

61(59’) 61’

Developing epiproct of mature male strongly slanted upward and truncate at tip (Fig. 3.145); shredders/detritivores; clingers in leaf packs, early instars in hyporhea; AL, AR, GA, IL, IN, KY, MO, MS, OH, TN, VA, WV ..................... Allocapnia mystica Tip of developing epiproct of male slightly, or not slanted, tip variable, usually either slightly enlarged on dorsal margin (Fig. 3.146), or notched along apical margin (Fig. 3.148) ....................................................................................................................... 62

62(61’)

Tip of developing male epiproct slightly enlarged, bulb-like (Fig. 3.146); galea without apical fringe hairs (Fig. 3.147); shredders/detritivores; clingers in leaf packs, early instars in hyporhea of small streams; AL, DC, GA, MD, MS, NC, PA, SC, TN, VA ................................................................................................................ Allocapnia aurora Tip of developing male epiproct not bulb-like, but tip notched in lateral aspect (Fig. 3.148); galea with a few apical fringe hairs forming a weak tuft (Fig. 3.149); shredders/ detritivores; clingers in leaf packs, early instars in hyporhea; AL, CT, DC, DE, IL, IN, KY, MA, MD, NB, NC, NJ, NS, NY, OH, PA, QC, RI, TN, VA, VT, WI, WV .............................................................................................................. Allocapnia nivicola

62’

CHLOROPERLIDAE, Alloperla

(After Stark & Kondratieff, 2010, with characters extracted from Brown & Stark, 1995; Fiance, 1977; Poulton & Stewart, 1991; Ray et al., 2010; Stewart & Stark, 2002. Suitable characters are available for nine of the 20 regional species; however, sample size for specimens used in developing this key is small and the key should be considered provisional.). Larvae often are absent in samples where adults are common. 63(12) 63’

Vertical setal fringe on apical four cercal segments poorly developed, sparse on ventral margin and reduced or absent on dorsal margin (Fig. 3.150); anterolateral margins of frons slightly angular from base of frontoclypeus to labrum (Fig. 3.151); likely predators/engulfers; hyporheic for most of larval development; AL, GA, KY, MD, NC, OH, PA, SC, TN, VA, WV . ................................................................... Alloperla usa Vertical setal fringe well developed on mid- to apical cercal segments (Fig. 3.152); anterolateral margins of frons rounded between frontoclypeus and labrum (Fig. 3.153) ....................................................................................................................... 64

64(63’) 64’ 65(64) 65’

Twelve or 13 cercal segments; Coastal Plain populations found south of Jackson (MS), Tuscaloosa (AL), Atlanta (GA) line . ................................................................................ 65 At least 14 cercal segments; populations found north of Jackson, Tuscaloosa, Atlanta line . ... 67 Thirteen cercal segments; ventral fringe on foretibiae absent (Fig. 3.155); known from central and southwest Mississippi; likely predators/ engulfers; hyporheic for most of larval development; MS . ......................................................................... Alloperla natchez Twelve cercal segments; sparse ventral fringe of fine setae usually present on foretibiae (Fig. 3.154) ....................................................................................................................... 66

66(65’)

Abdominal terga with pair of pale, round spots (Fig. 3.156); known from Coastal Plain of South Carolina; likely predators/engulfers; hyporheic for most of larval development; SC ........................................................................................................................ Alloperla furcula Abdominal terga without pair of pale round spots; known from southern Alabama and Florida Panhandle; likely predators/engulfers; hyporheic for most of larval development: AL, FL ............................................................................................... Alloperla prognoides

66’

Chapter 3 ~ Key to Stoneflies

Page 203

Fig. 3.146 Fig. 3.145

Fig. 3.147

Fig. 3.149

Fig. 3.148

Fig. 3.150

Fig. 3.152

Fig. 3.151

Fig. 3.153

Fig. 3.155

Fig. 3.154

Fig. 3.156

Figures 3.145–3.156, Allocapnia spp. (Capniidae) and Alloperla spp. (Chloroperlidae). 3.145, Allocapnia mystica, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.146, A. aurora, right sides of abdominal segments 8–10 and base of right cercus, developing male nymph, right lateral. 3.147, A. aurora, left maxilla with base of palp, ventral. 3.148, A. nivicola, right sides of abdominal segments 8–10, developing male nymph, right lateral. 3.149, A. nivicola, left maxilla, ventral. 3.150, Alloperla usa, apex of right cercus, right lateral. 3.151, A. usa, head, dorsal. 3.152, Alloperla furcula, right cercus, right lateral. 3.153, A. furcula, head, dorsal. 3.154, A. furcula, left foreleg, anterodorsal. 3.155, A. imbecilla, left foreleg, anterodorsal. 3.156, A. furcula, abdominal terga 6–7, dorsal. Figures 3.148–3.149 redrawn from Harper & Hynes (1971b); all other figures original.

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67(64’) 67’

Posterior pronotal fringe consisting of two rather widely spaced long setae at corners (Fig. 3.157) ....................................................................................................................... 68 Posterior pronotal fringe consisting of 1 or 2 short setae between longer ones at corners (Fig. 3.160) ....................................................................................................................... 69

68(67) 68’

Vertical cercal fringe beginning on segment 9 or 10 (Fig. 3.158); fewer than 10 outer marginal setae on mesonotum (Fig. 3.159); likely predators/ engulfers; hyporheic for most of larval development; KY, MD, NY, OH, PA, VA, WV............. Alloperla imbecilla Vertical cercal fringe beginning on segment 12; at least 10 outer marginal setae on mesonotum (Fig. 3.162); likely predators/engulfers; hyporheic for most of larval development; AL, CT, GA, MA, MD, ME, MI, MN, NB, NC, NH, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, VT ............................................................... Alloperla atlantica

69(67’)



Mid-fringe setae on posterior margin of abdominal terga 8 and 9 similar to those just laterad to innermost long thin setae (Fig. 3.161); mesonotum with about 8 outer marginal setae; vertical cercal fringe beginning on segment 9; likely predators/engulfers; hyporheic for most of larval development; GA, MA, ME, NB, NC, NF, NH, NL, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WV . ............................................................. Alloperla petasata Mid-fringe setae on posterior margin of abdominal terga 8 and 9 much thinner and more widely spaced than those laterad to innermost pair of long thin setae (Fig. 3.163); mesonotum with about 4, or about 12 outer marginal setae; vertical cercal setal fringe beginning beyond segment 10 .......................................................................................... 70

70(69’) 70’

Vertical cercal setal fringe beginning on segment 11; mesonotum with about 12 outer marginal setae; likely predators/engulfers; hyporheic for most of larval development; CT, MA, ME, NB, NH, NL, NS, NY, ON, PA, QC, VT, WV . .............. Alloperla concolor Vertical cercal setal fringe beginning on segment 13; mesonotum with about 4 outer marginal setae; likely predators/engulfers; hyporheic for most of larval development; NC, TN, VA ........................................................................................... Alloperla neglecta

69’

CHLOROPERLIDAE, Haploperla

(Characters are extracted from Brown & Stark, 1995; Fiance, 1977; Stewart & Stark, 2002. Suitable characters are available for two of the four regional species.) 71(15’)

71’

Pronotum mostly pale brown, but with a pale median band (Fig. 3.164); ventral margin of fore femora usually with 2 or 3 long setae (Fig. 3.166); predators/engulfers; clingers in gravel and organic substrate; widely distributed and often common; AB, AL, AR, BC, CT, DE, GA, IL, IN, KY, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NH, NJ, NS, NY, OH, OK, ON, PA, PE, QC, SC, SK, TN, VA, VT, WI, WV ................................................................................................................. Haploperla brevis Pronotum pale brown along median suture and paler on central disk and near lateral margins (Fig. 3.165); ventral margin of fore femora with several long setae (Fig. 3.167); predators/engulfers; clingers on gravel in first-order to third- order streams; known only from southwest MS . ........................................................................... Haploperla chukcho

Chapter 3 ~ Key to Stoneflies

Page 205

Fig. 3.158

Fig. 3.157 Fig. 3.159

Fig. 3.160

Fig. 3.161

Fig. 3.164

Fig. 3.163

Fig. 3.166

Fig. 3.162

Fig. 3.165

Fig. 3.167

Figures 3.157–3.167, Alloperla spp. and Haploperla spp. (Chloroperlidae). 3.157, Alloperla imbecilla, left half of pronotum, dorsal. 3.158, A. imbecilla, right cercus, right lateral. 3.159, A. imbecilla, left half of mesonotum, dorsal. 3.160, A. concolor, left half of pronotum, dorsal. 3.161, A. petasata, abdominal terga 8–9, dorsal. 3.162, A. atlantica, left half of mesonotum, dorsal. 3.163, A. neglecta, abdominal terga 8–9, dorsal. 3.164, Haploperla brevis, left half of pronotum, dorsal. 3.165, H. chukcho, left half of pronotum, dorsal. 3.166, H. brevis, left foreleg, anterodorsal. 3.167, H. chukcho, left foreleg, anterodorsal. All figures original.

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CHLOROPERLIDAE, Sweltsa

(After Stark et al., 2011. Suitable characters are available for eight of the 10 species known from the region and adjacent states. Only larvae of S. holstonensis [NC, VA] and S. voshelli [GA, NC, VA] are undescribed in Region IV. Sample size for some species is small and the key should be considered provisional.) 72(13) 72’

Tibial fringe setae sparse and about half as long as median outer marginal seta (Fig. 3.168) ....................................................................................................................... 73 Tibial fringe setae more abundant and about as long as median outer marginal seta (Fig. 3.171) ....................................................................................................................... 77

73(72) 73’

Metanotal wingpads with fewer than five intercalary setae (Fig. 3.169) . .............................. 74 Metanotal wingpads with at least 10 intercalary setae (Fig. 3.172) ....................................... 75

74(73) 74’

Tergum 10 without intercalary setae (Fig. 3.173); likely predators/ engulfers; clingers in small to medium-sized creeks, primarily in upper Potomac River and North and South Forks of Shenandoah River drainages; MD, PA, VA, WV ...................... Sweltsa palearata Tergum 10 usually with a pair of intercalary setae (Fig. 3.170); likely predators/engulfers; clingers in creeks and small rivers of the unglaciated region of New York, through the Interior Plateau in Indiana and south to northwestern Alabama; AL, IN, KY, NY, OH, PA, TN, WV . ........................................................................................... Sweltsa hoffmani

75(73’) 75’

Tergum 10 with about 10 to 12 intercalary setae in two lateral clusters (Fig. 3.174); likely predators/engulfers; clingers in higher elevation rheocrenes and small streams of western North Carolina and southwestern Virginia; NC, VA ...................... Sweltsa urticae Tergum 10 with, at most, two intercalary setae (Fig. 3.176) .................................................. 76

76(75’)

Longest dorsal setae on last five cercal segments subequal to two cercal segments in length (Fig. 3.175); likely predators/engulfers; clingers in headwater seeps, rheocrenes and small streams of Appalachians; CT, GA, MA, MD, ME, NB, NC, NH, NY, OH, PA, QC, SC, TN, VA, VT, WV ........................................................................ Sweltsa lateralis Longest dorsal setae on last five cercal segments about as long as 1.5 cercal segments (Fig. 3.177); likely predators/engulfers; clingers generally in headwater seeps and rheocrenes of the Ridge and Valley Province and Allegheny Plateau; MD, VA .............................................................................................................. Sweltsa pocahontas

76’ 77(72’) 77’

Longest dorsal setae on last five cercal segments about as long as 1.2-1.5 cercal segments (Fig. 3.178); likely predators/engulfers; clingers in creeks of the upper Tennessee River drainage southward through the mountains of the Carolinas, Tennessee and Virginia; AL, NC, SC, TN, VA ............................................................................... Sweltsa mediana Longest dorsal setae on last five cercal segments about as long as 2 cercal segments (Fig. 3.175) ....................................................................................................................... 78

Chapter 3 ~ Key to Stoneflies

Fig. 3.168

Page 207

Fig. 3.169

Fig. 3.170

Fig. 3.172

Fig. 3.171

Fig. 3.173

Fig. 3.175 Fig. 3.174

Fig. 3.177

Fig. 3.176

Fig. 3.178

Figures 3.168–3.178, Sweltsa spp. (Chloroperlidae). 3.168, Sweltsa hoffmani, left foreleg, anterodorsal. 3.169, S. hoffmani, left halves of meso- and metanota, dorsal. 3.170, S. hoffmani, abdominal terga 8–10, dorsal. 3.171, S. onkos, left foreleg, anterodorsal. 3.172, S. urticae, left halves of meso- and metanota, dorsal. 3.173, S. palearata, abdominal terga 8–10, dorsal. 3.174, S. urticae, abdominal terga 8–10, dorsal. 3.175, S. lateralis, apex of right cercus, right lateral. 3.176, S. hoffmani, abdominal terga 8–10, dorsal. 3.177, S. pocahontas, apex of right cercus, right lateral. 3.178, S. mediana, apex of right cercus, right lateral. All figures original.

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78(77’)

Abdominal terga dark brown (Fig. 3.180), occiput about as dark as frons (Fig. 3.179); meso- and metanota rather densely covered with long dark clothing hairs; likely predators/engulfers; clingers; CT, DE, IN, KY, MA, MD, ME, NB, NH, NL, NS, NY, OH, ON, PA, PE, QC, VA, VT, WV ............................................................. Sweltsa onkos Abdominal terga pale brown, occiput almost entirely pale (Fig. 3.181); meso- and metanota with sparse, brown clothing hairs; likely predators/engulfers; clingers; ME, NB, NH, NL, NS, NY, PA, PE, QC, VA, VT, WV ........................................................ Sweltsa naica

78’

LEUCTRIDAE, Leuctra

(After Harper & Hynes, 1971a; characters are extracted from Harrison & Stark, 2010; Stewart & Stark, 2002. Suitable characters are available for eight of the 25 regional species.) 79(18’) 79’

Cercal segments bearing prominent, long, bushy whorls consisting of 8 or more lateral setae on most segments (Fig. 3.182); likely shredders/detritivores; clingers in late instars; CT, DE, IL, IN, KY, MA, MD, ME, NB, NC, NY, OH, ON, PA, QC, TN, VA, WI, WV ............................................................................................................................ Leuctra sibleyi Cercal segments with less prominent whorls, usually consisting of 5 or fewer lateral setae on most segments (Fig. 3.184) . ........................................................................................ 80

80(79’) 80’

Abdominal terga 7 to 10 or 8 to 10 covered with short, stout hairs, more anterior terga with hairs reduced; hairs most conspicuous in lateral aspect (Fig. 3.183) ............................... 81 All abdominal terga covered with short, stout hairs ............................................................... 82

81(80) 81’

Mesonotum with numerous short, stout setae on anterolateral angles (Fig. 3.185); likely shredders/detritivores; clingers in late instars; CT, MA, MD, ME, MN, MS, NB, NJ, NL, NS, NY, ON, PA, QC, WI, WV ............................................................ Leuctra tenella Mesonotum with few short, stout setae on anterolateral angles (Fig. 3.187); likely shredders/ detritivores; clingers in late instars; CT, ME, NL, NY, PA, QC, VA, WV . ..................... ................................................................................................................... Leuctra truncata

82(80’) 82’

Short, stout setae absent on abdominal sterna, although one or more long setae may occur on posterior margins (Fig. 3.186) ..................................................................................... 83 Short, stout setae on at least sternum 9 (Fig. 3.189) ............................................................... 84

83(82)

A single long, or moderately long, seta present on posterior margin of at least abdominal sterna 7 to 9 (Fig. 3.188); likely shredders/detritivores; clingers in late instars; AL, CT, DE, FL, KY, MA, MD, ME, MI, MN, MS, NB, NC, NF, NS, NY, OH, PA, QC, SC, SK, VA, WI, WV . ................................................................................. Leuctra ferruginea A single long, or moderately long, seta present only on sternum 9 (Fig. 3.186); likely shredders/detritivores; clingers in late instars in spring seeps; MS ...... Leuctra colemanorum

83’

Chapter 3 ~ Key to Stoneflies

Page 209

Fig. 3.179

Fig. 3.180

Fig. 3.182

Fig. 3.181

Fig. 3.184

Fig. 3.183

Fig. 3.185

Fig. 3.186

Fig. 3.187

Fig. 3.188

Fig. 3.189

Figures 3.179–3.189, Sweltsa spp. (Chloroperlidae) and Leuctra spp. (Leuctridae). 3.179, Sweltsa onkos Chloroperlidae), head, dorsal. 3.180, S. onkos, abdominal terga 8–10, dorsal. 3.181, S. naica, head, dorsal. 3.182, Leuctra sibleyi (Leuctridae), basal segments of left cercus, left lateral. 3.183, L. tenella, abdominal segments 8–10, right lateral. 3.184, L. tenella, basal segments of left cercus, left lateral. 3.185, L. tenella, left half of mesonotum and wingpad, dorsal. 3.186, L. colemanorum, abdominal segments 8–10, right lateral. 3.187, L. truncata, left half of mesonotum and wingpad, dorsal. 3.188, L. ferruginea, abdominal segments 8–10, right lateral. 3.189, Leuctra tenuis, abdominal segments 8–10, right lateral. Figures 3.182, 3.185, 3.187-3.189 redrawn from Harper & Hynes (1971a); all other figures original.

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84(82’)

Short, stout setae abundant only on sternum 9, preceding segments may have a few (Fig. 3.189); likely shredders/detritivores; clingers in late instars; AL, AR, CT, DE, IA, IL, IN, KY, LA, MA, MD, ME, MI, MN, MO, MS, NB, NC, NJ, NL, NS, NY, OH, OK, PA, QC, VA, WI, WV . .......................................................................... Leuctra tenuis Abundant short, stout setae present on at least abdominal sterna 4 to 9 (Fig. 3.190) . ........... 85

84’ 85(84’) 85’

Posterolateral pronotal corner with several short marginal setae (Fig. 3.191); lateral margin of mesonotal wingpad bearing several short, stout setae (Fig. 3.191); likely shredders/ detritivores; clingers in late instars; CT, MD, ME, NB, NS, NY, ON, PA, PE, QC, VA,WV . .................................................................................................. Leuctra duplicata Posterolateral pronotal corner with at least one long seta (Fig. 3.192); lateral mesonotal wingpad margin with few, if any, short, stout setae (Fig. 3.192); likely shredders/ detritivores; clingers in late instars; CT, ME, NH, ON, PA, QC, VT, WV...... Leuctra maria

LEUCTRIDAE, Megaleuctra

(Larvae of the two eastern species are undescribed but can be separated on a geographical basis.) 86(16) 86’

Known from the Carolinas, north Georgia and the Mt. Rogers area of southern Virginia; likely detritivores in small spring seeps; GA, NC, SC, TN, VA ..................................... ....................................................................................................... Megaleuctra williamsae Known from West Virginia, Pennsylvania and the Shenandoah National Park of Virginia; likely detritivores in small spring seeps; MD, PA, VA, WV . ................. Megaleuctra flinti

LEUCTRIDAE, Zealeuctra

(Characters are extracted from Stewart & Stark, 2002. Suitable characters are available for one of the four regional species. See Grubbs et al., 2013 for an overview of the distributions of these species in EPA Region IV.) 87(17)

Paraprocts setose and partially fused (Fig. 3.193); longest setae on apical cercal segments about half as long as segments (Fig. 3.194); likely shredders /detritivores; late instars clingers; AL, AR, IL, IN, KS, KY, MO, OK, TN, TX, WV .............. Zealeuctra claasseni

Chapter 3 ~ Key to Stoneflies

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Fig. 3.190

Fig. 3.191

Fig. 3.192

Fig. 3.193

Fig. 3.194 Figures 3.190–3.194, Leuctra spp. and Zealeuctra claasseni (Leuctridae). 3.190, Leuctra duplicata, abdominal segments 8–10, right lateral. 3.191, L. duplicata, left halves of pro- and mesonota, dorsal. 3.192, L. maria, left halves of pro- and mesonota, dorsal. 3.193, Zealeuctra claasseni, abdominal sterna 8–10, ventral. 3.194, Z. claasseni, apical segments of right cercus, right lateral. Figures 3.190–3.192 redrawn from Harper & Hynes (1971a); figures 3.193–3.194 redrawn from Stewart & Stark (2002).

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NEMOURIDAE, Amphinemura

(After Harper & Hynes, 1971d; characters are extracted from Nelson, 1997; Stewart & Stark, 2002; Stark & Harrison, 2010. Suitable characters are available for five of the seven regional species. Only the larvae of A. appalachia [GA, NC, PA, SC, TN, VA] and A. varshava [IA, IL, IN, KY, WI] are unknown in Region IV.) 88(20’) 88’

Outer (anterior) gill tufts containing about half as many filaments as inner (posterior) tufts (Fig. 3.196); lateral margins of pronotum fringed with short, thick setae (Fig. 3.195); likely shredders/detritivores; clingers; CT, DE, GA, KY, MA, MD, ME, NB, NC, NJ, NS, NY, PA, PE, QC, SC, TN, VA, WV . .............................................. Amphinemura wui Outer gill tufts containing about as many filaments as inner tufts; lateral pronotal fringe consisting mostly of long setae (Fig. 3.197) ..................................................................... 89

89(88’) 89’

Longest setae in whorls at mid-cercal length usually less than half as long as segment (Fig. 3.198); likely shreddesr/detritivores; clingers; known only from TN ............................. ........................................................................................................... Amphinemura mockfordi Longest setae in whorls at mid-cercal length at least half as long as segment (Fig. 3.200) . ..... 90

90(89’) 90’

Longest setae in whorls at mid-cercal length about half as long as segment (Fig. 3.199); likely shredders/detritivores; clingers; AL, AR, GA, IA, IL, IN, KS, KY, MI, MO, NC, OH, OK, ON, PA, QC, TN, VA, WI, WV ........................................ Amphinemura delosa Longest setae in whorls at mid-cercal length at least ¾ as long as segment (Fig. 3.200) ...... 91

91(90’) 91’

Longest setae in whorls at mid-cercal length about as long as segment (Fig. 3.201); likely shredders/detritivores; clingers; AL, KY, MS ............................. Amphinemura alabama Longest setae in whorls at mid-cercal length about ¾ as long as segment (Fig. 3.202); likely shredders/detritivores; clingers; AL, AR, CT, DE, FL, GA, IL, IN, KY, MD, ME, MI, MO, MS, NB, NC, NL, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, WV ......................................................................................................... Amphinemura nigritta



NEMOURIDAE, Ostrocerca

(After Harper & Hynes, 1971d; characters are extracted from Stewart & Stark, 2002. Suitable characters are available for two of the four regional species.) 92(24) 92’

Developing epiproct on male tergum 10 about as wide as long (Fig. 3.203); most abdominal setae darker than tergal pigment and relatively conspicuous; cerci with inconspicuous pigment bands; likely shredders/ detritivores; clingers in first-order rheocrenes and springs; CT, MA, MD, ME, MI, NC, NH, NS, NY, ON, PA, QC, VA, WV ....................................................................................................... Ostrocerca albidipennis Developing epiproct on male tergum 10 longer than wide (Fig. 3.204); abdominal setae pale, relatively inconspicuous; cerci without bands; likely shredders/detritivores; clingers in first-order rheocrenes and springs; CT, IN, KY, MA, MD, ME, NC, NY, OH, ON, PA, QC, VA, WV ........................................................................................ Ostrocerca truncata

Chapter 3 ~ Key to Stoneflies

Page 213

Fig. 3.195

Fig. 3.196

Fig. 3.198

Fig. 3.197

Fig. 3.199

Fig. 3.200

Fig. 3.201

Fig. 3.202

Fig. 3.203

Fig. 3.204

Figures 3.195–3.204, Amphinemura spp. and Ostrocerca spp. (Nemouridae). 3.195, Amphinemura wui, left half of pronotum, dorsal. 3.196, A. wui, cervical gills, ventral. 3.197, A. alabama, left half of pronotum, dorsal. 3.198, A. mockfordi, segments of left cercus, left lateral. 3.199, A. delosa, segments of left cercus, left lateral. 3.200, A. nigritta, segments of left cercus, left lateral. 3.201, A. alabama, apical segments of left cercus, left lateral. 3.202, A. nigritta, apical segments of left cercus, left lateral. 3.203, Ostrocerca albidipennis, abdominal terga 8–10 of developing male nymph, dorsal. 3.204, O. truncata, abdominal terga 8–10 of developing male nymph, dorsal. Figure 3.198 redrawn from Nelson (1997); all other figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

NEMOURIDAE, Prostoia

(After Harper & Hynes, 1971d; characters are extracted from Stewart & Stark, 2002. Suitable characters are available for the three regional species. Slide mounts may be needed to observe intersegmental setae.) 93(23’) 93’

Intersegmental hairs on cercal segments poorly developed, or absent, on most segments (Fig. 3.205); likely shredders/detritivores; clingers; CT, DE, IL, IN, KY, MA, MD, ME, MI, MN, MO, NY, OH, ON, PA, QC, SC, VA, WI, WV . .................. Prostoia similis Intersegmental hairs on cercal segments present and well developed on several apical and middle segments (Fig. 3.207) ........................................................................................... 94

94(93’) 94’

Outer marginal tibial fringe poorly developed, comprising a few short setae (Fig. 3.206); outer marginal mesonotal setae absent between base and midlength of segment; likely shredders/detritivores; clingers; CT, GA, IL, MA, NC, VA . ................. ..................................................................................................................... Prostoia hallasi Outer marginal tibial fringe well developed, comprising mostly setae longer than tibial width (Fig. 3.208); outer marginal mesonotal setae scattered along entire length of segment (Fig. 3.209); likely shredders/detritivores; clingers; AL, DE, IA, IN, KY, MA, MD, ME, MI, MN, MS, NB, NC, NL, NS, OH, ON, PA, PE, QC, SC, TN, VA, WI, WV ................................................................................................................. Prostoia completa



NEMOURIDAE, Soyedina

(Characters are extracted from Harper & Hynes, 1971d; Stewart & Stark, 2002. Suitable characters are available for two of the six regional species.) 95(21) 95’

Most basal to mid-cercal segments with a few thick, intermediate setae similar to those in the apical segmental whorls (Fig. 3.210); likely shredders/detritivores; clingers; CT, IA, IL, IN, KY, MA, MD, ME, MI, NS, NY, OH, ON, PA, QC, TN, VA, WI, WV ............................................................................................................ Soyedina vallicularia Basal to mid-cercal segments without thick intermediate setae; likely shredders/detritivores; clingers; DE, MD, NC, PA, TN, VA, WV ............................................. Soyedina carolinensis

NEMOURIDAE, Zapada

(Characters are extracted from Grubbs et al., 2015; Ricker, 1952. Larvae of Zapada “species A” a species being described by Grubbs et al., 2015, are unknown, but the distinctive cervical gills of the adult also are expected to occur in the larvae and to permit the two species known from EPA Region IV to be distinguished.) 96(20) 96’

Cervical area bearing four unbranched tubular gills (Fig. 3.211); likely shredders/ detritivores; clingers; TN . .............................................................................. Zapada chila Cervical area bearing gills with three branches and a vestigial, lateral branch; likely shredders/detritivores; clingers; NC, TN, VA . ........................................ Zapada species A

Chapter 3 ~ Key to Stoneflies

Page 215

Fig. 3.205 Fig. 3.206

Fig. 3.207

Fig. 3.208

Fig. 3.209

Fig. 3.210

Fig. 3.211

Figures 3.205–3.211, Nemouridae genn. spp. 3.205, Prostoia similis, apical segments of right cercus, right lateral. 3.206, P. hallasi, left foreleg, anterodorsal. 3.207, P. completa, apical segments of right cercus, right lateral. 3.208, P. completa, left foreleg, anterodorsal. 3.209, P. completa, left halves of meso- and metanota, dorsal. 3.210, Soyedina vallicularia, apical segments of right cercus, right lateral. 3.211, Zapada chila, cervical gills, ventral. All figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

PERLIDAE, Acroneuria

(After Hitchcock, 1974; characters are extracted from Kondratieff & Kirchner, 1988; Stark & Kondratieff, 2004; Stewart & Stark, 2002. Characters based primarily on pigment patterns are available for 11 of the 16 regional species. These and the anal-gill character should be used with caution.) 97(36’) 97’

Anal gills present on paraprocts (Fig. 3.212) . ........................................................................ 98 Anal gills absent from paraprocts ......................................................................................... 105

98(97) 98’

M-line on head interrupted by dark pigment separating pale band into three pale spots (Fig. 3.213), or head almost completely dark ................................................................... 99 M-line on head complete (Fig. 3.214) .................................................................................. 100

99(98)

Abdominal terga without distinctive pigment bands; predators/engulfers; clingers; AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, NC, OH, OK, PA, TN, TX ...... Acroneuria evoluta

99’

Most abdominal terga with an incomplete, narrow, pale basal band (Fig. 3.218); predators/ engulfers; clingers; AL, CT, DC, DE, FL, GA, LA, MA, MD, ME, MS, NC, NJ, NY, PA, QC, SC, TX, VA, WV.................................................................... Acroneuria arenosa

100(98’) Abdominal terga with broad, pale basal and narrow, dark apical pigment bands (Fig. 3.215); predators/engulfers; clingers; CT, KY, IA, MB, ME, MI, MN, NB, NC, ND, NJ, NS, NY, OH, ON, PA, QC, SK, TN, VA, WI, WV ............................................ Acroneuria lycorias 100’ Abdominal terga typically with dark basal and pale apical pigment bands (Fig. 3.216) . .... 101 101(100’) Dark abdominal tergal bands approximately equal in width across segments (Fig. 3.216) . ....... 102 101’ Dark abdominal tergal bands slightly expanded near midlength and often notched at midlength (Fig. 3.217) .................................................................................................... 103 102(101) Tergum 10 apex with a small dark area and dark basal band usually divided; posterior tergal setal row set on narrow dark pigment band (Fig. 3.219); predators/engulfers; clingers; known from KY ............................................................................... Acroneuria hitchcocki 102’ Apex of tergum 10 pale, dark basal band complete; posterior tergal setal row set on pale pigment (Fig. 3.216); predators/engulfers; clingers; AL, AR, IL, IN, KS, KY, MA, MD, MI, MO, NC, OH, OK, ON, PA, TN, VA, WV ............................ Acroneuria frisoni 103(101’) Dark abdominal tergal bands strongly narrowed laterally on segments 8 and 9 (Fig. 3.217); predators/engulfers; clingers; AL, AR, DC, GA, IL, IN, KY, MO, OH, OK, PA, TN, WV ............................................................................................................ Acroneuria perplexa 103’ Dark abdominal tergal bands cover about half of segments 8 and 9 near lateral margins (Fig. 3.216) ..................................................................................................................... 104

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Fig. 3.212

Fig. 3.213

Fig. 3.215

Fig. 3.214

Fig. 3.216

Fig. 3.218

Fig. 3.217

Fig. 3.219

Figures 3.212–3.219, Agnetina capatata and Acroneuria spp. (Perlidae). 3.212, Agnetina capitata, anal gills, ventral. 3.213, Acroneuria arenosa, head, dorsal. 3.214, A. lycorias, head, dorsal. 3.215, A. lycorias, abdominal terga 8–10, dorsal. 3.216, A. frisoni, abdominal terga 8–10, dorsal. 3.217, A. perplexa, abdominal terga 8–10, dorsal. 3.218, A. arenosa, abdominal terga 8–10, dorsal. 3.219, A. hitchcocki, abdominal terga 8–10, dorsal. All figures original.

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104(103’) Median expansions of dark bands on abdominal terga 8 and 9 only slightly wider than lateral width; predators/engulfers; clingers; KY, PA, VA . ............... Acroneuria kirchneri 104’ Median expansions of dark bands on abdominal terga extending to, or near, posterior margins of some segments (Fig. 3.220); predators/engulfers; clingers; AL, AR, GA, IL, IN, KY, MD, MO, NC, OH, PA, SC, TN, VA, WV . .............................. Acroneuria filicis 105(97’) Abdominal terga uniformly brown, without distinctive pale banding; predators/engulfers; clingers; AB, AL, CO, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MS, MT, NB, NC, ND, NE, NL, NM, NY, OH, ON, PA, QC, SC, SD, SK, TN, UT, VA, WI, WV, WY ............................................................... Acroneuria abnormis 105’ Abdominal terga banded with pale and dark pigment .......................................................... 106 106(105’) Abdominal terga 8 and 9 with dark median or basal bands and pale, narrow, apical bands (Fig. 3.221); predators/engulfers; clingers; AR, IL, IN, KY, MI, MN, MO, OK, VA, WI, WV . ............................................................................................ Acroneuria internata 106’ Abdominal terga with pale basal and dark apical bands (Fig. 3.222); some southern populations have sparse anal gills and key to A. lycorias; predators/engulfers; clingers; AL, CT, GA, KY, MA, MB, MD, ME, MS, NC, NH, NJ, NY, OH, ON, PA, QC, SC, TN, VA, WV . ............................................................................................. Acroneuria carolinensis

PERLIDAE, Agnetina

(From Stark, 1986; All three regional species are included in the following key; however, sample size is small and identifications should be confirmed with adult males when possible.) 107(30’) Most of apex of tergum 10 dark (Fig. 3.223); predators/engulfers; clingers; AL, DC, FL, IN, LA, MD, MS, PA, SC, VA, WV .................................................... Agnetina annulipes 107’ Apex of tergum 10 pale (Fig. 3.225) .................................................................................... 108 108(107’) Arms of M-line on head directed essentially laterad (Fig. 3.224); dark pigment of tergum 10 usually continuous, narrowed over most of tergum 9 (Fig. 3.225); predators/ engulfers; clingers; AR, CT, IA, KY, MA, MD, ME, MI, MN, MO, NB, NC, NH, NS, NY, OH, ON, PA, PE, QC, VA, WI, WV ................................................ Agnetina capitata 108’ Arms of M-line on head directed posterolaterally (Fig. 3.226); dark pigment of tergum 10, when present, usually divided mesally, but covering most of tergum 9 (Fig. 3.227); predators/engulfers; clingers; AL, AR, GA, IL, IN, KY, MD, MI, MN, MO, NC, NY, OH, OK, PA, SC, TN, VA, WV . ......................................................... Agnetina flavescens

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Fig. 3.220

Fig. 3.223

Fig. 3.226

Fig. 3.221

Fig. 3.222

Fig. 3.224

Fig. 3.225

Fig. 3.227

Figures 3.220–3.227, Acroneuria spp. and Agnetina spp. (Perlidae). 3.220, Acroneuria filicis, abdominal terga 8–10, dorsal. 3.221, A. internata, abdominal terga 8–10, dorsal. 3.222, A. carolinensis, abdominal terga 8–10, dorsal. 3.223, Agnetina annulipes, abdominal terga 8–10, dorsal. 3.224, A. capitata, head, dorsal. 3.225, A. capitata, abdominal terga 8–10, dorsal. 3.226, A. flavescens, head, dorsal. 3.227, A. flavescens, abdominal terga 8–10, dorsal. All figures original.

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PERLIDAE, Beloneuria

(Characters are extracted from Stewart & Stark, 2002; Stark, unpublished. All three regional species are included in the following key; however, sample size is small and identifications should be confirmed when possible by examining eggs from gravid females.) 109(36) 109’

Known from mountain streams in eastern Alabama; predators/engulfers; clingers; AL .................................................................................................................... Beloneuria jamesae Known from sites north and east of Alabama . ..................................................................... 110

110(109’) Thick setae on lower half of hind femora distributed fairly evenly over much of basal three fourths of area (Fig. 3.228); predators/engulfers; clingers usually in lower elevation and piedmont streams; GA, NC, SC, TN . .......................................... Beloneuria stewarti 110’ Thick setae on lower half of hind femora less abundant, or absent beyond apical third of femoral length, except along margins (Fig. 3.229); predators/engulfers; clingers usually in higher elevation first-order and second-order seeps and streams; GA, NC .......................................................................................................... Beloneuria georgiana

PERLIDAE, Hansonoperla

(Characters are extracted from Kirchner & Kondratieff, 1985; Stewart & Stark, 2002. Suitable characters are available for only one of the three regional species, but a second species may be endemic to the Cheaha Mountain area of eastern Alabama and is separated on that basis.) 111(33’) Known from sites north and east of Alabama; anterior angles of pronotum bearing 2 or 3 long setae, lateral margins without setae (Fig. 3.230); tibiae without ventral silky setal fringe; likely predators/engulfers; early instars may occur in hyporhea, pre-emergent specimens found among roots, detritus, and bank undercuts and under large boulders in pristine streams; MA, NC, NH, PA, SC, TN, VA, WV ............. Hansonoperla appalachia 111’ Known from eastern Alabama; no larval specimens available; AL . ..... Hansonoperla cheaha

PERLIDAE, Neoperla

(After Poulton & Stewart, 1991; characters are extracted from Stark, 1995; Stewart & Stark, 2002. Characters based on subtle variations in pigment pattern are available for only four of the eleven regional species. Sample size for associated material is small; consequently, adult specimens are needed to confirm identifications.) 112(28) 112’

Dark transverse band on frons reaching lateral margins of head between eye and antennal base (Fig. 3.232) ............................................................................................................. 113 Dark transverse band on frons reaching lateral margins of head anterior to antennal base (Fig. 3.231) ..................................................................................................................... 114

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Fig. 3.228

Fig. 3.230

Fig. 3.229

Fig. 3.231

Fig. 3.232

Figures 3.228–3.232, Perlidae genn. spp. 3.228, Beloneuria stewarti, left hind femur, anterodorsal. 3.229, B. georgiana, left hind femur, anterodorsal. 3.230, Hansonoperla appalachia, head and pronotum, dorsal. 3.231, Neoperla clymene, head and pronotum, dorsal. 3.232, N. choctaw, head and pronotum, dorsal. All figures original.

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113(112) Dark pigment reaching anterolateral margins of head enclosing pale frontoclypeal pigment (Fig. 3.232); likely predators/engulfers; clingers; AR, KY, MO, OK, PA, WV ................................................................................................................. Neoperla choctaw 113’ Dark pigment not enclosing pale frontoclypeal pigment; likely predators/ engulfers; clingers; AL, AR, IL, IN, KY, MO, OH, OK, PA, SC, TX, VA, WV .... Neoperla catharae 114(112’) Dark pigment bands on abdominal terga 8 and 9 narrow (Fig. 3.234); dark pigment along posterior margin of pronotum very thin at median suture (Fig. 3.233); likely predators/ engulfers; clingers; MS . ................................................................................ Neoperla coxi 114’ Dark pigment bands on abdominal terga wide (Fig. 3.236); dark pigment on posterior margin of pronotum somewhat expanded near median suture (Fig. 3.235); likely predators/engulfers; clingers; AL, FL, GA, IA, IL, IN, KY, LA, MS, NC, OK, PA, SC, TX, VA, WV .................................................................................... Neoperla clymene or, AR, IL, MO, MS, OK, PA, TN, WI .................................................. Neoperla robisoni

PERLIDAE, Paragnetina

(After Stark & Szczytko, 1981. Suitable characters are available for all regional species.) 115(30) 115’

Femora distinctively patterned in yellow and dark brown (Fig. 3.237) ............................... 116 Femora almost entirely brown, without distinctive pattern .................................................. 118

116(115) Mid-femora with dark longitudinal band extending along two thirds of anteromedian surface (Fig. 3.237); M-line usually complete on head (Fig. 3.238); anal gills absent; predators/engulfers; clingers; CT, DE, GA, KY, MA, ME, NC, NH, NY, PA, QC, SC, TN, VA, WV .............................................................................. Paragnetina immarginata 116’ Mid-femora without dark longitudinal band, but transverse dark bands may be present (Fig. 3.239); M-line on head incomplete, may be reduced to 1 to 3 pale pigment spots near median ocellus (Fig. 3.242); anal gills present or absent . ...................................... 117 117(116’) M-line usually consisting of median and two lateral pale spots, often with a pale insertion from clypeal band extending to median ocellus (Fig. 3.240); several abdominal terga with extensive pale markings (Fig. 3.241); predators/engulfers; clingers, common, often abundant; AL, DC, FL, GA, LA, MS, NC, PA, SC, TX, VA ..... Paragnetina fumosa 117’ M-line usually consisting of pale spot around median ocellus (Fig. 3.242); abdominal terga typically with reduced pale markings (Fig. 3.243); predators/engulfers; clingers; AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, NC, SC............ Paragnetina kansensis (in part) 118(115’) Abdominal terga 4 to 9 typically with at least one or two short, thick intercalary setae similar to those in segmental whorls (Fig. 3.243); sparse anal gills usually present on paraprocts; predators/engulfers; clingers; AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, NC, SC ............................................................................ Paragnetina kansensis (in part) 118’ Abdominal terga 4 to 9 typically without short thick intercalary setae, but numerous short, thin setae are present in posterior segmental fringes (Fig. 3.244); anal gills usually absent from paraprocts ............................................................................................................... 119

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Fig. 3.233

Fig. 3.234

Fig. 3.235

Fig. 3.236

Fig. 3.237

Fig. 3.238

Fig. 3.239

Fig. 3.240

Fig. 3.241

Fig. 3.242

Fig. 3.243

Fig. 3.244

Figures 3.233–3.244, Neoperla spp. and Paragnetina spp. (Perlidae). 3.233, Neoperla coxi, head and pronotum, dorsal. 3.234, N. coxi, abdominal terga 8–10, dorsal. 3.235, N. clymene, head and pronotum, dorsal. 3.236, N. clymene, abdominal terga 9–10, dorsal. 3.237, Paragnetina immarginata, left forefemur, anterodorsal. 3.238, P. immarginata, head, dorsal. 3.239, P. fumosa, left midfemur, anterodorsal. 3.240, P. fumosa, head, dorsal. 3.241, P. fumosa, abdominal terga 8–10, dorsal. 3.242, P. kansensis, head, dorsal. 3.243, P. kansensis, abdominal terga 8–10, dorsal. 3.244, P. media, abdominal terga 8–10, dorsal. Figures 3.233–3.234 redrawn from Stark (1995); all other figures original.

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119(118’) Known only south of Virginia, West Virginia and Kentucky but sometimes overlapping P. media in North Carolina; egg chorion from pre-emergent larvae or emergent females smooth and collar very wide; predators/engulfers; clingers in small Appalachian rivers; NC, SC, TN, VA .................................................................................. Paragnetina ichusa 119’ Known generally from north of North Carolina and Tennessee; egg chorion from pre emergent larvae or emergent females ornate and collar narrow; predators/engulfers; clingers in medium-sized to large creeks and rivers; AR, CT, DE, IA, IL, IN, KY, MB, MD, ME, MI, MN, MO, NB, NH, NY, OH, ON, PA, QC, SK, VA, WI, WV ............................................................................................................... Paragnetina media

PERLIDAE, Perlesta

(Characters are extracted from Frison, 1942; Kirchner & Kondrateiff, 1997; Poulton & Stewart, 1991; Stark, 1989. Suitable characters are available for six of the seventeen regional species; however, sample size is small and identifications, when possible, should be confirmed with adults.) 120(34) 120’

Abdominal terga with dark, freckle-like spots around bases of intercalary setae (Fig. 3.245) ..................................................................................................................... 121 Abdominal terga without freckled appearance ..................................................................... 123

121(120) Dark transverse pigment band through ocellar area bearing scattered dark spots (Fig. 3.246); mature larvae predators/engulfers; clingers; AR, IA, IL, KY, MI, MO, MS, OK ................................................................................................................... Perlesta shubuta 121’ Dark transverse pigment band on head uniformly colored, without darker spots (Fig. 3.247) ..................................................................................................................... 122 122(121’) Pronotum with a pair of large, pale areas free of clothing hairs; occipital setal row not extending to ecydsial suture (Fig. 3.247); mature larvae predators/ engulfers; clingers; AR, IA, IL, KS, KY, MO, NE, OH, OK, WV ......................................... Perlesta cinctipes 122’ Pronotal clothing hairs rather evenly distributed; occipital setal row extending to ecdysial suture Fig 3.248); mature larvae predators/engulfers; clingers; AR, CO, IA, IL, IN, KY, MI, MO, NC, ND, NE, NM, OH, OK, PA, SD, TX, VA, WI, WV, WY ................................................................................................................. Perlesta decipiens 123(120’) Pronotum with a pair of dark parenthesis-shaped bars (Fig. 3.249); head without pale M-line: mature larvae predators/engulfers; clingers; IL, IN, KY, MD, OH, PA, TN, VA, WV ........................................................................................................................... Perlesta teaysia 123’ Pronotum without dark parenthesis-shaped bars (Fig. 3.251); head with pale M-line, but sometimes incomplete .................................................................................................... 124 124(123’) Most of dark pigment on head restricted to area of frons anterior to lateral ocelli (Fig. 3.251); known from the Appalachians and areas east of the mountains; mature larvae predators/engulfers; clingers; NC, SC, TN, VA, WV . .................... Perlesta frisoni 124’ Dark pigment on head covering most of frons anterior to ecdysial suture (Fig. 3.250); known from the Cumberland Plateau and areas to the north and west; mature larvae predators/ engulfers; clingers; IN, KY, OH, TN .......................................................... Perlesta adena

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Fig. 3.245

Fig. 3.246

Fig. 3.247

Fig. 3.249

Fig. 3.248

Fig. 3.250

Fig. 3.251

Figures 3.245–3.251, Perlesta spp. (Perlidae). 3.245, Perlesta shubuta, abdominal terga 8–10, dorsal. 3.246, P. shubuta, head, dorsal. 3.247, P. cinctipes, head, dorsal. 3.248, P. decipiens, head, dorsal. 3.249, P. teaysia, head and pronotum. 3.250, P. adena, head and pronotum. 3.251, P. frisoni, head and pronotum. Figures 3.245–3.248 redrawn from Stark (1989); figure 3.249 redrawn from Kirchner & Kondratieff (1997); all other figures original.

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PERLIDAE, Perlinella

(After Kondratieff et al., 1988. Suitable characters are available for all regional species.) 125(28’,33) Small anterior ocellus present in mature larvae Fig. 3.252); head and thorax with discernible dark and light pattern; predators/engulfers; early instars in hyporhea; AL, AR, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK, PA, QC, SC, TN, TX, VA, WI, WV ................................................ Perlinella drymo 125’ Anterior ocellus absent (Fig. 3.253); head and thorax brown, without distinct pattern on pronotum .................................................................................................................... 126 126(125’) Body yellow brown, frons rather uniformly pigmented; distribution widespread east of Rocky Mountains; predators/engulfers; early instars in hyporhea; AL, AR, CT, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NJ, NY, OH, OK, PA, RI, SC, TN, VA, WI, WV ................................................................. Perlinella ephyre 126’ Body dark brown; head with areas of dark and paler shading (Fig. 3.254); distribution southeastern U.S.; predators/engulfers; early instars in hyporhea; AL, FL, MS, SC, NC ................................................................................................................... Perlinella zwicki

PERLODIDAE, Cultus

(After Myers & Kondratieff, 2009. Suitable characters are available for both species known from the region.) 127(41’) Pale pronotal pigment area near posterior margin (Fig. 3.255); predators/engulfers; clingers in small Appalachian creeks and rivers; NC, NH, PA, QC, TN, VA, WV ...................... .................................................................................................................... Cultus verticalis 127’ Pale pronotal pigment area more anterior to posterior margin and sometimes diamond shaped (Fig. 3.256); predators/engulfers; clingers in larger creeks and rivers; GA, NC, VA ......................................................................................................... Cultus decisus isolatus

PERLODIDAE, Diploperla

(After Kondratieff et al., 1981. Characters are extracted from Kirchner & Kondratieff, 1984; Stewart & Stark, 2002. Suitable characters are available for all regional species.) 128(41) 128’

Dorsal abdominal terga brown, without distinctive pattern; head usually with pale, sometimes incomplete, M-line (Fig. 3.257); predators/engulfers; common and widespread clingers in small Appalachian, Piedmont, and Coastal Plain streams; AL, DE, GA, MD, MS, NC, PA, SC, TN, VA, WV . ................................ Diploperla duplicata Abdominal terga with distinctive pattern of yellow and brown pigment (Fig. 3.258); head usually without M-line . .................................................................................................. 129

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Fig. 3.252

Fig. 3.254

Fig. 3.253

Fig. 3.256 Fig. 3.255

Fig. 3.257

Fig. 3.258 Figures 3.252–3.258, Perlidae and Perlodidae genn. spp. 3.252, Perlinella drymo (Perlidae), head, dorsal. 3.253, P. ephyre, head and pronotum, dorsal. 3.254, P. zwicki, head, dorsal. 3.255, Cultus verticalis (Perlodidae), head and pronotum, dorsal. 3.256, C. decisus isolatus, pronotum, dorsal. 3.257, Diploperla duplicata (Perlodidae), head and pronotum, dorsal. 3.258, D. robusta, abdominal terga 8–10, dorsal. All figures original.

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129(128’) Hind femora without dark median longitudinal bands (Fig. 3.259); dark pigment bands on abdominal terga usually interrupted medially (Fig. 3.258); predators/engulfers; clingers, rare in streams south of Virginia/Kentucky; AL, CT, IN, KY, OH, PA, TN, VA, WV .................................................................................................................Diploperla robusta 129’ Hind femora with dark median longitudinal pigment bands, although sometimes diffuse (Fig. 3.260); dark pigment on abdominal terga continuous . .......................................... 130 130(129’) Dark pigment in ocellar region reaching ecdysial suture (Fig. 3.261); dark pigment on abdominal terga without posteromedial circular spots; predators/ engulfers; uncommon clingers; NC, VA .................................................................................. Diploperla morgani 130’ Dark pigment in ocellar region not reaching lateral ocelli (Fig. 3.262); dark pigment on abdominal terga includes small posteromedial circular spots; predators/engulfers; rare clingers; KY, NC, VA, WV ....................................................... Diploperla kanawholensis

PERLODIDAE, Helopicus

(After Stark & Ray, 1983. Suitable characters are available for both regional species.) 131(45) 131’

Anterior margin of dark pigment band on frons almost straight (Fig. 3.263); occiput predominantly brown; predators/engulfers; clingers; AL, CT, FL, KY, ME, NC, ON, PA, QC, SC, TN, VA, WV . .............................................................. Helopicus subvarians Dark pigment band on frons with a pair of small projecting lobes along anterior margin (Fig. 3.264); occiput predominantly pale constrasting sharply with dark frons; predators/engulfers; clingers in Coastal Plains streams; AL, FL, GA, LA, MS, NC, SC ............................................................................................................ Helopicus bogaloosa

PERLODIDAE, Hydroperla

(After Nelson, 1996. Suitable characters are available for all regional species.) 132(45’) Dark pigment band near anterior margin of frons complete (Fig. 3.265); abdominal terga 1 to 9 each with uniform pale pigmentation; predators/ engulfers; clingers; TN ................................................................................................................ Hydroperla rickeri 132’ Dark pigment on anterior frons consisting of 2 to 4 diffuse spots sometimes connected by a narrow band (Fig. 3.266); abdominal terga 1 to 9 each with transverse row of small dark pigment spots (Fig. 3.267) ...................................................................................... 133 133(132’) Dark pigment of anterior frons consisting of two diffuse spots (Fig. 3.268); anterior margin of labrum not expanded into a large mesal lobe; predators/ engulfers; clingers in larger streams, uncommon in region; AR, IA, IL, IN, KS, MO, MS, TN, TX .......................... ............................................................................................................. Hydroperla fugitans 133’ Dark pigment of anterior frons usually consisting of four diffuse pigment patches connected by a narrow band (Fig. 3.266); anterior margin of labrum expanded by a large mesal lobe; predators/engulfers; clingers in Coastal Plain streams; FL, NC, SC ..................... .......................................................................................................... Hydroperla phormidia

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Fig. 3.259

Fig. 3.261

Fig. 3.260

Fig. 3.262

Fig. 3.264

Fig. 3.263

Fig. 3.265

Fig. 3.267

Fig. 3.266

Fig. 3.268

Figures 3.259–3.268, Perlodidae genn. spp. 3.259, Diploperla robusta, left hind femur, anterodorsal. 3.260, D. morgani, left hind femur, anterodorsal. 3.261, D. morgani, head, dorsal. 3.262, D. kanawholensis, head, drosal. 3.263, Helopicus subvarians, head, dorsal. 3.264, H. bogaloosa, head, dorsal. 3.265, Hydroperla rickeri, head, dorsal. 3.266, H. phormidia, head, dorsal. 3.267, H. phormidia, abdominal terga 8–10, dorsal. 3.268, H. fugitans, head, dorsal. Figure 3.265 redrawn from Nelson (1996); all other figures original.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

PERLODIDAE, Isogenoides

(After Sandberg & Stewart, 2005. Suitable characters are available for both regional species.) 134(43) 134’

Head pattern with well-defined pale M-line completely enclosed with dark pigment (Fig. 3.269); abdominal terga 1 to 9 with dark anterior and posterior bands, but bands sometimes thin (Fig. 3.270); predators/engulfers; clingers; CT, MA, MD, ME, NB, NC, NS, NY, PA, QC, VA, WV .......................................................... Isogenoides hansoni Head pattern with diffuse M-line not completely enclosed with dark pigment (Fig. 3.271); abdominal terga 1 to 9 with only a narrow, diffuse, anterior pigment band (Fig. 3.272); predators/engulfers; some populations occur in hyporhea in Coastal Plain streams; IA, IL, IN, KS, MI, MN, MS, NC, SC, TN, VA . ..................................... Isogenoides varians

PERLODIDAE, Isoperla

(Characters are extracted from Frison, 1935, 1937, 1942; Hilsenhoff & Billmyer, 1973; Hitchcock, 1974; Nelson & Kondratieff, 1983; Poulton & Stewart, 1991; Stark et al., 1998; Stewart & Stark, 2002). Currently 41 regional Isoperla species are recognized (Table 3.1). The eastern Nearctic species recently have been revised by Szczytko & Kondratieff (2015), who advise some species formerly listed for the southeastern region do not occur here. In addition, many of those that do occur in the region are described for the first time in the Szczytko & Kondratieff monograph and a few others are now considered synonyms. Larvae for 16 of the 41 species currently reported for the region are included in the key. The following species, whose larvae are known, previously were reported from the region, but are not included in the key for the reasons indicated: Isoperla coushatta (synonym of I. davisi), I. cotta (regional record considered invalid), I. namata (regional record considered invalid), and I. richardsoni (regional record considered invalid.). Colleagues in North Carolina currently are working to associate and describe Isoperla larvae in that state. 135(38,47) Lacinia with a single, sometimes short tooth (Fig. 3.273, 3.275) ........................................ 136 135’ Lacinia with two teeth (Fig. 3.274) . ..................................................................................... 137 136(135) Lacinial tooth projecting clearly beyond primary setae (Fig. 3.273); predators/engulfers; clingers in small to medium-sized streams; IL, IN, KY, MI, NY, OH, ON, PA, QC, WI ........................................................................................................................ Isoperla nana 136’ Lacinial tooth short, sometimes absent or scarcely projecting beyond primary setae (Fig. 3.275); predators/engulfers; clingers in small streams; AL, IL, IN, KY, MI, MO, OH, ON, TN .............................................................................................. Isoperla decepta 137(135’) Body of lacinia quadrate with thick brush of primary setae along apical margin (Fig. 3.276); predators/engulfers; clingers in creeks and small rivers; ME, MI, MN, NB, NC, NL, NS, NY, ON, QC, TN, WI, WV . ..................................................................... Isoperla lata 137’ Body of lacinia more triangular, without thick anterior setal brush (Fig. 3.274, 3.277) ...... 138 138(137’) Body of lacinia subequal in length to longest tooth (Fig. 3.274) . ........................................ 139 138’ Body of lacinia distinctly longer than longest tooth (Fig. 3.277) ......................................... 140

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Fig. 3.269

Fig. 3.270

Fig. 3.271

Fig. 3.272 Fig. 3.273

Fig. 3.275

Fig. 3.276

Fig. 3.274

Fig. 3.277

Figures 3.269–3.277, Isogenoides spp. (Perlodidae). 3.269, Isogenoides hansoni, head, dorsal. 3.270, I. hansoni, abdominal terga 8–10, dorsal. 3.271, I. varians, head, dorsal. 3.272, I. varians, abdominal terga 8–10, dorsal. 3.273, Isoperla nana, right maxilla, ventral. 3.274, I. burksi, right maxilla, ventral. 3.275, I. decepta, right maxilla, ventral. 3.276, I. lata, right maxilla, ventral. 3.277, I. major, right lacinia, ventral. Figures 3.273, 3.275 redrawn from Frison (1935); 3.275, 3.276 redrawn from Frison (1942); 3.277 redrawn from Nelson & Kondratieff (1983); all other figures original.

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139(138) Abdominal terga with transverse pigment bands (Fig. 3.278); predators/ engulfers; clingers in small to medium-sized streams; AL, AR, IL, IN, KY, MD, MO, NC, OH, OK, SC, VA, WV . ...................................................................................................... Isoperla burksi 139’ Abdominal terga with longitudinal pigment stripes (Fig. 3.279); predators/ engulfers; clingers in medium-sized creeks to small rivers; NC, PA, TN, VA . ........... Isoperla orata 140(138’) Head with narrow, pale M-line completely enclosed in dark pigment (Fig. 3.280) ............. 141 140’ Head pattern variable but without narrow M-line enclosed in dark pigment ....................... 142 141(140) Abdominal terga brown, usually with pale spots but without dark stripes (Fig. 3.281); mature larva at least 18 mm long; predators/engulfers; clingers in headwater spring habitat, emerging in August and September; VA ........................................ Isoperla major 141’ Abdominal terga mostly brown but bearing slightly darker dorsolateral longitudinal pigment stripes (Fig. 3.282); mature larvae no more than 12 mm long; predators/ engulfers; clingers in headwater streams and rheocrenes, emerging April to June; CT, MD, ME, NH, NJ, NY, PA, VA . ................................................................. Isoperla similis 142(140’) Abdominal terga without distinct longitudinal dark stripes (Fig. 3.281, 3.283) . ................. 143 142’ Abdominal terga with distinct longitudinal dark stripes (Fig. 3.284) . ................................. 145 143(142) Abdominal terga dark, often patterned with pale spots (Fig. 3.283) but transverse banding, if present, consisting of two dark bands ......................................................................... 144 143’ Abdominal terga each with a single dark transverse band along posterior margin (Fig. 3.285); lacinia with approximately 6 major setae set near base of second tooth (Fig. 3.286); abdominal tergum 10 predominantly pale; predators/ engulfers; clingers in small creeks to larger rivers; AR, CT, IA, MB, ME, MI, MN, MO, NB, NS, NY, OH, OK, ON, PA, VA, WI, WV ....................................................................... Isoperla signata

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Fig. 3.278

Fig. 3.279

Fig. 3.280

Fig. 3.281

Fig. 3.282

Fig. 3.283

Fig. 3.284

Fig. 3.285

Fig. 3.286

Figures 3.278–3.286, Perlodidae, Isoperla spp. 3.278, Isoperla burksi, abdominal terga 8–10, dorsal. 3.279, I. orata, abdominal terga 8–10, dorsal. 3.280, I. major, head, dorsal. 3.281, I. major, abdominal terga 8–10, dorsal. 3.282, I. similis, abdominal terga 8–10, dorsal. 3.283, I. slossonae, abdominal terga 8–10, dorsal. 3.284, Isoperla sp., abdominal terga 7–10, dorsal. 3.285, I. signata, abdominal terga 8–10, dorsal. 3.286, I. signata, right maxilla. ventral. Figures 3.278, 3.279, 3.282, 3.283 redrawn from Frison (1942); 3.280, 3.281 redrawn from Nelson & Kondratieff (1983); 3.285, 3.286 redrawn from Poulton & Stewart, 1991; 3.284 original.

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144(143) Tergum 10 with dark pigment, other terga with dark anterior and posterior margins, or completely dark; lacinia with numerous major setae extending along inner margin to near midlength (Fig. 3.287); predators/engulfers; clingers in larger creeks and rivers; CO, IA, IL, IN, KY, MB, ME, MI, MN, NB, NE, NJ, NS, NY, ON, PA, QC, VA, WI, WV ................................................................................................................. Isoperla marlynia 144’ Apical half of tergum 10 pale, other terga dark, often with pale spots but without dark transverse anterior and posterior bands (Fig. 3.288); lacinia with major setae extending nearly to base; predators/engulfers; clingers in small to medium-sized streams; IA, ME, MI, MN, NB, NC, NH, NS, NY, PA, PE, QC, VA, WI, WV ................. Isoperla slossonae 145(142’) Median and lateral dark abdominal stripes similar in width and pigment intensity on terga 7 to 8 (Fig. 3.290) ........................................................................................................... 146 145’ Dark median stripe absent, or either wider, narrower, or more poorly defined than lateral stripes . ............................................................................................................... 148 146(145) Dark pronotal pigment forming four irregularly shaped sublateral spots, anterior projections of dark, transverse pigment band on frons not approaching anterior margin of frons and not connected anterior to median ocellus (Fig. 3.289); predators/engulfers; clingers in small to medium-sized rivers; CT, DE, GA, IN, ME, MI, MN, NB, NC, NS, NY, ON, PA, PE, QC, SC, TN, VA, WI . ................................................................... Isoperla frisoni 146’ Pronotum with extensive dark pigment adjacent to pale marginal flange (Fig. 3.292); anterior projections of dark transverse pigment bands reaching frontoclypeal area and often connecting, or nearly so, anterior to median ocellus (Fig.3.291) .......................... 147 147(146’) Darkest pigment on head usually connected anterior to median ocellus and behind lateral ocelli (Fig. 3.291); abdominal terga pale brown adjacent to dark stripes; lacinial setal row extending along most of inner margin; predators/ engulfers; clingers in small Coastal Plain creeks and rivers; AL, AR, DE, FL, LA, MS, NC, OK, SC, TX, VA ...................................................................................................................... Isoperla davisi 147’ Darkest pigment on head not usually connected anterior to median ocellus, or behind lateral ocelli (Fig. 3.292); abdominal terga white adjacent to dark stripes; lacinial setal row extending to midlength of inner margin; predators/ engulfers; clingers in small to medium-sized streams; BC, CT, DE, IA, KY, MB, ME, MI, MN, NB, NC, NH, NL, NS, NY, OH, ON, PA, PE, QC, SD, SK, WI, WV . ........................... Isoperla transmarina 148(145’) Dark median abdominal stripe wider and more prominent than lateral stripes (Fig. 3.293); abdominal terga dark on either side of median stripe and bearing small freckle-like spots; predators/engulfers; clingers in medium-sized to large creeks; AL, CT, FL, GA, IA, IN, KY, MA, MB, ME, MI, MN, MO, MS, NB, NC, NY, OH, ON, PA, QC, SC, TN, VA, WI, WV ......................................................................................... Isoperla dicala 148’ Dark median abdominal stripe narrower and often less prominent than lateral stripes (Fig. 3.294); abdominal terga pale on either side of median stripe ................................ 149

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Fig. 3.288 Fig. 3.287

Fig. 3.290

Fig. 3.289

Fig. 3.291

Fig. 3.293

Fig. 3.292

Fig. 3.294

Figures 3.287–3.294, Isoperla spp. (Perlodidae). 3.287, I. marlynia, right lacinia, ventral. 3.288, I. slossonae, abdominal terga 8–10, dorsal. 3.289, I. frisoni, head and pronotum, dorsal. 3.290, Isoperla sp., abdominal terga 8–10, dorsal. 3.291–3.292, heads, dorsal: 3.291, I. davisi; 3.292, I. transmarina. 3.293–3.294, abdominal terga 8–10, dorsal: 3.293, I. dicala. 3.294, I. holochlora. Figures 3.287–3.288, 3.292–3.294 redrawn from Frison, 1942; 3.289 redrawn from Frison, 1937, 3.290–291 original.

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149(148’) Ocellar triangle open across posterior margin (Fig. 3.295); pale areas on posterior abdominal terga small, generally smaller than dark lateral pigment; lacinial setal row extends along much of inner margin (Fig. 3.296); predators/engulfers; clingers in larger streams; IA, IL, IN, KS, MB, MI, MN, MO, MS, ND, NE, OH, SK, WI ................... Isoperla bilineata 149’ Ocellar triangle usually closed on posterior margin (Fig. 3.298); dark lateral abdominal stripes prominent but not as wide as pale areas on terga (Fig. 3.297); lacinial setal row extending along less than half the inner lacinial margin; predators/engulfers; clingers in small to medium-sized creeks and small rivers; AL, CT, DE, KY, MD, ME, NC, NS, NY, OH, PA, QC, SC, TN, VA, WV . .................................................. Isoperla holochlora PERLODIDAE, Remenus (Characters are extracted from Stewart & Stark, 2002. Characters are available for two of the three regional species.) 150(39) Basal and mid-cercal segments each with well-developed whorl of mixed long and short setae (Fig. 3.299); predators/engulfers; clingers; AL, CT, DE, KY, MD, NC, NY, PA, SC, TN, VA, WV................................................................................... Remenus bilobatus 150’ Basal and mid-cercal segments with poorly developed setal whorls (Fig. 3.300); predators/ engulfers; clingers; GA . ......................................................................... Remenus duffieldi PERLODIDAE, Yugus (Characters are extracted from Frison, 1942; Nelson, 2001; Stewart & Stark, 2002. Characters are available for all regional species.) 151(48’) Pale longitudinal band extending from median ocellus to anterior margin of frontoclypeus (Fig. 3.301); pale M-line not completely outlined by dark pigment on anterior margin; predators/engulfers; clingers; GA, NC, SC, TN, VA ..................................... Yugus arinus 151’ Pale longitudinal band absent between median ocellus and anterior margin of frontoclypeus (Fig. 3.302); pale M-line completely outlined by diffuse to dark pigment along anterior margin . .............................................................................................................. 152 152(151’) Marginal pigment outlining pale area in ocellar triangle not darker than surrounding band on head (Fig. 3.302); predators/engulfers; clingers; NC, VA ................ Yugus kondratieffi 152’ Marginal pigment outlining pale area in ocellar triangle darker than surrounding band on head (Fig. 3.303) ............................................................................................................. 153 153(152’) Entire anterior frontoclypeal margin dark (Fig. 3.304); predators/engulfers; clingers; KY, PA, VA, WV................................................................................................ Yugus kirchneri 153’ Anterior frontoclypeal margin with median pale spot (Fig. 3.303); predators/ engulfers; clingers; GA, NC, TN ................................................................................ Yugus bulbosus

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Fig. 3.295

Fig. 3.296

Fig. 3.297

Fig. 3.299

Fig. 3.298

Fig. 3.300

Fig. 3.301

Fig. 3.302

Fig. 3.303

Fig. 3.304

Figures 3.295–3.304, Perlodidae genn. spp. 3.295–3.296, Isoperla bilineata: 3.295, head, dorsal; 3.296, right lacinia, ventral. 3.297–3.298, I. holochlora: 3.297, abdominal terga 8–10, dorsal. 3.298, head and pronotum, dorsal. 3.299–3.300, basal and mid segments of left cerci, dorsal: 3.299, Remenus bilobatus. 3.300, R. duffieldi. 3.301–3.304, heads, dorsal: 3.301, Yugus arinus. 3.302, Y. kondratieffi. 3.303, Y. bulbosus. 3.304, Y. kirchneri. Figures 3.295–3.296 redrawn from Poulton & Stewart, 1991; 3.297–3.298 redrawn from Frison, 1942; all other figures original.

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PTERONARCYIDAE, Pteronarcys

(After Ricker, 1952. Characters are available for the six known regional species.) 154(2) 154’

Abdomen without lateral knobs on any segment (Fig. 3.305) . ............................................ 155 Abdomen with lateral knobs on several segments (Fig. 3.306) . .......................................... 156

155(154) Anterior projections on pronotum directed posterolaterally at tip; uncommon in southeastern region, shredders/detritivores; clingers; AR, CT, IL, IN, KS, KY, MB, MI, MN, MO, ND, NE, OH, PA, TN, WI .................................................................... Pteronarcys pictetii 155’ Anterior projections on pronotum directed anterolaterally; common throughout southeastern region; shredders/detritivores; clingers; AB, AK, AL, BC, CO, FL, GA, IL, IN, KS, KY, LA, MB, ME, MI, MN, MS, MT, NB, NC, NL, NT, NY, OH, ON, PA, QC, SC, SK, TN, VA, WI, WV, WY .................................................................. Pteronarcys dorsata 156(154’) Knobs on abdominal segments 7 and 8 reduced to slightly thickened areas, scarcely projecting beyond level of tergum (Fig. 3.307) .............................................................. 157 156’ Knobs on abdominal segments 7 and 8 more spine-like, projecting distinctly beyond level of tergum (Fig. 3.308) ..................................................................................................... 158 157(156) Knobs on abdominal segment 7 slightly smaller than, but similar to, those on segment 6 (Fig. 3.310); shredders/detritivores; clingers; KY, MB, MD, ME, NC, NH, NJ, NY, ON, PA, QC, VA, VT, WV.................................................................... Pteronarcys proteus 157’ Knobs on abdominal segment 7 reduced to thickened areas, slightly larger than, but similar to, those on segment 8 (Fig. 3.307); shredders/detritivores; clingers; GA, NC, PA, SC, TN, VA .................................................................................................... Pteronarcys scotti 158(156’) Mesonotum with small basolateral spine (Fig. 3.309); shredders/ detritivores; clingers; KY, MA, ME, NB, NC, NH, NS, NY, PA, VA, WV ...................... Pteronarcys comstocki 158’ Mesonotum without basolateral spine (Fig. 3.311); shredders/detritivores; clingers; AL, CT, GA, MA, MD, ME, NB, NC, NH, NS, NY, PA, PE, QC, SC, VA, WV ................................................................................................................ Pteronarcys biloba

TAENIOPTERYGIDAE, Strophopteryx

(After Earle & Stewart, 2008; Harper & Hynes, 1971c; Poulton & Stewart, 1991. The three species known from the region are included in the following key.) 159(51) 159’

Abdominal terga dark on anterior half, pale yellow brown on posterior half, and often bearing a median, transverse row of dark spots (Fig. 3.312); likely shredders/scrapers/ detritivores; clingers; AL, AR, CT, DE, IA, IL, IN, KS, KY, MB, ME, MI, MN, MO, MS, NC, ND, OH, OK, ON, PA, QC, SC, VA, WI, WV ............... Strophopteryx fasciata Abdominal tergal pattern without distinct dark transverse bands, dark pigment spots present or absent (Fig. 3.313) ...................................................................................................... 160

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Fig. 3.305

Fig. 3.306

Fig. 3.307

Fig. 3.308

Fig. 3.309

Fig. 3.310

Fig. 3.311

Fig. 3.312

Fig. 3.313

Figures 3.305–3.313, Pteronarcys spp. (Pteronarcyidae) and Strophopteryx spp. (Taeniopterygidae). 3.305–3.308, abdominal terga 7–10, dorsal: 3.305, Pteronarcys dorsata. 3.306, P. biloba. 3.307, P. scotti. 3.308, P. biloba. 3.309, P. comstocki, right halves of pro- and mesonota, dorsal. 3.310, P. proteus, abdominal terga 6–10, dorsal. 3.311, P. biloba, right halves of pro- and mesonota, dorsal. 3.312–3.313, abdominal terga 8–10, dorsal: 3.312, Strophopteryx fasciata; 3.313, S. appalachia. All figures original.

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160(159’) Fore and mid-tibiae bearing sparse fringe of fine, long setae along inner margins (Fig. 3.316); dark pigment spots on abdominal terga obscure or absent (Fig. 3.313); likely shredders/scrapers/detritivores; clingers; KY, NC, NY, PA, SC, TN, VA, WV .................................................................................................... Strophopteryx appalachia 160’ Fore and mid-tibiae without inner fringe of fine, long setae (Fig. 3.315); abdominal terga with row of small dark spots on pale brown background (Fig. 3.314); likely shredders/ scrapers/detritivores; clingers; NC, TN, VA . .................................... Strophopteryx limata

TAENIOPTERYGIDAE, Taeniopteryx

(After Fullington & Stewart, 1980; Harper & Hynes, 1971c; Kondratieff & Baumann, 1988; Poulton & Stewart, 1991. Characters are extracted from Kondratieff & Kirchner 1982, 1984. All species known from the region are included in the following key.) 161(49) 161’

Abdominal terga without pale, mid-dorsal stripe, if present, broken, incomplete, or obscure ......................................................................................................................................... 162 Abdominal terga with pale mid-dorsal stripe (Fig. 3.317) ................................................... 164

162(161) Cerci and basal third of antennae bearing segmental whorls of long setae (Fig. 3.318); body often covered with mixtures of detritus, sand, and periphyton; likely shredders/ detritivores; sprawlers/clingers; NC, VA ........................................... Taeniopteryx nelsoni 162’ Cercal and antennal segments lacking whorls of long setae; body not covered with detritus, sand, and periphyton ....................................................................................................... 163 163(162’) Head bearing a prominent broad pale chevron between eyes (Fig. 3.319); likely shredders/ detritivores; sprawlers/clingers; AL, GA, KY, PA, TN, VA ........................................... ............................................................................................... Taeniopteryx ugola (in part) 163’ Pale area between eyes narrow with expanded ends (Fig. 3.320); shredders/scrapers/ detritivores; sprawlers/clingers; AB, AR, CO, CT, GA, IL, IN, KY, MB, ME, MI, MN, MO, MS, NC, NM, NY, OH, ON, PA, QC, SC, TN, VA, WI, WV, WY ........................ ........................................................................................................... Taeniopteryx parvula 164(161’) Pale mid-dorsal abdominal stripe inconspicuous; pale area between eyes chevron shaped (Fig. 3.319); likely shredders/detritivores; sprawlesr/clingers; AL, GA, KY, PA, TN, VA ............................................................................................... Taeniopteryx ugola (in part) 164’ Pale mid-dorsal abdominal stripe conspicuous (Fig. 3.321); pale area between eyes if present, not chevron shaped............................................................................................. 165 165(164’) Posterior finge on abdominal terga consisting mostly of short peg setae (Fig. 3.322); pale mid-dorsal abdominal stripe often bordered by dark pigment (Fig. 3.321) . .................. 166 165’ Posterior fringe on abdominal terga consisting mostly of slender curved setae (Fig. 3.323); pale mid-dorsal abdominal stripe without dark margins .................................................167

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Fig. 3.316

Fig. 3.314 Fig. 3.315

Fig. 3.318

Fig. 3.317

Fig. 3.319

Fig. 3.320

Fig. 3.322

Fig. 3.321

Fig. 3.323

Figures 3.314–3.323, Strophopteryx spp. and Taeniopteryx spp. (Taeniopterygidae. 3.314, Strophopteryx limata, abdominal terga 8–10, dorsal. 3.315–3.316, left forelegs, anterodorsal: 3.315, S. limata. 3.316, S. appalachia. 3.317, Taeniopteryx maura, abdominal terga 8–10. 3.318, Taeniopteryx nelsoni, base of right antenna, dorsal. 3.319–3.320, heads, dorsal: 3.319, T. ugola. 3.320, T. parvula. 3.321, T. maura, abdominal terga 8–10, dorsal. 3.322–3.323, posterior fringes of abdominal setae: 3.322, T. maura. 3.323, T. lonicera. All figures original.

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166(165) Pre-emergent male larvae with developing spur visible through exoskeleton of inner margin of hind femora (Fig. 3.324); adult males may be needed to confirm identity; shredders/ scrapers/detritivores; sprawlers/ clingers; AL, AR, CT, DE, GA, IN, KY, MA, MD, ME, MI, MN, MS, NC, NY, OH, PA, SC, TN, TX, VA, WV ............ Taeniopteryx maura 166’ Pre-emergent male larvae without developing spur under exoskeleton on inner margin of hind femora; adult males may be needed to confirm identity; likely shreddersdetritivores; sprawlers/clingers; AL, AR, CO, CT, DE, FL, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, MS, NC, NE, NY, OH, OK, ON, PA, QC, SD, TN, TX, VA, WI, WV .............................................................................................................. Taeniopteryx burksi 167(165’) Frons with large pale spot over ocellar triangle (Fig. 3.325); pre-emergent male larvae with developing epiproct broad at apex .................................................................................. 168 167’ Frons without pale spot over ocellar triangle (Fig. 3.326); pre-emergent male larvae with developing epiproct narrowed apically . ......................................................................... 169 168(167) 168’

Frontoclypeal region usually occupied by dark U-shaped area (Fig. 3.325); pre-emergent male larvae at least 8.0 mm in length; likely shredders/detritivores; sprawlers/clingers; AL, AR, FL, IL, IN, KY, LA, MS, NC, NJ, OK, SC, TX, VA, WV .................... Taeniopteryx lita Frontoclypeal region usually occupied by pair of dark blotches (Fig. 3.327); pre-emergent male larvae less than 7.6 mm in length; likely shredders/ detritivores; sprawlers/clingers; AL, AR, FL, GA, LA, MD, MS, NC, SC, TN, TX, VA .................. Taeniopteryx lonicera

169(167’) Central frons pale brown with scattered darker areas; femora and tibiae pale; likely shredders/detritivores; sprawlers/clingers; SC ................................. Taeniopteryx robinae 169’ Central frons adjacent to anterior bases of antennae covered with dark brown pigment (Fig. 3.326); femora dark, tibiae and basal two tarsal segments pale; likely shredders/ detritivores; sprawlers/clingers; AL, AR, IL, IN, KS, KY, MO, NC, NY, OH, OK, ON, PA, VA, WV ...................................................................................... Taeniopteryx metequi

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Fig. 3.324 Fig. 3.325

Fig. 3.326

Fig. 3.327

Figures 3.324–3.327, Taeniopteryx spp. (Taeniopterygidae. 3.324, Taeniopteryx maura, left hind femur, anterodorsal. 3.325–3.327, heads, dorsal: 3.325, T. lita. 3.326, T. metequi. 3.327, T. lonicera. All figures original.

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References Cited Baumann, R.W. 1975. Revision of the stonefly family Nemouridae (Plecoptera): A study of the world fauna at the generic level. Smithsonian Contributions to Zoology 211: 1–74. Brown, LD., & B.P. Stark. 1995. Nymphs and eggs of Alloperla natchez and Haploperla chukcho (Plecoptera: Chloroperlidae). Journal of the Kansas Entomological Society 68: 120–125. DeWalt, R.E., M.D. Maehr, U. Neu-Becker, & G. Steuber. 2015. Plecoptera species file online.Version 5.0/5.0. Available from http://Plecoptera.SpeciesFile.org/HomePage.aspx (accessed 5 March 2015). Earle, J.I., & K.W. Stewart. 2008. Description of the nymph of Strophopteryx appalachia Ross and Ricker (Plecoptera: Taeniopterygidae), and key to Strophopteryx nymphs. Proceedings of the Entomological Society of Washington 110: 551–555. Fiance, S.B. 1977. The genera of eastern North American Chloroperlidae (Plecoptera): Key to larval stages. Psyche 84: 308–316. Fochetti, R., & J.M. Tierno de Figueroa. 2008. Global diversity of stoneflies (Plecoptera; Insecta) in freshwater. Hydrobiologia 595: 365–377. Frison, T.H. 1935. The stoneflies, or Plecoptera, of Illinois. Bulletin of the Illinois Natural History Survey 20: 281–471. Frison, T.H. 1937. Studies of Nearctic aquatic insects. II. Descriptions of Plecoptera with special reference to the Illinois species. Bulletin of the Illinois Natural History Survey 21: 78–99. Frison, T.H. 1942. Studies of North American Plecoptera with special reference to the fauna of Illinois. Bulletin of the Illinois Natural History Survey 22: 235–355. Fullington, K.E., & K.W. Stewart. 1980. Nymphs of the stonefly genus Taeniopteryx (Plecoptera: Taeniopterygidae) of North America. Journal of the Kansas Entomological Society 53: 237–259. Grubbs, S., B.C. Kondratieff, B.P. Stark, & R.E. DeWalt. 2013. A review of the Nearctic genus Zealeuctra Ricker (Plecoptera, Leuctridae), with the description of a new species from the Cumberland Plateau region of eastern North America. ZooKeys 344: 17–47. Grubbs, S., R. Baumann, & A. Sheldon. 2015. A review of eastern Nearctic Zapada (Plecoptera, Nemouridae) with a new species from the Great Smoky Mountains. Freshwater Science, 34:1312-1323. Harper, P.P., & H.B.N. Hynes. 1971a. The Leuctridae of eastern Canada (Insecta; Plecoptera). Canadian Journal of Zoology 49: 915–920. Harper, P.P., & H.B.N. Hynes. 1971b. The Capniidae of eastern Canada (Insecta; Plecoptera). Canadian Journal of Zoology 49: 921–940. Harper, P.P., & H.B.N. Hynes. 1971c. The nymphs of the Taeniopterygidae of eastern Canada (Insecta; Plecoptera). Canadian Journal of Zoology 49: 941–947. Harper, P.P., & H.B.N. Hynes. 1971d. The nymphs of Nemouridae of eastern Canada (Insecta: Plecoptera). Canadian Journal of Zoology 49: 1129–1142. Harrison, A.B., & B.P. Stark. 2010. Two new species of stoneflies in the Leuctra ferruginea group (Plecoptera: Leuctridae), with notes on the Leuctra species known for Mississippi and Alabama, U.S.A. Illiesia 6: 16–33. Hilsenhoff, W.L., & S.J. Billmyer. 1973. Perlodidae (Plecoptera) of Wisconsin. The Great Lakes Entomologist 6: 1–14. Hitchcock, S.W. 1974. Guide to the Insects of Connecticut. Part VII. The Plecoptera or Stoneflies of Connecticut. State Geological and Natural History Survey of Connecticut, Bulletin Number 107, Hartford, Connecticut. Kirchner, R.F., & B.C. Kondratieff. 1984. A new Diploperla from West Virginia (Plecoptera: Perlodidae). Proceedings of the Entomological Society of Washington 86: 648–652.

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Kirchner, R.F., & B.C. Kondratieff. 1985. The nymph of Hansonoperla appalachia Nelson (Plecoptera: Perlidae). Proceedings of the Entomological Society of Washington 87: 593–596. Kirchner, R.F., & B.C. Kondratieff. 1997. A new species of Nearctic Perlesta (Plecoptera: Perlidae) from Virginia. Proceedings of the Entomological Society of Washington 99: 290–293. Kondratieff, B.C. 2004. Perlodidae-Perlodinae (The Springflies). In: Stark, B.P., & B.J. Armitage (Eds.). Stoneflies (Plecoptera) of Eastern North America. Volume 2. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp.149–180. Kondratieff, B.C., & R.W. Baumann. 1988. Taeniopteryx of western North America (Plecoptera: Taeniopterygidae). Pan-Pacific Entomologist 64: 381–390. Kondratieff, B.C., & R.F. Kirchner. 1982. Taeniopteryx nelsoni, a new species of winter stonefly from Virginia (Plecoptera: Taeniopterygidae). Journal of the Kansas Entomological Society 55: 1–7. Kondratieff, B.C., & R.F. Kirchner. 1988. A new species of Acroneuria from Kentucky (Plecoptera: Perlidae) and new records of stoneflies from eastern North America. Journal of the Kansas Entomological Society 61: 201–207. Kondratieff, B.C., R.F. Kirchner, & K.W. Stewart. 1988. A review of Perlinella Banks (Plecoptera: Perlidae). Annals of the Entomological Society of America 81: 19–27. Kondratieff, B.C., R.F. Kirchner, & J.R. Voshell. 1981. Nymphs of Diploperla. Annals of the Entomological Society of America 74: 428–430. Myers, L.W., & B.C. Kondratieff. 2009. Descriptions of the nymphs of eastern North American species of Cultus (Plecoptera: Perlodidae). Entomologica Americana 115: 109–114. Needham, J.G., & P.W. Claassen. 1925. A Monograph of the Plecoptera or Stoneflies of America North of Mexico. Volume 2. Thomas Say Foundation, Lafayette, Indiana. Nelson, C.H. 1996. Placement of Helopicus rickeri Stark in Hydroperla Frison (Plecoptera: Perlodidae) with the description of the adult female, nymph and egg and a cladistic analysis of Hydroperla. Proceedings of the Entomological Society of Washington 98: 237–244. Nelson, C.H. 1997. Descriptions of the female, nymph, egg and redescription of the male of Amphinemura mockfordi (Plecoptera: Nemouridae). Entomological News 108: 107–112. Nelson, C.H. 2000. Pteronarcyidae (The Salmonflies). In Stark, B.P., & B.J. Armitage (Eds).. Stoneflies (Plecoptera) of Eastern North America. Volume 1. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp.29–39. Nelson, C.H. 2001. The Yugus bulbosus complex, with a comment on the phylogenetic position of Yugus within the eastern Perlodini (Plecoptera: Perlodidae: Perlodinae). Proceedings of the Entomological Society of Washington 103: 601–619. Nelson, C.H., & B.C. Kondratieff. 1983. Isoperla major, a new species of eastern Nearctic Isoperlinae (Plecoptera: Perlodidae). Annals of the Entomological Society of America 76: 270–273. Poulton, B.C., & K.W. Stewart. 1991. The Stoneflies of the Ozark and Ouachita Mountains (Plecoptera). Memoirs of the American Entomological Society, Number 38. Philadelphia, Pennsylvania. Ray, D. H., A.K. Rasmussen, J.G. Peters, &, B.P. Stark. 2010. The larva and egg of Alloperla prognoides (Plecoptera: Chloroperlidae), with ecological notes and new state records from Florida, U.S.A. Illiesia 6: 256–266. Ricker, W.E. 1952. Systematic Studies in Plecoptera. Indiana University Publications, Science Series No. 18. Indiana University Press, Bloomington, Indiana. Ross, H. H., & W.E. Ricker. 1971. The Classification, Evolution and Dispersal of the Winter Stonefly Genus Allocapnia. Illinois Biological Monographs, 45. University of Illinois Press, Urbana and Chicago, Illinois.

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Sandberg, J.B., & K.W. Stewart. 2005. Holomorphology and systematics of the stonefly genus Isogenoides (Plecoptera: Perlodidae). Transactions of the American Entomological Society 131: 269–345. Shepard, W.D., & K.W. Stewart. 1983. Comparative study of nymphal gills in North American stonefly (Plecoptera) genera and a new, proposed paradigm of Plecoptera gill evolution. Miscellaneous Publications of the Entomological Society of America 13: 1–57. Stark, B.P. 1986. The Nearctic species of Agnetina (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 59: 437–445. Stark, B.P. 1989. Perlesta placida (Hagen), an eastern Nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica 20: 263–286. Stark, B.P. 1995. A new species of Neoperla (Insecta: Plecoptera: Perlidae) from Mississippi. Proceedings of the Biological Society of Washington 108: 45–49. Stark, B.P. 2000. Peltoperlidae (The Roachflies). In Stark, B.P., & B.J. Armitage (Eds.). Stoneflies (Plecoptera) of Eastern North America. Volume 1. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp.41-54. Stark, B.P. 2004. Perlidae (The Stones). In Stark, B.P., & B.J. Armitage (Eds.). Stoneflies (Plecoptera) of Eastern North America. Volume 2. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp. 61-148. Stark, B.P., & B.J. Armitage (Eds.). 2000. Stoneflies (Plecoptera) of Eastern North America. Volume l. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Bulletin of the Ohio Biological Survey, New Series, Volume 14. Ohio Biological Survey, Columbus, Ohio. Stark, B.P., & B.J. Armitage (Eds.). 2004. Stoneflies (Plecoptera) of Eastern North America. Volume II. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Bulletin of the Ohio Biological Survey, New Series, Volume 14. Ohio Biological Survey, Columbus, Ohio. Stark, B.P., & A.R. Gaufin. 1976. The Nearctic genera of Perlidae (Plecoptera). Miscellaneous Publications of the Entomological Society of America 10: 1–77. Stark, B P., & Harrison, A.B. 2010. The larva of Amphinemura alabama Baumann and new records of Nemouridae (Plecoptera) from Mississippi, U.S.A. Illiesia 6: 234–240. Stark, B.P., & B.C. Kondratieff. 2004. Acroneuria kirchneri (Plecoptera: Perlidae), a new species from eastern North America. Annals of the Entomological Society of America 97: 393–396. Stark, B.P., & B.C. Kondratieff. 2010. Larvae of eight eastern Nearctic Alloperla species (Plecoptera: Chloroperlidae). Illiesia 6: 267–276. Stark, B.P., & J.W. Lacey. 2005. Larvae of the winter stonefly genus Allocapnia (Plecoptera: Capniidae) in Mississippi, U.S.A. Illiesia 1: 10–20. Stark, B.P., & D.H. Ray. 1983. A revision of the genus Helopicus (Plecoptera: Perlodidae). Freshwater Invertebrate Biology 2: 16–27. Stark, B.P., & K.W. Stewart. 1982. The nymph of Viehoperla ada (Plecoptera: Peltoperlidae). Journal of the Kansas Entomological Society 55: 494–498. Stark, B.P., & S.W. Szczytko. 1981. Contributions to the systematics of Paragnetina (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 54: 625–648. Stark, B.P., C. Froehlich, & M.C. Zúñiga. 2009. South American Stoneflies (Plecoptera). Volume 5. Aquatic Biodiversity in Latin America. Pensoft, Sofia-Moscow. Stark, B.P., S.W. Szczytko,, & C.R. Nelson. 1998. American Stoneflies: A Photographic Guide to the Plecoptera. The Caddis Press, Columbus, Ohio. Stark, B.P., B.C. Kondratieff, R.F. Kirchner, & K.W. Stewart. 2011. Larvae of eight eastern North American Sweltsa (Plecoptera: Chloroperlidae). Illiesia 7: 51–64.

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Stark, B.P., K.W. Stewart, S.W. Szczytko, R.W. Baumann, & B.C. Kondratieff. 2012. Scientific and common names of stoneflies (Plecoptera) from the United States and Canada, with corrections and additions to the list. Miscellaneous Contributions of the Caddis Press 1: 1–20. Stewart, K.W. 2000. Taeniopterygidae (The Willowflies). In Stark, B.P., & B.J. Armitage (Eds.). Stoneflies (Plecoptera) of Eastern North America. Volume 1. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp. 55–88. Stewart, K.W., & B.P. Stark. 2002. Nymphs of North American Stonefly Genera (Plecoptera). Second Edition. The Caddis Press, Columbus, Ohio. Stewart, K.W., & B.P. Stark. 2008. Plecoptera. In Merritt, R.W., K.W. Cummins, & M.B. Berg. (Eds.). An Introduction to the Aquatic Insects of North America. Fourth Edition. Kendall/Hunt Publishing Company, Dubuque, Iowa, pp. 311–384. Surdick, R.F. 2004. Chloroperlidae (The Sallflies). In Stark, B.P., & B.J. Armitage (Eds.). Stoneflies (Plecoptera) of Eastern North America. Volume 2. Chloroperlidae, Perlidae, and Perlodidae (Perlodinae). Bulletin of the Ohio Biological Survey, New Series. Volume 14. Ohio Biological Survey, Columbus, Ohio, pp. 1–60. Szczytko, S.W., & B.C. Kondratieff. 2015. A Review of the Eastern Nearctic Isoperlinae (Plecoptera: Perlodidae) with the Description of Twenty-two New Species. Monographs of Illiesia, Number 1. Ljubljana, Slovenia. Zwick, P. 1973. Insecta: Plecoptera. Phylogenetisches System und Katalog. Das Tierreich 94. Walter de Gruyter, Berlin. Zwick, P. 2004. Key to the west Palearctic genera of stoneflies (Plecoptera) in the larval stage. Limnologica 34: 315–348. Zwick, P. 2006. New family characters of larval Plecoptera, with an analysis of the Chloroperlidae: Paraperlinae. Aquatic Insects 28: 13–22.

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Chapter 4

TRICHOPTERA

John C. Morse , Ralph W. Holzenthal, Oyunchuluun Yadamsuren With over 15,000 known, extant species globally (Morse, 2015, and in preparation), the caddisflies, or Trichoptera, are the seventh largest order of insects, with more species than the combined number of species in the other primarily aquatic insect orders (Ephemeroptera, Odonata, Plecoptera, Megaloptera) (Morse, 2009). Among all insect orders, only the Diptera have more known freshwater species (Morse, 2009). The North American caddisfly fauna currently includes 1,887 species, or about 13% of the total world fauna (Rasmussen & Morse, 2014). With 626 species known for the southeastern states (“*” in Table 4.1) and 37 others (“+” in Table 4.1) likely to be found here in coming years, one-third of these Nearctic species occur in the Southeast. Among the 663 caddisfly species known or likely to occur in the southeastern states, the larvae of only 309 (47%) have been described sufficiently to provide at least tentative diagnoses—our continuing inability to identify a majority of our species as larvae remains a significant impediment for use of these insects at the species level in ecological research or in biomonitoring programs. The symbol “#” in Table 4.1 indicates species for which the larva has been described sufficiently to be identified at least tentatively in the subsequent key; notes in the key and lack of this symbol in Table 4.1 consequently implicate those species for which a search for immature stages and for their differentiating characters still is needed. Readers of this synthesis are urged to rear suspected unknown larvae using methods recommended by Merritt et al. (2008a, pp. 30, 36, Table 3H) and to sequence and analyze DNA of identifiable adults and yet-unknown larvae (e.g., Zhou et al., 2007a, 2007b) in order to associate these life history forms, thereby identifying the larvae; you are then urged to explore these newly associated larvae for reliable diagnostic characters and to describe these differences in publicly accessible venues. Table 4.1 reflects the classification of Trichoptera families, genera, and species found in the Trichoptera World Checklist (Morse, 2015), with families arranged in alphabetical order, genera alphabetically within families, and species alphabetically within genera. Other, intermediate, taxonomic categories (e.g., subfamilies, subgenera) are provided in the key where this information seemed potentially useful. Abbreviated bibliographic references for original descriptions and for synonyms in Table 4.1 can be found in the Trichoptera World Checklist (Morse, 2015); complete bibliographic references generally are available in the Trichoptera Literature Database (Holzenthal et al., 2015). Table 4.1 also provides for each species the standard 2-letter postal codes for states and provinces of the U.S.A. and Canada where species have been reported in publications and publicly accessible M.S. theses and Ph.D. dissertations; literature citations for detailed collection records for states and provinces are available in Trichoptera Nearctica (Rasmussen & Morse, 2014) and complete bibliographic details generally are available for those citations from the Trichoptera Literature Database (Holzenthal et al., 2015). Among those references are published checklists for the caddisfly species of the following southeastern states: Alabama (Lago & Harris, 1987, 1991; Harris & Lago, 1990; Harris et al., 1991; Hicks & Haynes, 2000); Florida (Blickle, 1962; Gordon 1984; Pescador et al., 1995; Harris et al., 1982b, 2012); Georgia, North Carolina, and South Carolina (Gordon & Wallace, 1975); Kentucky (Resh, 1975; Haag & Hill, 1983; Thoeny & Batch, 1983; Phillippi & Schuster, 1987; Floyd & Schuster, 1990; Floyd, 1992; Houp & Schuster, 1997; Houp, 1999; Hamilton et al., 2002; Etnier et al., 2006); Mississippi (Harris et al., 1982a; Holzenthal et al., 1982; Lago et al., 1982; Etnier, 2010; Harrison & Morse, 2012); North Carolina (Banks, 1908; Carpenter, 1933; Denning, 1950; Armitage & Tennessen, 1984; Lenat, 1987; Lenat & Penrose, 1987; Lenat et al., 2010; Beaty, 2010); North and South Carolina (Unzicker et al., 1982; Morse et al., 1989; Floyd et al., 1997); South Carolina (Morse et al., 1980; Smock, 1988; Floyd & Morse, 1993; Floyd et al., 1993; Biondi, 2010); and Tennessee

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(Carpenter, 1933; Edwards, 1955, 1956, 1966; Chandler, 1978; Etnier & Schuster, 1979; Hamilton et al., 2002; Etnier et al., 1998, 2006; Wiggins et al., 2001; Vlach et al. 2010). An overview of insect morphology (Cummins et al., 2008) and of caddisfly larval morphology (Wiggins & Currie, 2008) as mentioned and illustrated in the following key is found in the reference by Merritt et al. (2008b). A more detailed account was provided by Wiggins (1996). The head, thorax, and abdomen of a caddisfly larva are always readily distinguishable (Fig. 4.01). General features of the sclerotized head of a larva include a pair of lateral genae separated anterodorsally by a frontoclypeal apotome, or frontoclypeus, posterodorsally by a coronal suture, and ventrally or anteroventrally by a ventral apotome, collectively constituting a head capsule. The posterodorsal regions of the genae sometimes are called parietals and the posteroventral regions postgenae. Pairs of primary setae no. 1–6 occur on the frontoclypeus and no. 7–18 on the genae (Figs. 4.02, 4.03). Various numbers and shapes of secondary setae may also occur on the head. A pair of usually short and inconspicuous antennae anterior to a pair of clusters of stemmata, or eyes, occur anterolaterally and somewhat dorsally on the genae. Small patches of color called muscle scars may be either darker or lighter than the background colors of the head capsule, indicating locations of muscles that are attached to the inner walls of the sclerotized head. Mouthparts consist of an anterodorsal labrum, a pair of horizontally moving mandibles, followed by a pair of maxillae with apical palps, and a ventral, spinneret-bearing labium.

Fig. 4.01

Fig. 4.02

Fig. 4.03

Figures 4.01–4.03, hypothetical larva of Trichoptera. 4.01, larva, left lateral. 4.02, head, dorsal. 4.03, head, ventral.

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The caddisfly larval thorax is always 3-segmented, with a pair of segmented legs on each of the prothorax, mesothorax, and metathorax. Dorsally, the pronotum is always sclerotized with a medial, longitudinal mid-dorsal ecdysial line (Fig. 4.04). The mesonotum and metanotum can be membranous or with two to four pairs of sclerites. Pairs of primary setae occur singly or in groups that are localized in setal areas, with setal area 1 (sa1) anteromedial, setal area 2 (sa2) posteromedial and more widely separated than the sa1s, and setal area 3 (sa3) lateral. Laterally a thoracic segment has on each side an anterior episternum and posterior epimeron, separated from each other by a pleural suture. On the prothorax, a foretrochantin of unknown function projects anterad on each side, being fused with the episternum or separated from it by a suture. Ventrally, the prosternum has one or more sclerites and, in several of the case-making families, a single, median, finger-like prosternal horn of unknown function curved anterad. The mesosternum and metasternum may also have sclerites and filamentous gills. Thoracic legs are always 5-segmented, each including a basal coxa, a trochanter (usually secondarily subsegmented), a femur, a tibia (generally with 1 or 2 apical tibial spurs), and a tarsus terminated by a single tarsal claw bearing a basal seta (Fig. 4.05). The arrangement of major and minor setae (determined by length and thickness) and combs on the leg segments may have diagnostic value.

Fig. 4.04

Fig. 4.05

Figure 4.04–4.05, hypothetical larva of Trichoptera. 4.04, thorax, dorsal. 4.05, left foreleg, posterolateral. The abdomen of a caddisfly larva is always 10-segmented, the segments commonly numbered with Roman numerals (Fig. 4.01). Segment I in most case-making families bears a median dorsal spacing hump and a pair of lateral spacing humps to allow passage of oxygenated water through the case; these humps are extended or retracted in preserved specimens, so that the evidence for their presence may be only some folds at these positions in the larva’s integument. Diagnostic sclerites often are associated with these humps. Sclerites may also be present on other abdominal segments, especially dorsally on segment IX. Some sclerites may be difficult to detect if their color is similar to surrounding membrane, but their presence and shape can be ascertained by being slightly harder and shinier than the membrane. On each

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side of some abdominal segments of some families are a lateral fringe (or lateral line) of bifid or single filaments and/or lateral tubercles (also called bifid processes or forked lamellae) (Fig. 4.06). Round or oval patches of osmoregulatory chloride epithelium may occur ventrally and sometimes also dorsally and occasionally laterally above the lateral fringe and lateral tubercles on each side; each of these is marked by a very fine line of sclerotization. Slender tracheal gills sometimes are present; each gill can be single (single gills in Figs. 4.01 and 4.06), multiple (arising from a common base), or branched from a central stalk and occur either anteriorly or posteriorly in dorsomesal, dorsolateral, ventrolateral, or ventromesal positions on a given segment (Figs. 4.01, 4.06). Osmoregulatory anal papillae that resemble tracheal gills may be everted from the anus. Segment X is much reduced and bears a pair of anal prolegs. Prolegs may be long, separate, and independently mobile on membranous tubular projections (Rhyacophilidae, Hydrobiosidae, and families of retreat makers, as in Fig. 4.31), shortened and partially fused with the body of segment X (Glossosomatidae and some genera of Hydroptilidae, as in Fig. 4.30), or nearly completely fused with segment X (families of case makers and most genera of Hydroptilidae) (Figs. 4.01 and 4.07). Recognizable in all of these conditions is a dorsolateral “lateral sclerite,” a ventral sole plate, and an anal claw that sometimes has a dorsal accessory hook or ventral teeth (Fig. 4.07).

Fig. 4.06

Fig. 4.07

Figures 4.06–4.07, hypothetical larva of Trichoptera. 4.06, abdominal segment IV, left lateral. 4.07, posterior end of abdomen, left lateral. The key also includes notes about the larval retreat or case and the distribution of each species. As in previous chapters, the key couplets and figures are numbered sequentially from the beginning to the end of the key. Figures generally face the page on which they are first mentioned, rarely appearing on the preceding page. Illustrations to assist with identification of families and genera in this key are mainly copyrighted drawings by co-author Dr. Ralph W. Holzenthal, University of Minnesota, first provided by Morse & Holzenthal (1984, 1996, 2008). Other illustrations are in the public domain or are re-drawn from copyrighted publications with permission of the authors and copyright holders. References to the sources of these illustrations are indicated in the legends for the plates of figures. I am especially grateful to the authors referenced in the figure captions and to the following copyright holders for permission to redraw published, copyrighted illustrations: American Entomological Society (Entomological News), Center for Systematic Entomology (Insecta Mundi), Entomological Society of America (Annals), Entomological Society of Washington (Proceedings), Society for Freshwater Science (Journal of the North American Benthological Society), Georgia Entomological Society (Journal), Illinois Natural History Survey (Bulletin), Kansas Entomological Society (Journal), Michigan Entomological Society (Great Lakes Entomologist), New York Entomological Society (Entomologica Americana), NRC Research Press (Canadian Journal of Zoology),

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Ohio Biological Survey (Bulletin and Proceedings of the 8th International Symposium on Trichoptera), ProQuest LLC (M.S. thesis UMI 1492638), Royal Ontario Museum (Contributions and Life Sciences Miscellaneous Publications), Taylor & Francis (Aquatic Insects), and University of Toronto Press [Larvae of the North American Caddisfly Genera (Trichoptera)]. Co-author Dr. O. Yadamsuren prepared most of the redrawn figures using Adobe® Illustrator® CC software (and earlier versions). We are to Dr. David Etnier, Dr. Andy Rasmussen, and an anonymous reviewer who provided many useful recommendations for refining this chapter. Nevertheless, remaining errors and shortcomings are entirely ours. We warmly welcome recommendations for improving future versions. Hundreds of students have used earlier drafts of the following keys in Morse’s classes at Clemson University (Clemson, South Carolina) and Highlands Biological Station (Highlands, North Carolina) during the past 42 years and provided insight about the best ways to represent the diagnostic characters described and illustrated here—We are grateful to these institutions and to the students for their contributions to this chapter. Indeed, it is mostly for these students in their subsequent careers and for their successors that this project was undertaken.

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Table 4.1. List of families, genera, and species of Trichoptera of the Southeastern U.S.A. (AL, FL, GA, KY, MS, NC, SC, TN). # = larva described at least tentatively and included in the accompanying key. * = occurring in one or more of the southeastern states, + = occurring in one or more states adjacent to the Southeast and likely to be found in the southeastern states, Each species name is followed by the name of its original author(s) and the date of validating publication. Where the author(s) and date are in parentheses, the species name is now associated with a genus other than the one with which it was originally described and the name of the original genus is indicated. The 2-letter postal code is provided for each state and province of the U.S.A. and Canada in which the species has been reported in published literature. Indented names of species are considered synonyms of the unindented species name immediately preceding; the author and date of publication of the synonymy are in parentheses.

Apataniidae

# * Apatania incerta (Banks, 1897), Enoicyla— CT, MA, NC, NH, NY, PA, QC, SC, TN, VA, WI; Apatania blacki Sykora & Weaver, 1978 (Flint, 2007); Radema praevolans Morse, 1971 (Flint, 2007); Radema rossi Morse, 1971 (Flint, 2007). # * Manophylax altus (Huryn & Wallace, 1984), Madeophylax—NC, WV. # * Manophylax butleri Schuster, 1997—KY, TN, WV.

Beraeidae

# * Beraea gorteba Ross, 1944—GA.

Brachycentridae

# + Adicrophleps hitchcocki Flint, 1965—CT, PA, MD, NY, VA, WV.

# * Brachycentrus (Sphinctogaster) appalachia Flint, 1984—GA, MD, ME, NB, NC, NH, NJ, NS, NY, ON, PA, SC, TN, VA, WV. # * Brachycentrus (Sphinctogaster) chelatuse Ross, 1947—AL, FL, GA, NC, SC. # * Brachycentrus (Sphinctogaster) etowahensis Wallace, 1971—GA, NC, TN. # * Brachycentrus (Sphinctogaster) incanus Hagen, 1861—DC, MD, MI, NC, NJ, NY, ON, PA, QC, VA, WI.

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Brachycentridae continued

# * Brachycentrus (Sphinctogaster) lateralis (Say, 1823), Phryganea—AR, IL, IN, KY, MD, ME, MI, MO, NC, NH, NY, ON, PA, QC, SC, TN, VA, VT, WI, WV; Sphinctogaster lutescens Provancher, 1877 (Milne, 1936). # * Brachycentrus (Sphinctogaster) lunatus Harrington & Morse, 2004— NC, SC. # * Brachycentrus (B.) nigrosoma (Banks, 1905), Sphinctogaster—AL, GA, KY, MD, ME, NC, NY, PA, SC, TN, VA, WV; Brachycentrus adelus Ross, 1947 (Flint, 1984); Brachycentrus notabulus Milne, 1936 (Flint, 1966, 1984). # * Brachycentrus (Sphinctogaster) numerosus (Say, 1823), Phryganea—AL, AR, CO, CT, DE, FL, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NY, OH, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV. # + Brachycentrus (Sphinctogaster) solomoni Flint, 1984—NY, PA, QC, VA, VT, WV. # * Brachycentrus (Sphinctogaster) spinae Ross, 1948—GA, NC, SC, TN, VA. # * Micrasema bennetti Ross, 1947—GA, KY, NC, SC, TN, VA, WV. # * Micrasema burksi Ross & Unzicker, 1965—GA, NC, NY, SC, TN, VA.

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Brachycentridae continued

Dipseudopsidae continued

Calamoceratidae

Glossosomatidae

# * Micrasema charonis Banks, 1914—AL, CT, KY, MA, MD, NB, NC, NF, NS, NY, PA, QC, SC, TN, VA, VT, WV. # * Micrasema florida Chapin & Morse, 2011—AL, FL. # * Micrasema rickeri Ross & Unzicker, 1965—GA, NC, SC, TN. # * Micrasema rusticum (Hagen, 1868), Dasystoma—AL, AR, FL, GA, IL, IN, KY, MA, MB, ME, MI, MN, MO, NB, NC, NH, NS, NY, OH, OK, ON, PA, QC, SC, SK, TN, VA, VT, WI, WV; Micrasema falcatum Banks, 1914 (Ross, 1938). # * Micrasema scotti Ross, 1947—AL, GA, IN, KY, TN, VA, WV. # * Micrasema sprulesi Ross, 1941—CT, MA, ME, NB, NC, NH, NY, ON, PA, QC, SC, VA. # * Micrasema wataga Ross, 1938—AL, AR, CT, FL, GA, IL, KY, LA, MA, MB, MD, ME, MI, MN, MS, NB, NC, NF, NH, NS, NY, OH, OK, ON, PA, QC, SC, SK, TN, VA, VT, WI.

# * Anisocentropus (Anisocentropus) pyraloides (Walker, 1852), Notidobia—AL, FL, GA, DE, KY, MS, NC, NJ, SC, TN, VA. # * Heteroplectron americanum (Walker, 1852), Sericostoma—AL, CT, DE, FL, GA, KY, MA, ME, NC, NH, NJ, NY, PA, QC, SC, TN, VA, VT; Ganonema nigrum Lloyd, 1915 (Milne, 1936).

Dipseudopsidae

* Phylocentropus auriceps (Banks, 1905), Plectrocnemia—GA, NC, SC, TN, VA; Phylocentropus rabilis Milne, 1936 (Ross, 1944).

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# * Phylocentropus carolinus Carpenter, 1933—AL, CT, DE, FL, GA, KY, LA, ME, MS, NC, NH, NY, ON, QC, SC, TN, VA. * Phylocentropus harrisi Schuster & Hamilton, 1984—AL, TX. # * Phylocentropus lucidus (Hagen, 1861), Polycentropus—AL, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, MD, ME, MS, NB, NC, NH, NJ, NS, NY, OH, ON, PA, QC, RI, SC, TN, VA, VT, WV. # * Phylocentropus placidus (Banks, 1905), Holocentropus—AL, AR, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NH, NJ, NS, NY, OH, OK, ON, PA, QC, RI, SC, TN, TX, VA, VT, WI, WV; Phylocentropus hansoni Root, 1965 (Schuster & Hamilton, 1984); Phylocentropus irroratus Navás, 1934 (Schuster & Hamilton, 1984); Phylocentropus maximus Vorhies, 1909 (Milne, 1936).

* Agapetus alabamensis Harris, 1986—AL. * Agapetus aphallus Etnier, Baxter, & Parker, 2010, in Etnier, Parker, Baxter, & Long, 2010–TN. * Agapetus avitus Edwards, 1956—AL, KY, MS, TN. * Agapetus baueri Etnier, Parker, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–NC, TN, VA. * Agapetus crasmus Ross, 1939—AL, TN. # * Agapetus diacanthus Edwards, 1956—TN. * Agapetus flinti Parker, Etnier, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–NC. * Agapetus gelbae Ross, 1947—AL, IN, TN. * Agapetus harrisi Etnier, Parker, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–AL.

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* Agapetus hesperus Etnier, Baxter, & Parker, 2010, in Etnier, Parker, Baxter, & Long, 2010–AL. * Agapetus hessi Leonard & Leonard, 1949—AL, KY, ME, MI, NB, NC, NS, ON, QC, VA, WI, WV. * Agapetus ibis Etnier, Baxter, & Parker, 2010, in Etnier, Parker, Baxter, & Long, 2010–AL, GA, TN. # * Agapetus illini Ross, 1938—AR, ID, IL, IN, KS, KY, MO, OK, TN. * Agapetus iridis Ross, 1944—AL, CT, DE, MA, ME, NB, NC, NH, NY, PA, QC, SC, TN, VA. # * Agapetus jocassee Morse, 1989, in Morse, Hamilton, & Hoffman, 1989–GA, NC, SC, TN. * Agapetus kirchneri Parker, Etnier, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–KY, TN, VA. * Agapetus meridionalis Etnier, Parker, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–GA. # * Agapetus minutus Sibley, 1926—DE, KY, MA, NY, NC, ON, PA, TN, VA, WV. * Agapetus pegram Etnier, Baxter, & Parker, 2010, in Etnier, Parker, Baxter, & Long, 2010–TN. * Agapetus pinatus Ross, 1938—AL, CT, GA, MA, ME, MS, NB, NC, NH, NS, NY, PA, QC, SC, TN, VA, VT. * Agapetus ruiteri Parker, Etnier, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–TN. * Agapetus spinosus Etnier & Way, 1973— AL, SC, TN. * Agapetus stylifer Etnier, Baxter, & Parker, 2010, in Etnier, Parker, Baxter, & Long, 2010–KY, TN. * Agapetus tomus Ross, 1941—AL, GA, KY, MI, MN, NC, NJ, OH, PA, TN, VA. * Agapetus tricornutus Etnier, Parker, & Baxter, 2010, in Etnier, Parker, Baxter, & Long, 2010–AL. * Agapetus vireo Ross, 1941—AL, GA, TN.

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# * Agapetus walkeri Betten & Mosely, 1940– AL, CT, GA, MA, MD, ME, MN, NB, NC, NH, NS, NY, ON, PA, QC, SC, TN, VA, WV; Agapetus rossi Denning, 1941, MN (Etnier, Parker, Baxter, & Long, 2010). # * Culoptila plummerensis Blahnik & Holzenthal, 2006—MD, ME, NC, NY, VA. # * Glossosoma (Synafophora) intermedium (Klapálek, 1892), Mystrophora— AB, BC, IA, ID, IL, IN, KY, MB, ME, MI, MN, MO, MT, NF, NY, OH, ON, PA, QC, VA, VT, WI, YT. # * Glossosoma (Synafophora) lividum (Hagen, 1861), Tinodes—MA, ME, NC?, NH, NY, ON, QC, VA, VT; Glossosoma americanum Banks, 1897 (Ross, 1956). # * Glossosoma (Synafophora) nigrior Banks, 1911—AL, CT, DE, GA, IN, KY, LB, MA, MD, ME, MI, MN, MS, NB, NC, NF, NH, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV; Eomystra unica Denning, 1942 (Ross, 1944). # * Padunia jeanae (Ross, 1938), Protoptila— AL, GA, KY, NC, SC, TN, VA. * Protoptila alexanderi Ross, 1941—KY, TX. * Protoptila cahabensis Harris, 1989—AL. * Protoptila georgiana Denning, 1948—AL, GA, MD, NC, VA. * Protoptila lega Ross, 1941—AL, AR, IL, MO, MS, NB, NC, OH, OK, ON, VA, WI. # * Protoptila maculata (Hagen, 1861), Beraea ?—AL, AR, DC, IL, IN, KY, MA, MB, MD, ME, MI, MN, MO, NC, NH, NY, OH, OK, ON, PA, QC, TN, TX, VA, WI, WV; Protoptila expositionis Nimmo, 1966 (Schmid, 1982); Protoptila lloydi Mosely, 1934 (Milne, 1936).

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* Protoptila morettii Morse, 1990—NC, SC. * Protoptila palina Ross, 1941—AL, IN, KY, MA, MD, ME, NC, NH, NY, PA, SC, TN, VA, WV.

Goeridae

# * Goera calcarata Banks, 1899—AL, CT, DE, GA, KY, MA, ME, MN, NB, NC, NH, NJ, NS, NY, OH, PA, QC, SC, TN, VA, VT, WV. # * Goera fuscula Banks, 1905—GA, KY, MA, ME, NB, NC, NJ, NY, PA, QC, SC, TN, VA, VT. # * Goera stylata Ross, 1938—AL, CT, IN, KY, MA, MI, MN, NC, NH, NY, OH, ON, PA, TN, VA, WI, WV. * Goera townesi Morse, 1971—AL, GA, NC, SC. # * Goerita betteni Ross, 1962—KY, NC, OH, PA, TN, VA, WV. # * Goerita flinti Parker, 1999—NC, TN. # * Goerita semata Ross, 1938—MT, NC, TN, VA.

Helicopsychidae

# * Helicopsyche borealis (Hagen, 1861), Notidobia—AB, AL, AR, AZ, CA, CO, CT, DE, FL, GA, ID, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NF, NH, NJ, NM, NS, NT, NV, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY; Helicopsyche annulicornis Banks, 1904 (Betten, 1934; Ross, 1944); Va l vata arenifera L ea, 1834 (Johanson, 2002); Helicopsyche californica Banks, 1899 (Ross, 1944); Helicopsyche glabra Hagen, 1864 (Hagen, 1864; Ross, 1944); Paludina lustrica Say, 1821 (Johanson, 1995; Johanson, 1998; Opinion 1108). # * Helicopsyche paralimnella Hamilton, 1989—NC, SC.

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Hydropsychidae

# * Arctopsyche irrorata Banks, 1905—GA, NC, SC, TN, VA. # * Cheumatopsyche analis (Banks, 1903), Hydropsyche—AB, AL, AR, BC, CO, CT, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NF, NH, NJ, NS, NY, OH, OK, ON, OR, PA, PE, QC, SC, SD, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY; Hydropsyche pettiti Banks, 1908 (Ross, 1944; Hoffman & Parker, 1997). * Cheumatopsyche aphanta Ross, 1938— AR, IL, IN, KS, KY, ME, MI, MN, MO, NB, ND, NH, NY, OH, OK, PA, QC, TN, TX, VT, WI. * Cheumatopsyche bibbensis Gordon, Harris, & Lago, 1986—AL. * Cheumatopsyche burksi Ross, 1941—AL, AR, CT, FL, IL, IN, KY, LA, MS, NY, OK, TN, TX, VA, VT. * Cheumatopsyche cahaba Gordon, Harris, & Lago, 1986—AL. # * Cheumatopsyche campyla Ross, 1938— AB, AL, AR, AZ, BC, CA, CO, CT, DE, FL, GA, IA, ID, IL, IN, KS, KY, LB, MA, MB, ME, MI, MN, MO, MS, MT, NC, ND, NE, NF, NH, NJ, NM, NS, NV, NY, OH, OK, ON, OR, PA, QC, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY, YT. # * Cheumatopsyche edista Gordon, 1974— AL, FL, GA, SC. * Cheumatopsyche ela Denning, 1942—AL, DC, DE, GA, KY, MB, MD, ME, NC, NY, OH, ON, PA, QC, TN, VA, WV. # * Cheumatopsyche enigma Ross, Morse, & Gordon, 1971—AR, GA, MO, NC, PA, SC, TN, VA. # * Cheumatopsyche etrona Ross, 1941—GA, NC, SC, TN, VA. * Cheumatopsyche geora Denning, 1948— AL, CT, DE, GA, KY, MS, NC, OH, PA, SC, TN, VA.

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Hydropsychidae continued

* Cheumatopsyche gordonae Lago & Harris, 1983—FL. * Cheumatopsyche gracilis (Banks, 1899), Hydropsyche—AB, AL, AR, BC, CO, CT, KS, LB, MA, MB, ME, MI, MN, MO, MT, NC, ND, NF, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, UT, VA, VT, WI, WV, WY. * Cheumatopsyche gyra Ross, 1938—GA, KY, ME, NC, PA, SC, VA, WV. * Cheumatopsyche halima Denning, 1948— AR, KY, MA, ME, NB, NC, NY, OH, PA, QC, SC, VA, WV. # * Cheumatopsyche harwoodi Denning, 1949—AB, AL, CT, GA, IN, KY, MB, ME, NB, NC, NS, NY, OH, ON, PA, PE, RI, SC, TN, VA, WV. * Cheumatopsyche helma Ross, 1939—AL, AR, KY, ME, NC, NY, PA, TN, WV. * Cheumatopsyche kinlockensis Gordon, Harris, & Lago, 1986—AL. + Cheumatopsyche lasia Ross, 1938—AB, AR, AZ, CO, IA, IL, IN, KS, MN, MO, MT, ND, NE, NM, OH, OK, PA, SK, TX, WY. # * Cheumatopsyche minuscula (Banks, 1907), Hydropsyche—AL, AR, CT, DC, FL, GA, KS, KY, MA, MB, MD, ME, MN, MO, NC, ND, NH, NY, OH, OK, ON, PA, QC, SC, TN, VA, VT, WI, WV; Cheumatopsyche montrealensis Nimmo, 1966 (Gordon, 1974). * Cheumatopsyche oxa Ross, 1938—AB, AL, AR, BC, CT, GA, IL, IN, KS, KY, MB, ME, MI, MN, MO, MS, MT, NB, NC, NH, NY, OH, OK, ON, PA, QC, SC, SD, SK, TN, VA, VT, WI, WV, WY. * Cheumatopsyche parentum Gordon, 1974—MD, NC, VA. * Cheumatopsyche pasella Ross, 1941—AL, AR, CT, DE, FL, GA, IL, IN, KY, LA, MA, MD, ME, MN, MS, MT, NC, ND, NH, NJ, NY, OH, OK, ON, OR, PA, QC, RI, SC, TN, TX, VA, VT, WA, WI, WV.

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* Cheumatopsyche petersi Ross, Morse, & Gordon, 1971—AL, FL, MS. # * Cheumatopsyche pinaca Ross, 1941—AL, DE, FL, GA, KY, LA, MA, ME, MS, NC, NH, NJ, PA, RI, SC, TN, VA. # * Cheumatopsyche richardsoni Gordon, 1974—NC, SC. + Cheumatopsyche rossi Gordon, 1974— AR, KS, MO, OH, OK. # * Cheumatopsyche sordida (Hagen, 1861), Hydropsyche—AL, AR, CT, DC, GA, IL, IN, KY, LA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NH, NJ, NY, OK, ON, PA, QC, SC, TN, TX, VA, WI, WV. * Cheumatopsyche speciosa (Banks, 1904), Hydropsyche—AB, AR, CO, CT, DE, IA, IL, IN, KY, LB, MB, MD, MI, MN, MO, MT, NC, ND, NY, OH, OK, ON, PA, QC, SK, TN, VA, WI. # * Cheumatopsyche virginica Denning, 1949—AL, DE, FL, GA, LA, MS, NC, NJ, SC, VA. * Cheumatopsyche wrighti Ross, 1947— MA, ME, NC, NS, NY, PA, PE, TN, VA, WV. * Diplectrona marianae Reeves, 1999— GA, TN. # * Diplectrona metaqui Ross, 1970—AL, CT, GA, IL, IN, KY, MO, NC, OH, PA, SC, TN, VA, WV. # * Diplectrona modesta Banks, 1908—AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, MA, MD, ME, MI, MN, MO, MS, NC, NF, NH, NS, NY, OH, OK, ON, PA, QC, SC, SD, TN, TX, VA, VT, WI, WV. # + Diplectrona rossi Morse, 1990—LA. # * Homoplectra doringa (Milne, 1936), Diplectrona—AL, AR, IN, KY, MA, NC, NH, NY, TN, VA, WV; Aphropsyche aprilis Ross, 1941 (Flint, 1966).

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# * Homoplectra flinti Weaver, 1985—NC. # * Homoplectra monticola (Flint, 1965), Aphropsyche—NC, PA, VA, WV.

# * Hydropsyche aerata Ross, 1938—IL, IN, KY, MI, MO, OH, VT. * Hydropsyche alabama Lago & Harris, 1991—AL, FL. # * Hydropsyche alhedra Ross, 1939—AB, AK, BC, CO, CT, IA, KY, MA, ME, MB, MN, NC, ND, NY, ON, PA, QC, SD, SK, TN, VA, VT, WI, WV, WY, YT; Hydropsyche racona Denning, 1965 (Schefter, Wiggins, & Unzicker, 1986); Hydropsyche riola Denning, 1942 (Schefter, Wiggins, & Unzicker, 1986). # + Hydropsyche alternans (Walker, 1852), Philopotamus—AB, AK, BC, CT, IL, IN, MA, MB, ME, MI, MN, NF, NH, NJ, NS, NT, NY, OH, ON, OR, PA, QC, SK, VT, WI, YT; Hydropsyche codona Betten, 1934 (Milne, 1936, as synonym of H. slossonae; Ross, 1938, as synonym of H. recurvata); Philopotamus indecisus Walker, 1852 (McLachlan, 1863; Betten & Mosely, 1940); Hydropsyche slossonae var. recurvata Banks, 1914 (Hydropsyche recurvata is synonym of Hydropsyche slossonae according to Milne, 1936; Nimmo, 1981). # + Hydropsyche arinale Ross, 1938—AR, IL, IN, KS, MO, OK, ON, WI; Hydropsyche reiseni Denning, 1975 (Moulton & Stewart, 1996).. * Hydropsyche bassi Flint, Voshell, & Parker, 1979—TN, VA. # * Hydropsyche betteni Ross, 1938—AK, AL, AR, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, ND, NF, NH, NS, NY, OH, OK, ON, PA, PE, QC, SC, SK, TN, VA, VT, WI, WV.

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* Hydropsyche bidentata Denning, 1948—SC. # * Hydropsyche bronta Ross, 1938—AB, AR, CO, CT, DE, IA, IL, IN, KY, MA, MB, MD, MI, MN, MO, ME, MT, NB, NC, ND, NE, NH, NS, NY, OH, OK, ON, PA, QC, SC, SK, TN, VA, VT, WI, WV, WY. * Hydropsyche brunneipennis Flint & Butler, 1983—MD, PA, TN, VA, WV. # * Hydropsyche carolina Banks, 1938—GA, NC, SC, TN. * Hydropsyche catawba Ross, 1939—GA, NC, TN, VA. # * Hydropsyche cheilonis Ross, 1938—AL, IL, IN, KY, MI, NC, OH, PA, TN, VA, WI, WV. # * Hydropsyche cuanis Ross, 1938—AL, CT, IL, IN, KY, MA, MI, MN, OH, TN, VT, WI. # * Hydropsyche decalda Ross, 1947—AL, CT, DE, FL, GA, LA, NC, SC, TX, VA. # * Hydropsyche demora Ross, 1941—AL, GA, NC, TN. # * Hydropsyche depravata Hagen, 1861— AL, GA, IN, KY, MB, NC, OH, TN, VA, WV. # * Hydropsyche dicantha Ross, 1938—AL, DC, DE, GA, IN, KY, MD, MI, MN, NH, NY, OH, ON, PA, QC, TN, VA, WI, WV. # * Hydropsyche elissoma Ross, 1947—AL, FL, GA, LA, NC, SC. # * Hydropsyche etnieri (Schuster & Talak, 1977), Symphitopsyche—KY, TN, VA. * Hydropsyche fattigi Ross, 1941—AL, GA, NC, TN, VA. * Hydropsyche fenestra Lago & Harris, 2006—AL * Hydropsyche franclemonti Flint, 1992— CT, GA, NC, NY, QC, SC, TN, VA. # * Hydropsyche frisoni Ross, 1938—AL, CO, CT, IA, IL, IN, KS, KY, MI, MO, MN, NE, OH, TN, VA, WV.

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Hydropsychidae continued

# * Hydropsyche hageni Banks, 1905—AL, DC, IL, KY, MB, MD, MI, MN, NC, OH, ON, PA, QC, TN, VA, WI, WV. # + Hydropsyche hoffmani Ross, 1962—DE, MD, VA, WV. # * Hydropsyche incommoda Hagen, 1861— AL, AR, CO, FL, GA, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, MT, NC, ND, NE, NY, OH, OK, PA, QC, SC, SD, TN, TX, VA, WI, WV; Hydropsyche alvata Denning, 1949 (Korecki, 2006, synonymy debated); Hydropsyche bidens Ross, 1938 (Korecki, 2006, synonymy debated); Hydropsyche orris Ross, 1938 (Korecki, 2006, synonymy debated); Hydropsyche cornuta Ross, 1938 (preoccupied, replaced by H. orris by Ross, 1938). # * Hydropsyche leonardi Ross, 1938—DE, MI, NY, ON, PA, TN, VA, WI, WV. # * Hydropsyche macleodi Flint, 1965—GA, NC, TN, VA, WV. # * Hydropsyche mississippiensis Flint, 1972—AL, FL, KY, LA, MS, NC, SC, TN, TX, VA. # * Hydropsyche morosa Hagen, 1861—AB, AR, BC, CO, CT, DC, DE, GA, IA, ID, IL, IN, KS, KY, MA, MB, ME, MI, MN, MO, MT, NB, NC, ND, NH, NJ, NS, NT, NY, OH, OK, ON, PA, QC, SC, SK, TN, UT, VA, VT, WA, WI, WV, WY; Hydropsyche bifida Banks, 1905 (Schefter & Unzicker, 1984; Schefter & Wiggins, 1986); Hydropsyche chlorotica Hagen, 1861 (Ross, 1938). * Hydropsyche opthalmica Flint, 1965—DE, NC, PA, VA, WV. # * Hydropsyche patera Schuster & Etnier, 1978—KY, OH, TN.

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# * Hydropsyche phalerata Hagen, 1861—AL, DC, DE, FL, GA, IA, IL, IN, KY, KS, MA, MD, ME, MI, MN, MS, NC, NJ, NY, OH, ON, PA, QC, TN, VA, WI, WV. # + Hydropsyche piatrix Ross, 1938—AR, MO, ND, QC. # + Hydropsyche potomacensis Flint, 1965—VA. * Hydropsyche rotosa Ross, 1947—AL, TN, VA. # * Hydropsyche scalaris Hagen, 1861—AL, AR, CO, CT, DC, DE, GA, IL, IN, KS, MB, MD, ME, MN, MO, NC, NE, NJ, NM, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV, WY. # * Hydropsyche simulans Ross, 1938—AL, AR, CO, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, MI, MN, MO, MS, MT, NC, ND, NE, NY, OH, OK, ON, SC, TN, TX, VA, WI, WV, WY; Hydropsyche rossi Flint, Voshell, & Parker, 1979 (Korecki, 2006, synonymy debated). # * Hydropsyche slossonae Banks, 1905—AB, AR, BC, CO, CT, DE, IL, IN, KY, LB, MA, MB, ME, MI, MN, MO, MT, NB, NC, ND, NF, NH, NJ, NS, NT, NY, OH, ON, PA, QC, SC, SD, SK, TN, VA, VT, WI, WV, WY. # * Hydropsyche sparna Ross, 1938—AL, CT, DE, GA, IL, IN, KS, KY, LB, MA, MB, MD, ME, MI, MN, MS, NB, NC, NF, NH, NJ, NS, NY, OH, ON, PA, PE, QC, SC, TN, VA, VT, WI, WV. # * Hydropsyche valanis Ross, 1938—IA, IL, IN, KS, KY, MI, MN, NY, OH, PA, VT, WI. # * Hydropsyche ventura Ross, 1941—CT, KY, MA, ME, MN, NC, NF, NY, OH, ON, PA, QC, TN, VA, VT, WV. # * Hydropsyche venularis Banks, 1914—AL, CT, DC, GA, IL, KY, MD, MO, NC, NY, PA, SC, TN, VA, VT, WI, WV.

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# * Hydropsyche walkeri Betten & Mosely, 1940—AB, CT, DE, MA, MB, ME, MI, MN, NC, ND, NH, NY, OH, ON, PA, QC, SK, VA, VT, WI, WV; Hydropsyche maculicornis Walker, 1852 (preoccupied by Pictet, 1834).

# * Macrostemum carolina (Banks, 1909), Macronema—AL, AR, DE, FL, GA, IL, IN, LA, MD, MO, MS, NC, NJ, NY, OK, PA, SC, TN, TX, VA. # * Macrostemum transversum (Walker, 1852), Hydropsyche—AL, GA, IN, MD, MS, OH, VA; Macronema polygrammatum McLachlan, 1871 (Milne, 1936). # * Macrostemum zebratum (Hagen, 1861), Macronema—AL, CT, DC, DE, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, NC, ND, NH, NJ, NY, OH, ON, PA, QC, SC, TN, UT, VA, VT, WI, WV; Macronema australe McLachlan, 1862 (Neboiss, 1984); Monopseudopsis inscriptus Walker, 1852. * Oropsyche howellae Ross, 1941—NC. # * Parapsyche apicalis (Banks, 1908), Arctopsyche—CO, CT, MA, ME, MI, MN, NC, NF, NH, NS, NY, OH, ON, PA, PE, QC, RI, SC, TN, VA, VT, WI, WV. # * Parapsyche cardis Ross, 1938, —GA, KY, NC, SC, TN, VA. * Potamyia flava (Hagen, 1861), Macronema—AL, AR, CO, DE, FL, GA, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, MT, NC, ND, NE, NY, OH, OK, ON, PA, QC, SC, SD, TN, TX, VA, WI, WV; Hydropsyche kansensis Banks, 1905 (Milne, 1936).

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Hydroptilidae

# * Agraylea multipunctata Curtis, 1834—BC, CO, CT, DE, IL, IN, KY, MA, ME, MB, MI, MN, MT, NB, ND, NE, NH, NM, NS, NT, NY, OH, ON, OR, PA, QC, SD, TN, UT, VA, VT, WA, WI, WY, YT; Allotrichia flavida Banks, 1907 (Ross, 1944); Agraylea fraterna Banks, 1907 (Milne, 1936); Agraylea multiguttata Uljanin, 1869 (Fischer, 1961); Allotrichia signata Banks, 1904 (Milne, 1936). # * Dibusa angata Ross, 1939—AL, AR, GA, IN, KY, MA, ME, NC, NH, OH, OK, ON, PA, SC, TN, VA, VT, WV. * Hydroptila acadia Ross, 1941—AL, FL, NS, VA. * Hydroptila ajax Ross, 1938—AZ, CA, CO, FL, IA, ID, IL, IN, KS, KY, MB, MN, MO, MT, NE, NV, NY, OH, OK, OR, PA, TX, UT, VA, WA, WI, WV, WY. * Hydroptila alabama Harris & Kelley, 1984—AL, FL, GA, ME, NC, NY, PA, TN, TX. * Hydroptila albicornis Hagen, 1861—AR, IL, IN, MB, ME, MN, MO, NY, OH, OK, ON, QC, TN, WI. * Hydroptila amoena Ross, 1938—AL, AR, IL, IN, KY, MN, MO, NH, OH, OK, PA, QC, SC, TN, VA, WI. * Hydroptila ampoda Ross, 1941—CT, KY, ME, MN, NB, NC, NH, NS, NY, PA, QC, TN. * Hydroptila angusta Ross, 1938—AL, AR, CO, IA, IL, IN, KS, KY, MB, MO, MS, NC, NM, OH, OK, PA, SC, TN, TX, WY. + Hydroptila anisoforficata Parker & Voshell, 1979—VA. * Hydroptila apalachicola Harris, Pescador & Rasmussen, 1998—FL.

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Hydroptilidae continued

# * Hydroptila armata Ross, 1938—AL, AR, IL, IN, FL, KS, KY, MB, MD, MI, MN, MO, MS, NC, NH, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, WI, WV. * Hydroptila artesa Mathis & Bowles, 1990—AR, MO, TN, VA. * Hydroptila auriscuspa Harris, Rasmussen, & Denson, 2012–FL. * Hydroptila berneri Ross, 1941—AL, AR, FL, LA, MN, NC, QC, SC, TX, WI. * Hydroptila bribriae Harris, 2002—FL. + Hydroptila broweri Blickle, 1963—AR, ME, MO, NH. * Hydroptila callia Denning, 1948–AL, CO, MI, MN, NC, NH, NS, OH, PA, QC, SC, TN, VA, WI, WY. * Hydroptila carolae Holzenthal & Kelley, 1983—SC. * Hydroptila chattanooga Frazer & Harris, 1991—AL, ME, OH, PA, TN. * Hydroptila cheaha Harris, 1991—AL, KY. * Hydroptila chelops Harris, 1985—AL. * Hydroptila circangula Harris, 1985— AL, FL. # * Hydroptila consimilis Morton, 1905—AB, AL, AR, AZ, BC, CO, CT, DE, IA, ID, IL, IN, KS, KY, MB, ME, MI, MN, MO, MT, NB, ND, NH, NM, NT, NV, NY, OH, OK, ON, OR, PA, QC, SC, TN, TX, UT, VA, VT, WA, WI, WY. * Hydroptila cottaquilla Harris, 1994—AL; Hydroptila setigera Harris, 1986 (preoccupied by Wells, 1984). * Hydroptila coweetensis Huryn, 1985—AL, KY, NC, VA. * Hydroptila cretosa Harris, 1985—AL, MS. * Hydroptila decia Etnier & Way, 1973— AL, KY, TN; Hydroptila choccolocco Harris, 1985 (Harris & Etnier, 1994). * Hydroptila delineata Morton, 1905—AL, AR, IN, KY, ME, MN, NC, NH, NS, NY, OH, PA, QC, SC, TN, VA, WV. + Hydroptila dentata Ross, 1938—ME, NH, NY, PA, VA.

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* Hydroptila disgalera Holzenthal & Kelley, 1983—AL, FL, SC. * Hydroptila ebroensis Harris, Rasmussen, & Denson, 2012–FL. * Hydroptila eglinensis Harris, 2002—FL. * Hydroptila englishi Hamilton, 1989— NC, SC. + Hydroptila eramosa Harper, 1973—ON, VA. * Hydroptila fiskei Blickle, 1963—KY, ME, NC, NH, NY, PA, TN, VA. + Hydroptila fowlesi Harris & Sykora, 1996—OH, WV. * Hydroptila fuscina Harris, 1985—AL. * Hydroptila grandiosa Ross, 1938—AL, AR, IL, IN, KS, KY, MB, MN, MO, MS, OH, OK, PA, TN, TX, VA, WI, WV. * Hydroptila gunda Milne, 1936—AL, CT, GA, IN, FL, KY, ME, MS, NC, NH, NJ, OH, PA, QC, SC, TN, VA; Hydroptila dodgei Denning, 1947 (Blickle, 1979). * Hydroptila hamata Morton, 1905—AL, AR, AZ, CA, CO, CT, FL, GA, ID, IL, IN, KY, LA, MB, ME, MI, MN, MO, MS, MT, NC, NH, NM, NY, OH, OK, ON, OR, PA, QC, SC, TN, TX, UT, VA, VT, WA, WI, WV, WY. * Hydroptila hamiltoni Harris, 2002—FL. * Hydroptila holzenthali Sykora & Harris, 1994—MS. * Hydroptila homochitta Harris & Sykora, 1996—MS. * Hydroptila howelli Houp, Houp, & Harris, 1998—KY. * Hydroptila icona Mosely, 1937—AZ, CA, CO, FL, HI, NM, OK, TX. * Hydroptila jackmanni Blickle, 1963—AL, IN, KY, MB, ME, MN, NH, OH, PA, QC, TN, VA, WI, WY. * Hydroptila kuehnei Houp, Houp, & Harris, 1998—KY, TN. * Hydroptila lagoi Harris, 1985—AL. * Hydroptila latosa Ross, 1947—AL, FL, GA, SC.

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* Hydroptila lennoxi Blickle, 1969—AL, KY, NH, OH, VA. * Hydroptila licina Frazer & Harris, 1991— AL, GA. * Hydroptila lloganae Blickle, 1961—FL, LA, NC, SC, TX; Hydroptila morsei Sykora & Harris, 1994 (Etnier & Baxter, 1999). * Hydroptila lonchera Blickle & Morse, 1954—AL, NH, NY, OH, VA. * Hydroptila maculata Banks, 1904—DC, FL, ME, NC, NH, VA; Hydroptila transversa Banks, 1907 (Ross, 1938). * Hydroptila mcgregori Harris & Huryn, 2000–AL, TN. + Hydroptila metoeca Blickle & Morse, 1954—DE, ME, MN, NF, NH, NJ, NY, OH, PA, VA. * Hydroptila metteei Harris, 1991—AL, FL. * Hydroptila micropotamis Harris, 1989— AL, GA. * Hydroptila molsonae Blickle, 1961—AL, FL, LA, MS. * Hydroptila murtlei Harris, Rasmussen, & Denson, 2012–FL. * Hydroptila novicola Blickle & Morse, 1954—AL, FL, GA, LA, ME, MN, MS, NH, NY, QC, TN, TX. * Hydroptila oakmulgeensis Harris, 1985—AL. * Hydroptila okaloosa Harris, 2002—FL. * Hydroptila oneili Harris, 1985—AL, AR, GA, KY, TN. * Hydroptila paralatosa Harris, 1985— AL, FL. * Hydroptila paramoena Harris, 1985—AL, GA, KY, TN. * Hydroptila parastrepha Kelley & Harris, 1983—AL, FL. * Hydroptila paraxella Harris & Armitage, 2011—KY, OH. * Hydroptila patriciae Harris, 1985—AL, TN. # * Hydroptila perdita Morton, 1905—AL, AR, DE, IL, IN, KS, KY, MB, MD, MI, MN, MO, NH, NY, OH, OK, ON, PA, QC, TN, VA, WI, WV.

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* Hydroptila poirrieri Holzenthal & Kelley, 1983—LA, MO, MS. * Hydroptila quinola Ross, 1947—AL, AR, CT, FL, GA, KY, ME, MN, MS, NC, NH, ON, PA, QC, SC, TN, TX, VA. * Hydroptila recurvata Harris & Kelley, 1984— AL. * Hydroptila remita Blickle & Morse, 1954— AL, AR, FL, KY, LA, ME, MS, NC, NH, NJ, PA, SC, TN, TX, VA. * Hydroptila roberta Hamilton & Holzenthal, 1986—GA. * Hydroptila sandersoni Mathis & Bowles, 1990—AL, AR, KY, MO, OH, OK, TN. * Hydroptila santarosa Harris, Rasmussen, & Denson, 2012–FL. * Hydroptila sarahae Harris, 2002—FL. * Hydroptila scheiringi Harris, 1986— AL, FL. * Hydroptila scolops Ross, 1938—IL, KS, KY, MB, MN, TX, WI. * Hydroptila spatulata Morton, 1905—AL, AR, DE, IL, IN, KY, MB, MD, MI, MN, MO, NH, NY, OH, OK, PA, QC, TN, VA, WI. * Hydroptila spinata Blickle & Morse, 1954—AL, ME, NH, NY, PA, QC, TN, VA. * Hydroptila strepha Ross, 1941—AR, FL, ME, MN, MO, MS, NH, OH, PA, SC, WI. * Hydroptila sykorai Harris, 2002—FL. * Hydroptila talladega Harris, 1985— AL, GA, KY, NC, OH, PA, SC, TN, VA. * Hydroptila tortosa Ross, 1938—ME, MN, NC, NH, SC, VA. * Hydroptila tridentata Holzenthal & Kelley, 1983—SC. * Hydroptila tusculum Ross, 1947—AL, LA, MO, MS, TN. * Hydroptila vala Ross, 1938—AL, AR, IL, IN, KY, MB, OH, OK, TN, VA.

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Hydroptilidae continued

* Hydroptila valhalla Denning, 1947—AL, ME, MI, MN, NH, OH, TN, WI. * Hydroptila virgata Ross, 1938—AL, AR, DE, IL, KY, MB, MN, MO, NH, OH, OK, QC, TN, VA, WI. * Hydroptila wakulla Denning, 1947—FL. * Hydroptila waskesia Ross, 1944—AL, KY, MN, OH, PA, QC, SK, TN. * Hydroptila waubesiana Betten, 1934—AL, AR, CO, DE, FL, GA, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, MT, NC, ND, NJ, NY, OH, OK, ON, PA, QC, SC, SK, TN, TX, VA, WI. * Hydroptila wetumpka Harris, 1991— AL, FL. * Hydroptila xella Ross, 1941—AL, IL, NH, TN. # * Ithytrichia clavata Morton, 1905—AB, AR, AZ, BC, CA, FL, IL, KS, MB, ME, MI, MN, MO, MT, NH, NY, OH, OK, ON, PA, QC, TN, TX, UT, VA, WI, WY. * Ithytrichia mazon Ross, 1944—AR, IL, KY, OH, OK. # * Leucotrichia pictipes (Banks, 1911), Orthotrichia—AL, AZ, CA, CO, CT, DE, ID, IL, KS, KY, MA, MI, MN, MO, MT, NC, NM, NV, NY, OR, PA, TN, UT, VA, VT, WA, WI, WV, WY.

# * Mayatrichia ayama Mosely, 1937—AB, AL, AR, AZ, CO, FL, GA, IA, IL, IN, KS, KY, MB, ME, MI, MN, MO, MS, MT, NB, NC, NH, NT, NV, NY, OH, OK, ON, PA, QC, SC, SK, TN, TX, UT, VA, VT, WI, WY. * Mayatrichia tuscaloosa Harris & Sykora, 1996—AL. * Metrichia sp.—KY.

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* Neotrichia alabamensis Kelley & Harris, 1983—AL, FL, MS. * Neotrichia armitagei Harris, 1991—AL, FL, GA, MS, TX. * Neotrichia collata Morton, 1905—AL, AR, IL, KY, ME, NY, SC, UT, VT. * Neotrichia falca Ross, 1938—IL, KS, MN, OH, SC, WI. * Neotrichia mentonensis Frazer & Harris, 1991—AL, GA. * Neotrichia minutisimella (Chambers, 1873), Cyllene—AL, AR, FL, GA, IL, IN, KS, KY, LA, MB, MN, MO, MS, NC, OK, SC, TX. * Neotrichia mobilensis Harris, 1985— AL, TX. * Neotrichia okopa Ross, 1939—AL, AR, AZ, CA, CO, FL, IL, IN, KS, KY, MB, ME, MN, MO, NH, NY, OH, OK, ON, OR, PA, QC, TX, WI. * Neotrichia rasmusseni Harris & Keth, 2002—FL. * Neotrichia riegeli Ross, 1941—AR, IL, KY, OK. * Neotrichia sepulga Harris, 1991—AL. * Neotrichia vibrans Ross, 1938—AL, AR, FL, GA, IL, KS, KY, ME, MN, MO, MS, NH, NY, OH, OK, SC, TN, TX, VA, WI, WV; Neotrichia ranea Denning, 1947 (Ross, 1948). * Ochrotrichia anisca (Ross, 1941), Polytrichia—AR, IL, KS, KY, MO, OK, TX. * Ochrotrichia apalachicola Harris, Pescador & Ramussen, 1998—FL. * Ochrotrichia arva (Ross, 1941), Polytrichia—AL, AR, KY, MI, MO, OH, TN, VA. # * Ochrotrichia confusa (Morton, 1905), Ithytrichia—AL, FL, KY, NC, NY, OH, ON, PA, SC, TN, TX. * Ochrotrichia dardeni Harris, 1986— AL, WV.

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+ Ochrotrichia denningi Blickle & Morse, 1957—ME, NH, PA, VA, WV. * Ochrotrichia eliaga (Ross, 1941), Polytrichia—AL, AR, KY, IL, IN, MO, OH, OK, TN. * Ochrotrichia elongiralla Harris, 1986— AL, TN. * Ochrotrichia graysoni Parker & Voshell, 1980—AL, PA, TN, VA, WV. * Ochrotrichia okaloosa Harris, 1987—FL. * Ochrotrichia provosti Blickle, 1961—FL. * Ochrotrichia riesi Ross, 1944—AL, AR, KY, IL, IN, MO, TN. * Ochrotrichia shawnee (Ross, 1938), Polytrichia—AL, IL, KY, NY, PA, TN. * Ochrotrichia spinosa (Ross, 1938), Polytrichia—AR, IL, KY, MB, MN, MO, NT, OH, OK, PA, TN, WI. * Ochrotrichia tarsalis (Hagen, 1861), Hydroptila—AL, AR, AZ, FL, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, NC, NE, NY, OH, OK, ON, PA, SC, TN, TX, VA, VT, WI, WV. * Ochrotrichia transylvanica Harris & Floyd, 1997—NC. * Ochrotrichia tuscaloosa Harris & Kelley, 1984—AL. + Ochrotrichia unio (Ross, 1941), Polytrichia—IL, MO, OH. * Ochrotrichia weoka Harris, 1989—AL. * Ochrotrichia wojcickyi Blickle, 1963—IN, KY, MB, ME, MN, NH, OH, PA, TN, VA. * Ochrotrichia xena (Ross, 1938), Polytrichia—AR, IL, IN, KY, MO, OH, TN. # * Orthotrichia aegerfasciella (Chambers, 1873), Clymene— AL, AR, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NY, OH, OK, ON, QC, SC, TN, TX, VA, WA, WI;

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Orthotrichia americana, Banks, 1904 (Flint, 1966); Orthotrichia brachiata Morton, 1905 (Ross, 1938); Oxyethira dorsalis Banks, 1904 (Ross, 1944). * Orthotrichia baldufi Kingsolver & Ross, 1961—AL, FL, ME, MN, NH, NY, QC, TX, WI. * Orthotrichia cristata Morton, 1905—AL, AR, BC, CO, DE, FL, GA, IL, IN, KS, KY, LA, MB, ME, MI, MN, MO, MS, MT, NC, NH, NY, OH, OK, ON, PA, QC, TN, TX, VA, WA, WI. * Orthotrichia curta Kingsolver & Ross, 1961—AL, AR, FL, KY, LA, MN, QC, TX. * Orthotrichia dentata Kingsolver & Ross, 1961—FL, MS, SC, VA. * Orthotrichia instabilis Denning, 1948— AL, AR, FL, NH, SC, TX. * Oxyethira (Oxytrichia) abacatia Denning, 1947—AL, FL, GA, MN, NC, NH, SC, TX. * Oxyethira (Dampfitrichia) aculea Ross, 1941—AL, AZ, NM, NV, OK, TX. * Oxyethira (Oxytrichia) anabola Blickle, 1966—AL, ME, MI, MN, NF, NH, NJ, ON, PA, QC, VA, WI. * Oxyethira (Dampfitrichia) arizona Ross, 1948—AZ, CA, FL, NV; Oxyethira cirrifera Flint, 1964 (Kelley & Morse, 1982). * Oxyethira (Loxotrichia) azteca (Mosely, 1937), Loxotrichia—NC, OK, SC, TX. * Oxyethira (Holarctotrichia) chrysocara Harris, 2002—FL. * Oxyethira (O.) coercens Morton, 1905— AL, AR, IL, IN, MB, ME, MN, MO, MT, NH, NY, OK, QC, TX, VA, WI.

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* Oxyethira (Oxytrichia) dualis Morton, 1905—AL, AR, AZ, CA, CO, IL, IN, KS, MO, MT, NH, NM, NY, OH, OK, OR, PA, SC, TX, UT, VA, VT, WA, WY; Oxyethira allosi Blickle, 1980 (Kelley & Morse, 1982). * Oxyethira (Holarctotrichia) dunbartonensis Kelley, 1981—GA, SC. * Oxyethira (Holarctotrichia?) elerobi (Blickle, 1961), Neotrichia—AL, FL, LA. * Oxyethira (Dampfitrichia) florida Denning, 1947—FL, TX. # * Oxyethira (Holarctotrichia) forcipata Mosely, 1934—AL, AR, CT, GA, KY, IL, ME, MI, MN, MO, NC, NH, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, WI. * Oxyethira (Argyrobothrus) glasa (Ross, 1941), Loxotrichia—AL, AR, FL, GA, LA, MS, NC, OK, SC, TN, TX, VA. * Oxyethira (O.) grisea Betten, 1934—AL, DE, FL, GA, IL, IN, KY, LA, ME, MI, MS, NC, NH, NJ, NS, NY, OH, PA, QC, SC, TN, VA. * Oxyethira (Loxotrichia) janella Denning, 1948—AL, AR, FL, GA, LA, MS, NC, OK, SC, TX, VA; Oxyethira neglecta Flint, 1964 (Blickle, 1979). * Oxyethira (Holarctotrichia) kelleyi Harris, 1987—FL. * Oxyethira (Dactylotrichia) kingi Holzenthal & Kelley, 1983—FL. * Oxyethira (O.) lumipollex Kelley & Harris, 1983—AL. * Oxyethira (O.) lumosa Ross, 1948—AL, FL, GA, SC, TX. * Oxyethira (Dampfitrichia) maya Denning, 1947—AL, FL, GA, HI, TX. * Oxyethira (Oxytrichia) mcgregori Harris & Huryn, 2000—AL, TN.

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* Oxyethira (Holarctotrichia) michiganensis Mosely, 1934—AL, BC, CT, GA, ME, MI, MN, NC, NH, NY, OH, PA, QC, SC, TN, VA, WI; Oxyethira sodalis Ross & Spencer, 1952 (Blickle, 1979). * Oxyethira (O.) novasota Ross, 1944—AL, AR, FL, GA, KY, LA, MS, NC, OH, SC, TN, TX. # * Oxyethira (Dampfitrichia) pallida (Banks, 1904), Orthotrichia—AL, AR, AZ, CA, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, MS, NB, NC, NE, NH, NY, OH, OK, PA, QC, SC, TN, TX, VA, WI, WV, WY; Oxyethira cibola Denning, 1948 (Ross, 1948); Oxyethira viminalis Morton, 1905 (Ross, 1938). * Oxyethira (Loxotrichia) parce (Edwards & Arnold, 1961), Protoptila—AL, TX. * Oxyethira (O.) pescadori Harris & Keth, 2002—AL, FL, KY, NC, TN, VA. * Oxyethira (O.) rivicola Blickle & Morse, 1954—AL, AR, GA, KY, ME, MN, NH, NY, PA, QC, SC, TN, VA, WI. # * Oxyethira (Argyrobothrus) roberti Roy & Harper, 1980—AL, FL, QC, SC, TX; Oxyethira leonensis Kelley, 1981 (Kelley, 1984). * Oxyethira (O.) rossi Blickle & Morse, 1957— KY, ME, MN, NH, NY, TN, WI; Oxyethira berneri Etnier, 1965 (Etnier, 1968). * Oxyethira (O.) savanniensis Kelley & Harris, 1983—AL, FL, SC. * Oxyethira (Holarctotrichia) serrata Ross, 1938—AB, BC, CT, ID, IL, IN, MB, ME, MI, MN, ND, NH, NY, QC, SK, TN, UT, WA, WI, WY. * Oxyethira (Holarctotrichia) setosa Denning, 1947—AL, FL, GA. * Oxyethira (Dampfitrichia) simulatrix Flint, 1968–FL.

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Hydroptilidae continued

* Oxyethira (Oxytrichia) sininsigne Kelley, 1981—AL, FL, LA, SC. * Oxyethira (Dampfitrichia) verna Ross, 1938—AL, FL, IL, LA, ME, MI, MN, NB, NC, NH, NJ, NS, ON, SC, TN, TX, WI. * Oxyethira (Argyrobothrus) zeronia Ross, 1941—AL, AR, FL, GA, IL, KS, KY, LA, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK , PA, QC, SC, TN, TX, VA, WA, WI; Oxyethira walteri Denning, 1947 (Blickle, 1979). # * Palaeagapetus celsus (Ross, 1938), Paragapetus—KY, ME, NB, NC, NH, NY, OK, PA, QC, TN, VA, VT, WV. * Stactobiella cahaba Harris, 1985—AL. * Stactobiella delira (Ross, 1938), Stactobia— AL, AR, BC, CA, CO, ID, IL, IN, KS, KY, ME, MN, MO, MT, NB, NC, NH, NY, OH, OK, OR, PA, SC, TN, VA, WA, WI, WV, WY, YT. * Stactobiella martynovi Blickle & Denning, 1977—AL, KY, NC, NY, OH, PA, SC, TN, VA; Stactobiella solzhenitsyni Sykora & Weaver, 1978 (Weaver & Sykora, 1990). # * Stactobiella palmata (Ross, 1938), Stactobia—AB, AL, AR, IL, KS, KY, MB, ME, MN, MO, NC, NH, NS, NY, OH, OK, OR, PA, SD, TN, VA, WI.

Lepidostomatidae

* Lepidostoma americanum (Banks, 1897), Olemira—CT, DE, GA, MA, MB, ME, MN, NC, NH, NY, PA, QC, SC, TN, VA, VT. * Lepidostoma bryanti (Banks, 1908), Alepomyia—CT, GA, MA, MB, ME, MI, MN, NB, NC, NF, NH, NS, NY, ON, PA, QC, RI, SC, TN, VA, WI;

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Lepidostoma wisconsinense Vorhies, 1909 (Ross, 1938). * Lepidostoma carolina (Banks, 1911), Notiopsyche—NC, SC. * Lepidostoma carrolli Flint, 1958—AR, CT, KY, MD, ME, NC, NJ, OH, PA, SC, TN, VA. * Lepidostoma compressum Etnier & Way, 1973—AL, TN. * Lepidostoma etnieri Weaver, 1988— KY, TN. * Lepidostoma excavatum Flint & Wiggins, 1961—NC, TN, VA. * Lepidostoma flinti Wallace & Sherberger, 1972—NC, SC. * Lepidostoma frosti (Milne, 1936), Atomyia—CT, MA, ME, NC, NH, NS, NY, PA, QC, RI, TN, VT. * Lepidostoma glenni Wallace & Sherberger, 1972—GA. * Lepidostoma griseum (Banks, 1911), Phanopsyche—AL, AR, CO, CT, DE, FL, GA, KY, MA, ME, MI, MN, NC, NH, NJ, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. * Lepidostoma latipenne (Banks, 1905), Notiopsyche—AL, DE, FL, GA, MA, NC, NH, NS, NY, PA, QC, SC, TN, VA, VT. + Lepidostoma liba Ross, 1941—AR, IL, MN, MO, OH, WI. * Lepidostoma lobatum Wallace & Sherberger, 1972—GA, NC, TN. * Lepidostoma lydia Ross, 1939—GA, KY, MA, NC, NF, NH, NS, NY, OH, PA, QC, SC, TN, VA, VT. * Lepidostoma mitchelli Flint & Wiggins, 1961—NC, TN, VA. * Lepidostoma modestum (Banks, 1905), Atomyia—NC, NH, QC, SC, TN, VA. * Lepidostoma morsei Weaver, 1988—FL, MS, TX. * Lepidostoma ontario Ross, 1941—AL, CT, MA, ME, NB, NC, NF, NH, NS, NY, ON, PA, QC, SC, TN, VA, VT.

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Lepidostomatidae continued

* Lepidostoma pictile (Banks, 1899), Mormonia—CT, KY, MA, ME, NB, NC, NF, NH, NJ, NS, NY, PA, QC, TN, VA, WV; Lepidostoma swannanoa Ross, 1939 (Weaver, 1988). * Lepidostoma sackeni (Banks, 1936), Mormomyia—CT, KY, MA, ME, MI, MN, ND, NF, NH, NS, NY, OH, ON, PA, QC, TN, VT, WI, WV. * Lepidostoma sommermanae Ross, 1946— CT, DE, IN, MA, NC, NH, NY, OH, ON, PA, QC, VA, VT. * Lepidostoma styliferum Flint & Wiggins, 1961—NC, SC, TN, WV. * Lepidostoma tibiale (Carpenter, 1933), Neuropsyche—AL, GA, NC, SC, TN, VA; Lepidostoma rileyi Denning, 1948 (Denning, 1954; Weaver, 1988). * Lepidostoma togatum (Hagen, 1861), Mormomyia—AB, AL, AR, CT, DC, DE, GA, KY, MA, MB, MD, ME, MI, MN, MO, NB, NC, ND, NF, NH, NJ, NS, NT, NY, OH, OK, ON, PA, QC, RI, SC, SK, TN, VA, VT, WI, WV; Silo pallidus Banks, 1897 (Milne, 1936; Ross, 1944); Pristosilo canadensis Banks, 1899 (Fischer, 1970). * Lepidostoma vernale (Banks, 1897), Mormonia—CT, MA, MD, ME, MI, NC, NF, NH, NS, NY, OH, ON, PA, QC, VA, VT, WI, WV. * Lepidostoma weaveri Harris, 1986—AL. * Lepidostoma wigginsi Weaver, 2002—CT, FL, LA, NC, PA, SC, TN, VA; Lepidostoma serratum Flint & Wiggins, 1961 (Weaver, 2002). * Theliopsyche corona Ross, 1938—NC, TN. * Theliopsyche epilonis Ross, 1938— NC, TN.

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* Theliopsyche grisea (Hagen, 1861), Silo— NC, NH, NJ, NY, PA, SC, TN, VA. # * Theliopsyche melas Edwards, 1956—AL, KY, TN, VA, WV. * Theliopsyche tallapoosa Harris, 1986— AL.

Leptoceridae

* Ceraclea (Ceraclea) alabamae Harris, 1989—AL, KY, PA, TN. # * Ceraclea (Ceraclea) alces (Ross, 1941), Athripsodes—AL, MA, ME, MN, NY, ON, QC, WI. # * Ceraclea (Athripsodina) ancylus (Vorhies, 1909), Leptocerus—AL, AR, CT, DE, GA, IL, IN, KS, KY, MA, MB, ME, MN, MO, NC, ND, NY, OH, OK, ON, PA, QC, SC, TN, VA, WI. * Ceraclea (Ceraclea) cama (Flint, 1965), Athripsodes—NC. # * Ceraclea (Ceraclea) cancellata (Betten, 1934), Leptocerus—AL, AR, BC, CT, DE, FL, GA, IL, IN, KS, KY, LA, MA, MB, ME, MI, MN, MO, MS, MT, NC, ND, NF, NH, NS, NY, OH, OK, ON, PA, QC, SC, TN, VA, WI, WV, YT; Athripsodes improcerus Edwards, 1956 (Morse, 1975). # * Ceraclea (Athripsodina) diluta (Hagen, 1861), Leptocerus—AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, LA, MA, MB, ME, MI, MN, MO, MS, MT, NB, NC, ND, NF, NH, NY, OH, OK, ON, PA, QC, SC, TN, VA, VT, WI, WV; Leptocerus retactus Banks, 1914 (Morse, 1975). # * Ceraclea (Ceraclea) enodis Whitlock & Morse, 1994—CT, GA, IL, KY, NC, NY, ON, SC, TN, VA. # * Ceraclea (Athripsodina) flava (Banks, 1904), Leptocerus—AL, AR, DC, DE, FL, IA, IL, IN, KS, KY, MD, MI, MN, MO, MS, NC, NH, NY, OH, OK, ON, PA, SC, TN, TX, VA, WI, WV.

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

* Ceraclea (Ceraclea) floridana (Banks, 1903), Leptocerus—FL. # * Ceraclea (Athripsodina) joannae Morse & Lenat, 2006—NC, TN. * Ceraclea (Ceraclea) limnetes Rasmussen & Harris, 2008, in Rasmussen, Denson, & Harris, 2008—FL. # * Ceraclea (Ceraclea) maculata (Banks, 1899), Leptocerus—AL, AR, BC, CA, CO, CT, DC, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NH, NJ, NY, OH, OK, ON, OR, PA, QC, SC, TN, TX, VA, VT, WI, WV; Leptocerus inornatus Banks, 1914 (Flint, 1966). # * Ceraclea (Ceraclea) mentiea (Walker, 1852), Leptocerus—AL, DC, IL, IN, MI, MN, NC, NH, NJ, NY, ON, PA, QC, VA, WI; Leptocerus vanus Betten, 1934 (Betten & Mosely, 1940). # * Ceraclea (Athripsodina) neffi (Resh, 1974), Athripsodes—AL, KS, KY, MI, MN, NC, OH, TN, VA, WV. # * Ceraclea (Athripsodina) nepha (Ross, 1944), Athripsodes—AL, AR, DE, FL, GA, IL, KS, KY, MI, MN, MO, MS, NC, OK, SC, TN, TX, VA, WI. * Ceraclea (Ceraclea) ophioderus (Ross, 1938), Athripsodes—AL, AR, FL, IL, IN, KY, LA, MO, MS, NC, TN, TX, VA, WV. * Ceraclea (Athripsodina) protonepha Morse & Ross, in Morse, 1975—AL, DE, FL, GA, KS, KY, LA, MS, NC, OH, SC, TN, TX, VA. # * Ceraclea (Ceraclea) punctata (Banks, 1894), Mystacides—AR, DC, IL, KS, KY, MD, ME, MI, MO, NC, NH, NY, OH, OK, PA, QC, TN, VA, WI.

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# * Ceraclea (Ceraclea) resurgens (Walker, 1852), Leptocerus—AL, BC, CO, CT, DC, FL, IL, IN, KY, LA, MB, ME, MI, MN, MT, NC, ND, NH, NJ, NY, OH, OK, ON, OR, PA, QC, SC, SK, TN, VA, WI, WY, YT; Leptocerus aspinosus Betten, 1934 (Milne, 1936); Leptocerus variegatus Hagen, 1861 (Ross, 1938). # * Ceraclea (Ceraclea) slossonae (Banks, 1938), Athripsodes—CT, FL, GA, IA, MD, MS, NC, NY, PA, QC, VA, WV; Athripsodes daggyi Denning, 1948 (Flint, 1966). # * Ceraclea (Ceraclea) spongillovorax (Resh, 1974), Athripsodes—AL, FL, IL, IN, KS, KY, LA, MS, VA. # * Ceraclea (Ceraclea) submacula (Walker, 1852), Leptocerus—NY, OH, ON, QC, TN, VT. # * Ceraclea (Athripsodina) tarsipunctata (Vorhies, 1909), Leptocerus—AL, AR, CA, CO, CT, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MB, ME, MI, MN, MO, MS, MT, NB, NC, ND, NH, NV, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, VA, VT, WA, WI, WV, WY. # * Ceraclea (Ceraclea) transversa (Hagen, 1861), Leptocerus—AL, AR, CT, DC, DE, FL, GA, ID, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NF, NH, NV, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV; Leptocerus angustus Banks, 1914 (Morse, 1975). # * Leptocerus americanus (Banks, 1899), Setodes—AL, AR, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, ME, MI, MN, MS, NC, ND, NH, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV.

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# * Mystacides sepulchralis (Walker, 1852), Leptocerus—AB, AL, AK, AR, BC, CA, CT, DE, GA, IL, IN, KY, MA, MB, ME, MI, MN, MO, MT, NB, NC, ND, NF, NH, NJ, NS, NT, NY, OH, OK, ON, PA, QC, SC, SD, SK, TN, VA, VT, WI, WV, WY, YT; Setodes longula Navas, 1934 (Schmid, 1950).

# * Nectopsyche albida (Walker, 1852), Leptocerus—AK, BC, DC, GA, IA, IL, IN, KS, KY, MA, MB, ME, MI, MN, MO, MS, MT, NC, ND, NH, NJ, NY, OH, ON, PA, QC, SC, SK, TX, UT, VA, VT, WI; Leptocella coloradensis Banks, 1899 (Flint, 1966); Leptocella gracilis Banks, 1904 (preoccupied by Banks, 1901); Setodes nivea Hagen, 1861 (Milne, 1934); Mystacides uwarowii Kolenati, 1859 (Milne, 1934). # * Nectopsyche candida (Hagen, 1861), Setodes—AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, OH, OK, ON, PA, QC, SC, TN, TX, VA, WI, WV. # + Nectopsyche diarina (Ross, 1944), Leptocella—CO, DE, ID, IL, IN, KS, MB, MI, MN, MO, MT, ND, NE, NY, OH, OK, ON, SD, SK, TX, UT, VT, WI, WY. # * Nectopsyche exquisita (Walker, 1852), Leptocerus—AL, AR, CT, DC, DE, FL, GA, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, ND, NE, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV; Setodes piffardii McLachlan, 1863 (Haddock, 1977). # * Nectopsyche paludicola Harris, 1986— AL, FL.

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# * Nectopsyche pavida (Hagen, 1861), Setodes—AL, AR, CT, DC, FL, GA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NY, OH, OK, ON, QC, SC, TN, TX, VA, WI, WV. * Nectopsyche spiloma (Ross, 1944), Leptocella—AL, AR, IL, KS, LA, MO, MS, OK, SD, TX. # * Nectopsyche tavara (Ross, 1944), Leptocella—FL. # * Nectopsyche waccamawensis Glover & Floyd, 2004—NC. # * Oecetis (Pseudosetodes) avara (Banks, 1895), Setodes—AB, AL, AR, AZ, BC, CA, CO, CT, DC, DE, FL, GA, IA, ID, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, MT, NC, ND, NH, NJ, NM, NS, NY, OH, OK, ON, OR, PA, QC, SC, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY. # * Oecetis (Pleurograpta) cinerascens (Hagen, 1861), Setodes—AB, AL, AR, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NB, NC, ND, NE, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, SD, SK, TN, TX, VA, VT, WA, WI, WV; Oecetina floridana Banks, 1899 (Milne, 1934 errata); Setodes floridana Banks, 1905 (Holzenthal, 1982); Oecetina fumosa Banks, 1899 (Ross, 1938). * Oecetis (Pseudosetodes) daytona Ross, 1947—AL, FL, MS, SC. * Oecetis (O.) ditissa Ross, 1966—AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, MN, MO, MS, NC, OH, OK, PA, SC, TN, TX, VA. # * Oecetis (Pleurograpta) georgia Ross, 1941—AL, FL, GA, LA, NC, SC, TN, TX.

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# * Oecetis (O.) inconspicua (Walker, 1852), Leptocerus –AB, AK, AL, AR, AZ, BC, CA, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NF, NH, NJ, NM, NS, NT, NV, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY, YT; Oecetina antillana Banks, 1938 (Flint, 1967); Oecetina apicalis Banks, 1907 (Milne, 1934 errata; Ross, 1944); Setodes flaveolata Hagen, 1861 (Milne, 1934 errata; Ross, 1938); Oecetina flavida Banks, 1899 (Milne, 1934 errata; Ross, 1938); Oecetina inornata Banks, 1907 (Milne, 1934 errata; Ross, 1944); Setodes micans Hagen, 1861 (Milne, 1934 errata; Ross, 1938); Oecetina parvula Banks, 1899 (Milne, 1934 errata; Ross, 1938); Setodes sagitta Hagen, 1861 (Milne, 1934 errata; Ross, 1938). # * Oecetis (Quaria) morsei Bueno-Soria, 1981—AL, FL, SC. # * Oecetis (O.) nocturna Ross, 1966—AL, AR, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MB, MD, MN, MO, MS, NC, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WV. # * Oecetis (O.) ochracea (Curtis, 1825), Leptocerus—AB, AK, BC, CA, CO, MB, MN, MT, NC, ND, NT, OH, ON, SD, SK, TN, VA, WI, WY, YT; Oecetis ochracea carri Milne, 1934 (Ross, 1944); Phryganea hectica Zetterstedt, 1840 (Fischer, 1966); Mystacida obsoleta Rambur, 1842 (Fischer, 1966); Phryganea pilosa Müller, 1776 (Wallengren, 1870; Fischer, 1966);

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Oecetis vicaria Neave, 1934 (Fischer, 1966). # * Oecetis (Pleurograpta) osteni Milne, 1934—AL, AR, CT, FL, IL, IN, LA, MA, ME, MI, MN, MS, NB, NC, NF, NH, NJ, NY, OH, OK, ON, PA, QC, RI, SC, TX, VA, WI. # * Oecetis (Pseudosetodes) parva (Banks, 1907), Setodina—AL, FL, GA, SC. # * Oecetis (Pleurograpta) persimilis (Banks, 1907), Oecetina—AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NF, NH, NJ, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV. # * Oecetis (O.) porteri Ross, 1947—AL, FL, NC, NS, SC. * Oecetis (O.) pratelia Denning, 1948—FL. * Oecetis (Quaria) scala Milne, 1934—AL, AR, KY, MD, NC, NH, NJ, NY, PA, QC, SC. # * Oecetis (Quaria) sphyra Ross, 1941—AL, FL, GA, KY, LA, MS, NC, SC, TN, TX, VA. # * Setodes arenatus Holzenthal, 1982— NC, SC. * Setodes chipolanus Rasmussen & Harris, 2008, in Rasmussen, Denson, & Harris, 2008—FL. # * Setodes dixiensis Holzenthal, 1982—AL, LA, MS. # * Setodes epicampes Edwards, 1956—AL, KY, TN. # * Setodes guttatus (Banks, 1900), Oecetina— AL, CT, DC, FL, MD, ME, MN, NJ, PA, QC, VA. # * Setodes incertus (Walker, 1852), Leptocerus—AL, CT, GA, KY, MA, MD, ME, MI, MN, NC, NH, NJ, NY, ON, PA, QC, SC, TN, VA, WI, WV; Setodes autumnalis Banks, 1907 (Holzenthal, 1982); Setodes vernalis Banks, 1907 (Milne, 1934).

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# * Setodes oligius (Ross, 1938), Leptocerus— IL, IN, ME, MI, MN, NY, ON, PA, QC, TN, WI. # + Setodes oxapius (Ross, 1938), Leptocerus— AR, MO, OK, PA. # * Setodes stehri (Ross, 1941), Leptocerus— AL, GA, NC, SC, TN.

# * Triaenodes (T.) aba Milne, 1935—AL, AR, DE, FL, GA, IL, IN, KY, LA, MA, MB, ME, MI, MN, MS, NC, NH, NY, OH, ON, PA, QC, SC, TN, VA, WI. # * Triaenodes (T.) bicornus Manuel, 2010— AL, FL, GA, SC. # * Triaenodes (T.) cumberlandensis Etnier & Way, 1973—AL, AR, GA, KY, OK, TN. # * Triaenodes (T.) dendyi Manuel, 2010— AL, FL, GA, SC. * Triaenodes (T.) dipsia Ross, 1938—AL, AR, KS, KY, MB, MI, MN, ND, NY, OH, OK, ON, PA, QC, TN, VA, WI. # * Triaenodes (T.) flavescens Banks, 1900— AL, BC, CT, FL, GA, IL, IN, KY, MD, ME, MI, MN, MO, MS, NC, NJ, NY, OH, OK, ON, PA, QC, TN, TX, VA, WI, WV. Triaenodes venustus Smith, 1900 (Fischer, 1965). # * Triaenodes (new subgenus A) florida Ross, 1941—AL, FL, GA. # * Triaenodes (T.) furcella Ross, 1959—FL. # * Triaenodes (T.) helo Milne, 1934—AL, FL, GA, NC, OK, SC. # * Triaenodes (T.) ignita (Walker, 1852), Leptocerus—AL, AR, CT, DE, FL, GA, IL, IN, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK, PA, QC, SC, TN, TX, VA, WI, WV; Triaenodes dentatus Banks, 1914 (Ross, 1944).

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# * Triaenodes (T.) injusta (Hagen, 1861), Setodes—AB, AL, AR, BC, CT, DC, DE, FL, GA, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, NC, NF, NH, NJ, NS, NY, OH, OK, ON, OR, PA, QC, SC, SK, TN, TX, VA, VT, WI, WV. * Triaenodes (T.) lagarto Rasmussen & Harris, 2010, in Manuel, 2010— FL. # * Triaenodes (T.) marginata Sibley, 1926— AB, AL, AR, BC, CT, DC, DE, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, NC, ND, NF, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, SD, SK, TN, VA, WI, WV; Triaenodes inflexa Morse, 1971 (Manuel, 2010). # * Triaenodes (T.) melaca Ross, 1947—AB, AL, IL, IN, KS, KY, MN, MO, MS, NC, OH, ON, TN. # * Triaenodes (T.) milnei Manuel, 2010—AL, FL, GA, SC. # * Triaenodes (T.) morsei Manuel, 2010—SC. # * Triaenodes (T.) nox Ross, 1941—AB, AL, BC, CT, FL, GA, IL, IN, KY, MA, MB, ME, MI, MN, MO, MS, NB, NH, NJ, NS, NY, OH, ON, PA, QC, SC, SK, TN, VA, WI. # * Triaenodes (T.) ochracea (Betten & Mosely, 1940), Triaenodella—AL, AR, CT, DE, FL, GA, KY, MA, MD, ME, MS, NC, NJ, OH, SC, TN, TX, VA. # * Triaenodes (T.) perna Ross, 1938—AL, AR, DE, FL, GA, IL, KS, KY, LA, MA, MD, MO, MS, NC, NH, NJ, NY, OH, ON, OK, PA, QC, SC, TN, TX, VA, WV. * Triaenodes (T.) rossi Manuel, 2010—SC. * Triaenodes (T.) smithi Ross, 1959—AL, AR, FL, GA, IL, LA, MS, SC, TX, VA. # * Triaenodes (T.) taenia Ross, 1938—AL, FL, GA, NC, SC, TN, VA.

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Leptoceridae continued

# * Triaenodes (T.) tarda Milne, 1934—AB, AK, AL, AR, AZ, BC, CA, CO, CT, DC, DE, FL, GA, IA, ID, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NH, NJ, NS, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, UT, VA, VT, WA, WI, WV, WY, YT; Triaenodes mephitus Milne, 1936 (Ross, 1944); Triaenodes vorhiesi Betten, 1934 (Ross, 1944). * Triaenodes (T.) tridonta Ross, 1938—AL, FL, OK.

Limnephilidae

+ Anabolia apora Parker, 1984—VA. # * Anabolia consocia (Walker, 1852), Limnephilus (Goniotaulius)—AB, AK, CO, CT, IA, IL, IN, MA, MB, ME, MI, MN, NC, ND, NE, NH, NT, NY, OH, ON, PA, QC, SD, SK, TN, VA, VT, WI, WV, WY; Colpotaulius medialis Banks, 1905 (Milne, 1935); Anabolia oslari Ling, 1938 (Ross, 1944). # * Frenesia difficilis (Walker, 1852), Limnephilus—CT, DC, DE, KY, MA, ME, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, TN, VA, VT, WV; Cryptothrix coagulatus Provancher, 1877 (Milne, 1935); Chilostigma pallida Banks, 1899 (Milne, 1935). # * Frenesia missa Milne, 1935—AR, CT, DC, DE, IA, IL, IN, MA, MD, ME, MI, MN, MO, NC, ND, NH, NY, OH, ON, PA, PE, QC, VA, VT, WI. # * Glyphopsyche sequatchie Etnier & Hix, 1999—TN.

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# * Hydatophylax argus (Harris, 1869), Phryganea—AL, CT, DE, FL, IN, MA, MB, MD, ME, MI, MN, NB, NC, NF, NH, NJ, NY, OH, ON, PA, PE, QC, SC, TN, VA, WA, WI, WV.

* Ironoquia kaskaskia (Ross, 1944), Caborius—AL, AR, DE, IN, IL, KY, LA, MD, MS, OH, TN, VA, WV; Ironoquia brysoni Flint, 1972 (Etnier & Way, 1973). * Ironoquia lyrata (Ross, 1938), Caborius— CT, IL, IN, KY, ME, MN, NY, OH, PA, QC, VA, WI. # + Ironoquia parvula (Banks, 1900), Chaetopterygopsis—CT, DE, NB, NH, NJ, NY, OH, PA, VA. # * Ironoquia punctatissima (Walker, 1852), Halesus—AK, AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV. # * Limnephilus indivisus Walker, 1852—AB, BC, CO, CT, IL, IN, KY, MA, MB, ME, MI, MN, MT, ND, NF, NH, NS, NT, NY, OH, ON, PA, QC, SK, TN, UT, VA, VT, WI, WV, WY, YT; Limnephilus selatus Denning, 1966 (Ruiter, 1995); Limnephilus subguttatus Walker, 1852 (Betten & Mosely, 1940). * Limnephilus moestus Banks, 1908—AB, AZ, BC, CO, CT, DE, IL, MA, MB, ME, MI, MN, MT, NB, NC, ND, NF, NH, NM, NS, NY, OH, ON, PA, QC, UT, VA, VT, WA, WI, WV, WY; Limnephilus gioia Denning, 1948 (Ruiter, 1995); Limnephilus hingstoni Mosely, 1929 (Forsslund, 1932); Limnephilus valhalla Nimmo, 1971 (Ruiter, 1995).

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+ Limnephilus ornatus Banks, 1897—AB, AK, CT, DE, IL, IN, MA, MB, ME, MI, MN, MT, ND, NF, NH, NS, NT, NY, OH, ON, PA, PE, QC, SK, VT, WA, WI, WV. # + Limnephilus rhombicus Linnaeus, 1758— AB, AK, BC, CO, IA, IL, IN, MA, MB, ME, MI, MN, ND, NF, NJ, NS, NY, OH, ON, PA, PE, QC, SK, VT, WI, WV, WY, YT; Limnephilus chilcotinensis Nimmo, 1991 (Ruiter, 1995); Limnephilus combinatus Walker, 1852 (McLachlan, 1875); Phryganea rhomboidica Berkenhout, 1795 (Fischer, 1968). # + Limnephilus sericeus (Say, 1824), Phryganea—AB, AK, BC, CO, CT, ID, MA, MB, ME, MI, MN, NF, NH, NS, NT, NY, OH, ON, OR, PA, QC, VT, WA, WI, WV, WY, YT; Anabolia decepta Banks, 1899 (Flint, 1966); Limnephilus despectus Walker, 1852 (Schmid, 1955); Limnephilus eminens Betten, 1934 (Milne, 1935; Betten & Mosely, 1940; Ross, 1944); Apatania fuscostigma Matsumura, 1931 (Nozaki & Tanida, 1996); Limnephilus multifarius Walker, 1852 (Milne, 1935; Betten & Mosely, 1940; Ross, 1944); Limnephilus perforatus Walker, 1852 (Betten & Mosely, 1940; Ross, 1944) Phryganea pilosula Zetterstedt, 1840 (Fischer, 1968); Limnophilus shimushirensis Tsuda, 1942 (Schmid, 1955). # * Limnephilus submonilifer Walker, 1852— AR, CO, CT, DC, DE, IA, IL, IN, KY, MA, MB, MD, ME, MI, MN, NC, ND, NF, NH, NJ, NS, NY, OH, ON, PA, QC, RI, SC, SD, TN, VA, VT, WA, WI, WV;

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Limnophilus (Goniotaulius) pudicus Hagen, 1861 (Milne, 1935; Ross, 1938). # + Nemotaulius (Macrotaulius) hostilis (Hagen, 1873), Glyphotaelius— AB, AK, BC, CO, CT, MA, MB, ME, MI, MN, ND, NF, NH, NS, NT, NY, OH, ON, OR, PA, QC, SK, UT, VA, VT, WI, WV, WY, YT. + Platycentropus amicus (Hagen, 1861), Hallesus—AB, LA, MB, MI, MN, NJ, ON, QC, RI, SK, WI; Platycentropus plectrus Ross, 1938 (Flint, 1966). # * Platycentropus radiatus (Say, 1824), Phryganea—AL, AR, CT, DE, GA, IA, IL, IN, KY, MA, MB, MD, ME, MI, MN, MS, NC, ND, NF, NH, NJ, NS, NT, NY, OH, ON, PA, QC, SC, TN, VA, WA, WI, WV; Halesus hostis Hagen, 1861 (Smith, 1910; Ross, 1944); Limnephilus indicans Walker, 1852 (Ross, 1944); Hallesus maculipennis Kolenati, 1859 (Ross, 1944). # * Pseudostenophylax sparsus (Banks, 1908), Halesus—CT, MA, ME, MN, NC, NF, NH, NY, OH, ON, PA, QC, VA, VT, WI, WV; Stenophylax calypso Banks, 1911 (Ross, 1938). # * Pseudostenophylax uniformis (Betten, 1934), Drusinus—AL, CT, IL, IN, KY, MA, ME, MN, MO, NC, NH, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. # * Pycnopsyche antica (Walker, 1852), Neuronia—AL, FL, GA, IL, IN, KY, LA, MA, MI, MS, NC, NH, NJ, NY, ON, PA, SC, TN, TX, VA, WI, WV;

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Pycnopsyche minima Banks, 1943 (Wojtowicz, 1982). * Pycnopsyche circularis (Provancher, 1877), Platyphylax—CT, KY, MA, MD, ME, NC, NH, NS, NY, OH, PA, QC, RI, TN, VA, WI, WV. # * Pycnopsyche conspersa Banks, 1943—NC, NH, NY, ON, PA, QC, TN, VA. # * Pycnopsyche divergens (Walker, 1852), Limnephilus—CO, CT, MA, ME, NC, NH, NS, NY, OH, PA, TN, VA, VT, WV; Halesus dan Sibley, 1926 (Betten & Mosely, 1940). # * Pycnopsyche flavata (Banks, 1914), Stenophylax—GA, KY, NC, SC, TN, VA. # * Pycnopsyche gentilis (McLachlan, 1871), Stenophylax—AL, CT, DE, GA, KY, MA, MD, ME, MS, NC, NF, NH, NJ, NS, NY, OH, PA, PE, QC, SC, TN, VA, VT, WV. # * Pycnopsyche guttifera (Walker, 1852), Halesus—AB, AL, AR, CO, CT, DE, GA, IL, IN, KS, KY, LA, LB, MA, MB, ME, MI, MN, MO, MT, NC, ND, NE, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, SD, SK, TN, VA, VT, WA, WI, WV, WY; Pycnopsyche similis Banks, 1907 (Ross, 1938). # * Pycnopsyche indiana (Ross, 1938), Stenophylax—AL, AR, DE, FL, GA, IL, IN, KY, LA, MO, NC, NY, OH, SC, TN, TX, VA, WV. # * Pycnopsyche lepida (Hagen, 1861), Enoicyla—AL, AR, CT, DE, GA, IA, IL, IN, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, ND, NF, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, SD, TN, TX, VA, VT, WI, WV. * Pycnopsyche limbata (McLachlan, 1871), Stenophylax –MA, ME, MN, NF, NH, NS, NY, ON, QC, RI, TN, VT, WI.

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# * Pycnopsyche luculenta (Betten, 1934), Stenophylax—AL, CT, DE, GA, IN, KY, MA, ME, MS, NC, NH, NJ, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. # * Pycnopsyche pani Wojtowicz & Flint, 2007—NC, VA. # * Pycnopsyche rossi Betten, 1950– AR, IL, IN, KY, MO, OH, TN. # * Pycnopsyche scabripennis (Rambur, 1842), Limnephila—AK, AL, AR, CT, DE, GA, IL, IN, KY, LA, MA, MD, ME, MI, MN, MS, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, VA, VT, WI, WV; Pycnopsyche minima Banks, 1943 (Betten, 1950); Pycnopsyche perplexa Betten & Mosely, 1940 (Betten, 1950). # * Pycnopsyche sonso (Milne, 1935), Stenophylax—GA, NC, SC, TN, VA. # * Pycnopsyche subfasciata (Say, 1828), Phryganea -- AB, AR, CO, DE, GA, IA, IL, IN, KS, KY, MA, MB, MD, MI, MN, MO, NC, NH, NJ, NT, NY, OH, OK, ON, PA, QC, SC, SD, TN, VA, VT, WI, WV, WY. # * Pycnopsyche virginica (Banks, 1900), Potamorites—AL, KY, NC, SC, TN, VA.

Molannidae

# * Molanna (Molanneria) blenda Sibley, 1926—AL, AR, CT, DE, FL, GA, IL, IN, KY, LA, MA, MI, MN, MO, MS, NC, NF, NH, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. # * Molanna (Molanna) tryphena Betten, 1934—AL, CT, DE, FL, GA, KY, LA, MA, ME, MI, MN, MS, NC, NH, NY, OH, PA, QC, SC, TN, TX, VA, VT, WI.

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Molannidae continued

# * Molanna (Molanna) ulmerina Navás, 1934—AL, AR, CT, DE, FL, IL, IN, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV; Molanna musetta Betten, 1934 (Schuster, 1979). # * Molanna (Molanna) uniophila Vorhies, 1909—AR, IL, IN, MA, MB, ME, MI, MN, NB, NC, NF, NH, NY, OH, OK, ON, PA, QC, VA, VT, WI.

Odontoceridae

# + Marilia flexuosa Ulmer, 1905—AR, AZ, CA, IN, MO, NM, NV, OK, ON, TX, VT; Anisocentropus fuscus Banks, 1905 (Betten, 1934; Milne, 1936). # * Pseudogoera singularis Carpenter, 1933— GA, NC, SC, TN. # * Psilotreta amera Ross, 1939—GA, NC, SC, TN. # * Psilotreta frontalis Banks, 1899—AL, CT, DE, FL, GA, KY, MA, ME, MS, NB, NC, NF, NH, NJ, NS, NY, PA, QC, SC, TN, VA, VT; Heteroplectron gameta Ross, 1939 (Ross, 1944); Psilotreta hansoni Denning, 1948 (probably a synonym, Parker & Wiggins, 1987). # * Psilotreta indecisa (Walker, 1852), Goera—ME, MN, NB, NC, NH, NJ, NS, NY, OH, ON, PA, QC, WI; Heteroplectron borealis Provancher, 1877 (Schmid, 1983; Parker & Wiggins, 1987). # * Psilotreta labida Ross, 1944—AL, CT, GA, KY, MA, MD, ME, NB, NC, NH, NJ, NY, ON, PA, TN, VA, VT, WV. # * Psilotreta rossi Wallace, 1971—NC, TN, VA, WV.

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Odontoceridae continued

# * Psilotreta rufa (Hagen, 1861), Molanna— AL, CT, DE, KY, MA, MS, NC, NY, OH, PA, TN, VA, VT; Astoplectron connexa Banks, 1914 (Ross, 1944); “Heteroplectron? dissimilis” Banks, 1897 (Parker & Wiggins, 1987).

Philopotamidae

# * Chimarra aterrima Hagen, 1861—AL, AR, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NF, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV. # * Chimarra augusta Morse, 1971—AL, NC, SC, TN, VA, WV. * Chimarra falculata Lago & Harris, 1987— AL, FL, GA, MS. # * Chimarra feria Ross, 1941—AL, AR, IA, IL, IN, KS, KY, MI, MN, MO, MS, NE, NF, OH, OK, ON, QC, TN, TX, WI. # * Chimarra florida Ross, 1944—AL, FL, GA, LA, MS, NC, NJ, SC, VA. # + Chimarra holzenthali Lago & Harris, 1987—LA, TX. * Chimarra moselyi Denning, 1948—AL, FL, GA, IL, IN, KY, LA, MO, MS, NC, SC, TN, TX, VA; Chimarra perigua Ross, 1948 (Denning, 1950). # * Chimarra obscura (Walker, 1852), ? Beraea—AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, LB, MA, MB, MD, ME, MI, MN, MO, MS, NC, NE, NF, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, TX, VA, VT, WI, WV; Chimarra lucia Betten, 1934 (syn. of C. plutonis according to Milne, 1936; Ross, 1938); Wormaldia plutonis Banks, 1911 (Betten & Mosely, 1940).

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Philopotamidae continued

* Chimarra parasocia Lago & Harris, 1987—AL, AR, FL, KY, LA, MO, MS, TN, TX. # * Chimarra socia Hagen, 1861—AL, AR, CT, DC, FL, GA, IN, KY, LA, LB, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV; Wormaldia femoralis Banks, 1911 (Milne, 1936; Ross, 1938). # * Dolophilodes distincta (Walker, 1852), Philopotamus—AL, CT, DC, DE, GA, IN, KY, LB, MA, MD, ME, MI, MN, MS, NB, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV; Philopotamus americanus Banks, 1895 (Milne, 1936). # * Fumonta major (Banks, 1914), Dolophilus—AL, GA, KY, NC, SC, TN, VA.

* Sisko sisko (Ross, 1949), Wormaldia—CA, NC, OR, SC, TN, VA. # * Wormaldia moesta (Banks, 1914), Paragapetus—AL, AR, CT, DE, FL, GA, IL, IN, KY, MA, MB, MD, ME, MN, MO, MS, NB, NC, NF, NH, NJ, NS, NT, NY, OH, OK, ON, PA, QC, SC, TN, VA, VT, WI, WV; Dolophilus breviatus Banks, 1914 (Milne, 1936). * Wormaldia mohri (Ross, 1948), Gatlinia— NC, SC, TN, VA. * Wormaldia oconee Morse, 1989—NC, SC. # * Wormaldia shawnee (Ross, 1938), Dolophilus—AL, CT, IL, KY, MO, NC, NH, OH, PA, SC, TN, VA, WV. * Wormaldia thyria Denning, 1950—AL, KY, NC, SC, TN, VA.

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Phryganeidae

# * Agrypnia improba (Hagen, 1873), Phryganea—AB, AK, AL?, BC, ID, MA, MB, ME, MI, MN, MT, NC?, ND, NF, NH, NS, NT, NY, ON, OR, QC, SK, TN?, UT, VT, WA, WI, WY, YT [doubtfully southeastern—Etnier et al., 1998]; Dasystegia improba sackeni Banks, 1943 (Wiggins, 1998). # * Agrypnia vestita (Walker, 1852), Neuronia—AB, AL, AR, BC, CT, DC, DE, FL, GA, IL, IN, KS, KY, LB, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NF, NH, NJ, NS, NT, NY, OH, OK, ON, OR, PA, QC, SC, TN, TX, VA, VT, WA, WI, WV; Neuronia commixta Walker, 1852 (Betten & Mosely, 1940). * Banksiola concatenata (Walker, 1852), Neuronia—AL, CT, FL, GA, IL, MA, MB, ME, MI, NH, NJ, NS, NY, QC, SC; Banksiola cornuta Banks, 1951 (Wiggins, 1956); Neuronia irrorata Hagen, 1861 (Fischer, 1964). * Banksiola dossuaria (Say, 1828), Phryganea –CT, KY, MA, ME, MN, NB, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, VA, VT, WI, WV. # * Oligostomis ocelligera (Walker, 1852), Neuronia—CT, DE, IN, KY, MA, ME, MI, NF, NH, NJ, NS, NY, ON, PA, QC, TN, WI; Neuronia stygipes Hagen, 1873 (Milne, 1934). # * Oligostomis pardalis (Walker, 1852), Neuronia—CT, LB, MA, ME, NC, NH, NS, NY, OH, ON, PA, QC, TN, VA, WV; Neuronia pardalis redmani Betten & Mosely, 1940 (Ross, 1944).

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Chapter 4 ~ Key to Caddisflies

Phryganeidae continued

* Phryganea cinerea Walker, 1852—AB, AK, BC, CA, CO, CT, IA, ID, IL, IN, KY, MA, MB, ME, MI, MN, MT, NB, ND, NE, NF, NH, NS, NT, NY, OH, ON, OR, PA, PE, QC, SD, SK, UT, WA, WI, WY, YT; Phryganea divulsa Walker, 1860 (Fischer, 1964. # * Phryganea sayi Milne, 1931—AL, AR, CT, DC, DE, IA, IL, IN, KS, KY, MA, MD, ME, MI, MN, MO, NC, ND, NJ, NY, OH, ON, PA, QC, SC, SK, TN, TX, VA, WI, WV; Phryganea interrupta Say, 1828 (preoccupied by Fabricius). * Ptilostomis ocellifera (Walker, 1852), Neuronia—AK, AL, AR, BC, CA, CT, DC, DE, FL, GA, IA, ID, IL, IN, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, NB, NC, ND, NF, NH, NJ, NS, NT, NY, OH, OK, ON, OR, PA, QC, SC, SD, TN, TX, VA, VT, WA, WI, WV; Ptilostomis kovalevskii variety b Kolenati, 1859 (Wiggins, 1998); Neuronia simulans Betten & Mosely, 1940 (Ross, 1944). * Ptilostomis postica (Walker, 1852), Neuronia—AL, AR, CT, DC, DE, FL, GA, IL, IN, KY, LA, MA, MB, MD, ME, MI, MO, MS, NC, NH, NJ, NS, NY, OH, OK, ON, PA, QC, RI, SC, TN, TX, VA, WI, WV; Ptilostomis kovalevskii variety a Kolenati, 1859 (Fischer, 1964). * Ptilostomis semifasciata (Say, 1828), Phryganea—BC, CO, CT, DC, GA, IA, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MT, ND, NE, NF, NH, NJ, NS, NT, NY, OH, ON, PA, QC, SD, SK, TX, VA, WI, WV, WY, YT; Neuronia dubitans Betten & Mosely, 1940 (Ross, 1944); Neuronia fusca Walker, 1852 (Hagen, 1861; Betten & Mosely, 1940).

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Polycentropodidae

* Cernotina calcea Ross, 1938—AL, AR, FL, IL, KS, KY, LA, MO, MS, NC, OK, TN, TX, VA. + Cernotina pallida (Banks, 1904), Cyrnus— DC, MD, NH, OH, ON, VA. * Cernotina spicata Ross, 1938 –AL, AR, CT, DE, FL, GA, IL, IN, KS, KY, LA, MA, ME, MI, MS, NC, NH, OK, ON, SC, TN, TX, VA. * Cernotina truncona Ross, 1947—AL, FL, NC, VA. # * Cyrnellus fraternus (Banks, 1905), Cyrnus—AL, AR, CO, DE, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, MS, NC, NE, NY, OH, OK, PA, SC, TN, TX, VA, WI, WV; Nyctiophylax marginalis Banks, 1930 (Flint, 1964); Cyrnellus minimus Banks, 1913 (Flint, 1971); Cyrnellus zernyi Mosely, 1934 (Ross, 1938). # * Holocentropus flavus Banks, 1908—AB, AK, BC, CA, IA, IL, IN, MA, MB, ME, MI, MN, MT, ND, NE, NF, NH, NT, NV, NY, OH, OK, ON, PA, PE, QC, SD, SK, TN, TX, WA, WI, YT. # * Holocentropus interruptus Banks, 1914— AB, AL, BC, CO, CT, DE, IL, IN, MA, MB, ME, MI, MN, NC, ND, NE, NF, NH, NS, NT, NY, OH, ON, PA, QC, SK, TN, VA, VT, WA, WI; Holocentropus longus Banks, 1914 (Ross, 1938); Holocentropus orotus Banks, 1914 (Ross, 1938).

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Polycentropodidae continued

# * Neureclipsis crepuscularis (Walker, 1852), Brachycentrus—AB, AL, AR, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, LB, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, NF, NH, NS, NY, OH, OK, ON, PA, QC, SC, SK, TN, TX, VA, VT, WI, WV, WY; Neureclipsis parvula Banks, 1907 (Milne, 1936). * Neureclipsis melco Ross, 1947—AL, FL, GA, LA, NC, SC, TN, TX. * Neureclipsis piersoni Frazer & Harris, 1991—AL, AR, GA, KY, TN. * Nyctiophylax affinis (Banks, 1897), Polycentropus—AB, AL, AR, BC, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, ME, MI, MN, MO, MS, MT, NB, NC, ND, NF, NH, NJ, NS, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, VA, VT, WA, WI, WV, WY; Nyctiophylax zelenus Denning, 1950 (Morse, 1972). * Nyctiophylax banksi Morse, 1972—AL, CT, KY, MA, ME, MN, MS, NS, NY, ON, PA, QC, SC, TN, VA. * Nyctiophylax barrorum Morse, 1990— AL, GA. # * Nyctiophylax celta Denning, 1948—AL, AR, CT, FL, GA, IL, KY, MB, MD, ME, MI, MN, MS, NC, NY, OK, ON, PA, QC, SC, TN, TX, VA, WI, WV. * Nyctiophylax denningi Morse, 1972—AL, DE, GA, MD, MS, NC, PA, SC, TN, VA. * Nyctiophylax moestus Banks, 1911—AB, AK, AL, AR, BC, CT, DE, FL, IL, IN, KY, MA, ME, MI, MN, MT, NB, NC, NH, NJ, NS, NY, OH, OK, ON, OR, PA, QC, SC, SK, TN, VA, WI, WV. # * Nyctiophylax morsei Lago & Harris, 1983—AL, FL.

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# * Nyctiophylax nephophilus Flint, 1964— GA, NC, NY, PA, SC, TN, VA. * Nyctiophylax serratus Lago & Harris, 1985—AL, AR, FL, KY, MO, MS, OK, TN, TX. * Nyctiophylax uncus Ross, 1944—KY, MA, ME, MI, NH, NS, NY, ON, PA, QC, TN, WI.

# * Plectrocnemia cinerea (Hagen, 1861), Polycentropus—AB, AK, AL, AR, BC, CO, CT, DC, DE, FL, IA, ID, IL, IN, KS, KY, LA, MA, MB, MD, ME, MI, MN, MO, MS, MT, NB, NC, ND, NE, NF, NH, NJ, NS, NT, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, TX, UT, VA, VT, WA, WI, WV; Polycentropus canadensis Banks, 1897 (Milne, 1936); Holocentropus flavicornis Banks, 1907 (as syn. of P. canadensis ?, Betten, 1934); Plectrocnemia lutea Betten, 1934 (Milne, 1936); Plectrocnemia pallescens Banks, 1930 (Milne, 1936). * Plectrocnemia clinei Milne, 1936—AL, CT, DE, FL, MA, MD, MN, MS, NB, NC, NF, NH, NS, NY, OH, ON, PA, VA, WV. * Plectrocnemia crassicornis (Walker, 1852), Polycentropus—AL, AR, CT, DC, DE, FL, GA, ID, IL, IN, KS, KY, LA, MA, MB, ME, MI, MN, MO, MS, MT, NC, NH, NS, NY, OH, OK, ON, PA, QC, SD, SK, TN, TX, VA, WI; Plectrocnemia adironica Banks, 1914 (synonym of P. australis according to Milne, 1936); Plectrocnemia australis Banks, 1907 (Ross, 1944). * Plectrocnemia harpi Moulton & Stewart, 1993—AR, MO, OK, TN, TX, VA.

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Polycentropodidae continued

* Plectrocnemia nascotia (Ross, 1941), Polycentropus—AL, DE, FL, GA, IN, KS, KY, MA, ME, MI, MN, NB, NC, NH, NS, NY, OH, OK, ON, PA, QC, SC, TX, VA, WI. # * Plectrocnemia remota (Banks, 1911), Polycentropus—AB, AK, BC, CT, DE, IL, IN, KY, MA, MB, ME, MI, MN, MT, NF, NH, NY, OH, ON, PA, QC, SK, WA, WI, YT.

* Polycentropus alabamensis Hamilton, Harris, & Lago, 1990—AL. * Polycentropus barri Ross & Yamamoto, 1965—AL, KY, NC, PA, TN. * Polycentropus blicklei Ross & Yamamoto, 1965—AL, DE, FL, GA, KY, MD, ME, MS, NB, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, RI, SC, TN, VA. * Polycentropus carlsoni Morse, 1971—AL, NC, SC, TN, VA. * Polycentropus carolinensis Banks, 1905— KY, NC, NH, OH, ON, PA, QC, TN, VA, VT. # * Polycentropus centralis Banks, 1914—AL, AR, IL, IN, KS, KY, MN, MO, MS, NF, NS, NY, OH, OK, ON, PA, TN, TX, VA, WI. * Polycentropus chelatus Ross & Yamamoto, 1965—AL, IN, KY, MO, TN. * Polycentropus colei Ross, 1941—KY, NC, PA, QC, TN, WV. * Polycentropus confusus Hagen, 1861—AB, AL, AR, CT, DC, DE, GA, IN, KY, MA, ME, MI, MN, MO, MS, NB, NC, NF, NH, NJ, NS, NY, OH, OK, ON, PA, QC, SC, TN, VA, VT, WI, WV. * Polycentropus elarus Ross, 1944—AL, GA, IN, KY, MA, NC, NH, NY, OH, ON, PA, QC, TN, VA. * Polycentropus floridensis Lago & Harris, 1983—AL, FL. * Polycentropus maculatus Banks, 1908— CT, KY, MA, ME, NC, NF, NH, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WV.

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Polycentropodidae continued

* Polycentropus neiswanderi Ross, 1947— KY, OH. * Polycentropus pentus Ross, 1941—AL, CT, IL, KY, MB, ME, MI, MN, NH, NJ, NS, NY, OH, ON, PA, QC, TN, VA, VT, WI, WV, WY. * Polycentropus rickeri Yamamoto, 1966— AL, KY, NC, PA, TN, VA. * Polycentropus thaxtoni Hamilton & Holzenthal, 1986—GA. * Polycentropus vernus Hamilton, Harris, & Lago, 1990—AL.

Psychomyiidae

# * Lype diversa (Banks, 1914), Psychomyia— AB, AL, AR, CT, DE, FL, GA, IL, IN, KY, LA, MA, ME, MI, MN, MO, MS, NB, NC, NH, NJ, NY, OH, ON, PA, PE, QC, SC, TN, TX, VA, VT, WI, WV; Lype griselda Betten, 1934 (Milne, 1936). # + Paduniella nearctica Flint, 1967–AR, MO # * Psychomyia flavida Hagen, 1861—AB, AL, AR, AZ, BC, CA, CO, CT, DC, DE, FL, ID, IL, IN, KS, KY, MA, MB, MD, ME, MI, MN, MO, MT, NB, NC, ND, NH, NJ, NS, NY, OH, OK, ON, OR, PA, QC, SC, SD, SK, TN, UT, VA, VT, WA, WI, WV, WY; Psychomyiella composita Martynov, 1910 (Schmid, 1965); Psychomyia moesta Banks, 1907 (Milne, 1936); Psychomyia pulchella Banks, 1899 (Milne, 1936). # * Psychomyia nomada (Ross, 1938), Psychomyiella—AL, MA, ME, NC, NY, OR, PA, QC, SC, TN, VA.

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Rhyacophilidae

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

* Rhyacophila accola Flint, 1972—NC. # * Rhyacophila acutiloba Morse & Ross, 1971—MA, ME, NB, NC, NH, NY, SC, TN, VA, VT, WV. * Rhyacophila alabama Harris, 1991—AL. # * Rhyacophila amicis Ross, 1956—NC, TN. # * Rhyacophila appalachia Morse & Ross, 1971—KY, NC, SC, TN, VA. # * Rhyacophila atrata Banks, 1911—CO, CT, MA, NC, NH, NS, NY, PA, SC, TN, VA, VT. # * Rhyacophila banksi Ross, 1944—AL, AR, MD, MO, NH, NY, OH, ON, PA, QC, TN, VA, VT, WV. * Rhyacophila carolae Harris, 1989—AL. # * Rhyacophila carolina Banks, 1911—AL, CT, DE, FL, GA, KY, MA, ME, MO, MS, NB, NC, NF, NH, NJ, NY, OH, ON, PA, QC, SC, TN, VA, VT, WV; Rhyacophila carula Denning, 1948 (Ross, 1956); Rhyacophila gordoni Sibley, 1926 (Milne, 1936); # * Rhyacophila carpenteri Milne, 1936—KY, MA, NC, NF, NH, NY, OH, PA, QC, TN, VA, VT, WV. # * Rhyacophila celadon Etnier, Stocks, & Parker, 2004 (in Etnier et al., 2004)—NC, TN. # * Rhyacophila fenestra Ross, 1938—AL, AR, IL, IN, KY, MO, NC, OH, SC, TN, VT. # * Rhyacophila formosa Banks, 1911—AL, DE, GA, MA, ME, NC, NJ, NY, ON, PA, QC, SC, TN, VA, VT, WV; Rhyacophila vuphipes Milne, 1936 (Weaver, 1990). # * Rhyacophila fuscula (Walker, 1852), Neuronia—AL, CT, DE, GA, IL, KY, LB, MA, ME, MI, MN, NB, NC, NF, NH, NJ, NS, NY, ON, PA, QC, SC, TN, VA, VT, WI, WV, WY.

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Rhyacophilidae continued

# * Rhyacophila glaberrima Ulmer, 1907— AL, AR, CT, GA, IL, IN, KY, MA, ME, MO, MS, NC, NH, NY, NS, OH, PA, QC, SC, TN, VA, VT, WV; Rhyacophila andrea Betten, 1934 (Milne, 1936); Rhyacophila fairchildi Banks, 1930 (Milne, 1936). * Rhyacophila invaria (Walker, 1852), Polycentropus—CT, DE, KY, MA, MD, ME, NB, NC, NF, NH, NS, NY, ON, PA, QC, VA, VT, WI, WV; Rhyacophila luctuosa Banks, 1911 (Milne, 1936). * Rhyacophila kondratieffi Parker, 1986— NC, TN, VA. # * Rhyacophila ledra Ross, 1939—AL, DE, GA, IL, IN, KY, MI, MS, NC, OH, PA, SC, TN, VA, WV; Rhyacophila hardeni Denning, 1948 (Ross, 1956). # * Rhyacophila lobifera Betten, 1934—AL, AR, IL, IN, KS, KY, MO, OH, OK, ON, PA, TN, WI, WV. # + Rhyacophila mainensis Banks, 1911—MA, ME, MI, NB, NF, NH, NJ, NY, ON, PA, QC, VA, VT, WV; Rhyacophila melita Ross, 1938 (Weaver, 1990). # * Rhyacophila minora Banks, 1924—AL, CT, KY, MA, ME, NC, NF, NH, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WV. # * Rhyacophila montana Carpenter, 1933— NC, TN. * Rhyacophila mycta Ross, 1941—NC, SC, TN, VA. # * Rhyacophila nigrita Banks, 1907—AL, GA, KY, MA, ME, NC, NF, NH, NJ, NY, ON, PA, QC, SC, TN, VA, VT, WV. * Rhyacophila otica Etnier & Way, 1973— KY, PA, TN; Rhyacophila pennsylvanica Sykora & Weaver, 1976 (Weaver & Sykora, 1979).

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Rhyacophilidae continued

# * Rhyacophila parantra Ross, 1948—IN, KY, MO, OH, QC, TN, VA. # + Rhyacophila shenandoahensis Flint, 1958—ON, VA. # + Rhyacophila simmonsi Armitage, 2008— VA. * Rhyacophila teddyi Ross, 1939—AL, GA, NC, SC, TN. # * Rhyacophila torva Hagen, 1861—AL, CT, DC, DE, KY, MA, ME, NC, NF, NH, NJ, NS, NY, OH, PA, QC, SC, TN, VA, VT, WI, WV, WY; Rhyacophila terminata Banks, 1907 (Fischer, 1960); Rhyacophila vinura Milne, 1936 (Ross, 1944). * Rhyacophila tricornuta Sykora & McCabe, 1995—NC, VA. * Rhyacophila vibox Milne, 1936—CT, IL, IN, KY, MA, MI, MN, NC, NF, NH, NJ, NY, OH, ON, PA, QC, TN, VA, VT, WI, WV.

Sericostomatidae

* Agarodes (A.) alabamensis Harris, 1986— AL. # * Agarodes (A.) crassicornis (Walker, 1852), Hydropsyche—AL, FL, GA, LA, MD, MS, NC, SC, TX, VA. # * Agarodes (Psiloneura) distinctus Ulmer, 1905—CT, GA, MA, ME, MN, NH, NY, ON, QC, SC, TN, WI; Schizopelex lobata Banks, 1911 (Milne, 1936); Psiloneura moesta Banks, 1914 (Milne, 1936). # * Agarodes (A.) georgia Ross & Scott, 1974—GA, TN. # * Agarodes (A.) griseus Banks, 1899—AL, CT, DC, DE, FL, GA, MA, ME, MS, NC, NH, NS, NY, PA, QC, SC, VA; Notidobia americana Banks, 1900 (Milne, 1936).

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# * Agarodes (Psiloneura) libalis Ross & Scott, 1974—AL, DE, FL, LA, MS, NC, SC, TX, VA. # * Agarodes (A.) logani Keth & Harris, 1999—FL. # * Agarodes (A.) stannardi (Ross, 1962), Sericostoma—AL, KY, MS, TN. # * Agarodes (A.) tetron (Ross, 1948), Sericostoma—GA, NC, SC, TN. * Agarodes (A.) tuskaloosa Keth & Harris, 1999—AL. # * Agarodes (A.) wallacei Ross & Scott, 1974—FL, NC, SC. * Agarodes (A.) ziczac Ross & Scott, 1974— FL. # * Fattigia pele (Ross, 1938), Notidobia— GA, NC, SC, TN, VA.

Thremmatidae

# * Neophylax acutus Vineyard & Wiggins, 1987—AL, KY, TN, VA. # * Neophylax aniqua Ross, 1947—KY, ME, NB, NC, NF, NH, NS, NY, ON, PA, PE, QC, TN, VA, VT, WV. #* Neophylax atlanta Ross, 1947—AL, GA, MS, NC, SC, VA; Neophylax saloris Denning, 1948 (Denning, 1948). # * Neophylax ayanus Ross, 1938—IN, KY, MI, OH, TN. # * Neophylax concinnus McLachlan, 1871— AL, AR, CT, DE, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, MT, NC, NH, NJ, NY, OH, ON, PA, QC, RI, SC, TN, VA, VT, WI, WV, WY; Neophylax autumnus Vorhies, 1909 (Kimmins & Denning, 1951). # * Neophylax consimilis Betten, 1934—GA, KY, MA, MD, ME, NC, NH, NS, NY, PA, QC, SC, TN, VA, VT, WV; Neophylax slossonae Banks, 1943 (Vineyard et al., 2005). # * Neophylax etnieri Vineyard & Wiggins, 1987—KY, TN, VA; Neophylax auris Vineyard & Wiggins, 1987 (Vineyard et al., 2005).

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Thremmatidae continued

# * Neophylax fuscus Banks, 1903—AL, AR, CT, IL, IN, KY, MA, ME, MI, MN, MO, NB, NC, NH, NS, NY, OH, ON, PA, QC, TN, VA, VT, WI, WV. * Neophylax harrisi Schefter & Frania, 2005, in Vineyard et al., 2005–AL. * Neophylax kolodskii Parker, 2000—TN. # * Neophylax lewisae Etnier, 2006—KY, TN. # * Neophylax mitchelli Carpenter, 1933—DE, GA, KY, MA, MD, ME, MN, NB, NC, NF, NH, NS, NT, NY, ON, PA, QC, SC, TN, VA, VT; Neophylax delicatus Banks, 1943 (Vineyard et al., 2005); Neophylax nacatus Denning, 1941 (Vineyard et al., 2005). # * Neophylax oligius Ross, 1938—AL, CT, DE, GA, MA, MD, ME, MI, MN, MS, NC, NF, NH, NJ, NS, NY, OH, ON, PA, QC, SC, TN, VA, VT, WI, WV. # * Neophylax ornatus Banks, 1920—AL, GA, NB, NC, NF, NH, NJ, NS, NY, ON, PA, PE, QC, SC, TN, VA, VT, WV. # * Neophylax securis Vineyard & Wiggins, 1987—AL, TN. # * Neophylax stolus Ross, 1938—NY, PA, TN, VA, WV. # * Neophylax toshioi Vineyard & Wiggins, 1987—TN, VA. # * Neophylax wigginsi Sykora & Weaver, 1978—KY, OH, PA, TN, VA, WV.

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KEY to families (some genera and species) of TRICHOPTERA LARVAE FAMILIES

(adapted from the key of Wiggins & Currie, 2008) 1 1’



2(1’) 2’

Larva constructing snail-shell-shaped portable case of sand grains or small rock fragments (Fig. 4.1); each anal claw comb-like, with end hook no larger than other teeth (Fig. 4.2) ......................................................................... HELICOPSYCHIDAE, Helicopsyche, 61 Larva constructing cylindrical, dome-shaped, or purse-like portable case of various materials or not constructing portable case; each anal claw without subapical teeth (similar to Fig. 4.3) or with ventral subapical teeth (Figs. 4.4, 4.6) or dorsal accessory spines (Fig. 4.5), and always ending in conspicuously larger, stout apical hook ............... 2 Metanotum entirely covered by 1 sclerite or 1 pair of sclerites (Fig. 4.7) ................................ 3 Metanotum entirely membranous (Fig. 4.8) or with more than 2 smaller sclerites (Fig. 4.9) or with 2 narrow transverse sclerites (Figs. 4.596, 4.597, 4.602–4.605) ............................ 4

3(2) Abdomen with ventromesal rows of branched gills (Fig. 4.10) and with prominent tuft of long setae at base of each anal proleg; living in fixed retreat of detritus and rock fragments and with accompanying filter net ............................ HYDROPSYCHIDAE, 62 3’ Abdomen lacking ventromesal rows of branched gills and tufts of setae at bases of prolegs; free-living or living in flattened (Figs. 4.11, 4.12) or cylindrical (Fig. 4.13) portable case or in flattened silken dome fastened to rock (Fig. 4.14)...... HYDROPTILIDAE, 117

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Fig. 4.1

Fig. 4.2

Fig. 4.3

Fig. 4.6

Fig. 4.4 Fig. 4.5

Fig. 4.7

Fig. 4.11

Fig. 4.8

Fig. 4.12

Fig. 4.9

Fig. 4.13

Fig. 4.10

Fig. 4.14

Figures 4.1–4.14, Helicopsychidae, Psychomyiidae, Polycentropodidae, Limnephilidae, Hydropsychidae, and Hydroptilidae genn. spp. 4.1–4.2, Helicopsyche borealis (Helicopsychidae): 4.1, case; 4.2, left anal claw, left lateral. 4.3–4.5, left anal claw, left lateral: 4.3, Tinodes sp. (Psychomyiidae); 4.4, Polyplectropus sp. (Polycentropodidae); 4.5, Polycentropus sp. (Polycentropodidae). 4.6, Neureclipsis sp. (Polycentropodidae) left anal proleg, anal claw enlarged, left lateral. 4.7–4.9: thoracic nota, dorsal: 4.7, Palaeagapetus celsus (Hydroptilidae); 4.8, Polycentropus sp. (Polycentropodidae); 4.9, Anabolia bimaculata (Limnephilidae). 4.10, Parapsyche cardis (Hydropsychidae), abdominal segment IV, left lateral. 4.11–4.14, Hydroptilidae, cases: 4.11, Stactobiella delira; 4.12, Oxyethira sp.; 4.13, Mayatrichia ayama; 4.14, Leucotrichia sp. All figures © R.W. Holzenthal 2006.

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4(2’)

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species



Antennae markedly elongate and prominent, at least 6 times as long as wide (similar to Fig. 4.15) and/or sclerotized plates on mesonotum lightly pigmented except for pair of dark, curved lines, concave laterally, on posterior halves of plates (Fig. 4.17); constructing case of various shapes and materials (Fig. 4.18) ........................... LEPTOCERIDAE, 139 Antennae not more than 3 times as long as wide (Fig. 4.16) or not evident; mesonotum never with pair of dark, curved lines ................................................................................... 5

5(4’) 5’

Mesonotum without sclerites or with sclerites small, not covering more than half of mesonotum (Fig. 4.19); pronotum without anterolateral lobes (Fig. 4.23).......................... 6 Mesonotum mostly covered by 1–4 pairs of sclerotized plates (Figs. 4.20, 4.21); pronotum sometimes with prominent anterolateral lobes (Figs. 4.24) .............................................. 12

6(5) 6’

Abdominal tergum IX with sclerite (Fig. 4.25), sometimes pale or covered by tergum VIII if segment IX is partially retracted ...................................................................................... 7 Abdominal tergum IX entirely membranous ............................................................................ 9

7(6)

Metanotal sa3 each usually with cluster of setae arising from small rounded or ovoid sclerite (similar to Fig. 4.22); prosternal horn present (Fig. 4.26); constructing tubular portable case of plant materials (Fig. 4.27) .................................................PHRYGANEIDAE, 254

4’



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Fig. 4.19

Page 287

Fig. 4.16

Fig. 4.17

Fig. 4.20

Fig. 4.23

Fig. 4.18

Fig. 4.21

Fig. 4.24

Fig. 4.22

Fig. 4.25

Fig. 4.26 Fig. 4.27 Figures 4.15–4.27, Leptoceridae, Phryganeidae, Glossosomatidae, Limnephilidae, and Calamoceratidae genn. spp. 4.15, Triaenodes frontalis (Leptoceridae), head, dorsal. 4.16, Oligostomis ocelligera (Phryganeidae), right anterolateral corner of head, dorsal. 4.17, Ceraclea sp. (Leptoceridae), thorax, dorsal. 4.18, Leptoceridae genn. spp., larval cases. 4.19–4.22, mesonota, dorsal: 4.19, Agapetus sp. (Glossosomatidae); 4.20, Anabolia bimaculata (Limnephilidae); 4.21, Goeracea genota (Goeridae); 4.22, Hagenella canadensis (Phryganeidae). 4.23–4.24, pronota, dorsal: 4.23, Nemotaulius hostilis (Limnephilidae); 4.24, Heteroplectron americanum (Calamoceratidae). 4.25, Glossosoma nigrior (Glossosomatidae), abdominal segments IX and X, caudal. 4.26, Agrypnia vestita (Phryganeidae), prosternum, ventral. 4.27, Phryganeidae genn. spp., 3 cases. All figures © R.W. Holzenthal 2006.

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7’



8(7’) 8’

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Metanotal sa3 each with single seta and with or without sclerite (Fig. 4.28); free-living or retreat-making or constructing domed, portable case of rock fragments (Fig. 4.29) .......... 8 Anal prolegs each with basal half broadly fused with segment IX (Fig. 4.30); dorsal edge of each anal claw with at least one tiny accessory spine arising from it; constructing small domed, portable case of rock fragments (Fig. 4.29) ..... GLOSSOSOMATIDAE, 48 Anal prolegs each mostly free from segment IX (similar to Fig. 4.31); dorsal edges of anal claws without accessory hooks, although ventral teeth and/or a secondary, lateral hook may be present; free-living, without fixed retreat or case until pupation .......................................................................... RHYACOPHILIDAE, Rhyacophila, 279

9(6’) 9’

Labrum pale, membranous, and vaguely T-shaped (Fig. 4.32), often withdrawn from view in preserved specimens and apparently absent; constructing fixed sac-like, fine-meshed filter net of silk on undersides of rocks ................................... PHILOPOTAMIDAE, 243 Labrum sclerotized, rounded, articulated on anterior edge of frontoclypeus, never withdrawn .... 10

10(9’) 10’

Foretrochantin broadened apically, hatchet-shaped (Fig. 4.33); constructing fixed tubular retreat of sand and debris on rocks and logs .............................. PSYCHOMYIIDAE, 276 Foretrochantin acute apically (similar to Fig. 4.34) ................................................................ 11

11(10’)

Tarsi of all legs conspicuously flattened, tibiae shorter than tarsi (Fig. 4.35); constructing branching-tube retreat of silk covered with sand and buried in fine sand, except tips of branches exposed; collectors-filterers; burrowers usually in fine sediments of loticdepositional habitats ........................................ DIPSEUDOPSIDAE, Phylocentropus, 46 Tarsi of all legs tubular, tibiae longer than tarsi (Fig. 4.36); constructing exposed flattened, tubular, or funnel-like fixed retreat ................................ POLYCENTROPODIDAE, 263

11’

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Fig. 4.28 Fig. 4.29

Fig. 4.31

Fig. 4.34

Fig. 4.32

Fig. 4.30

Fig. 4.33

Fig. 4.35 Fig. 4.36

Figures 4.28–4.36, Rhyacophilidae, Glossosomatidae, Polycentropodidae, Philopotamidae, Psychomyiidae, Philopotamidae, and Dipseudopsidae genn. spp. 4.28, Rhyacophila sp. (Rhyacophilidae), metanotum, dorsal. 4.29, Protoptila sp. (Glossosomatidae), case, ventrolateral. 4.30, Glossosoma nigrior (Glossosomatidae), terminal abdominal segments and anal claws, caudal. 4.31, Neureclipsis sp. (Polycentropodidae), left anal proleg, left lateral. 4.32, Chimarra sp. (Philopotamidae), labrum and anterior edge of frontoclypeus, dorsal. 4.33, Psychomyiidae gen. sp., right foretrochantin, right lateral. 4.34, Dolophilodes sp. (Philopotamidae), right foretrochantin and prothoracic coxa, right lateral. 4.35–4.36, Dipseudopsidae and Polycentropodidae genn. spp., left prothoracic tibiae, tarsi, and tarsal claws, left lateral: 4.35, Phylocentropus sp. (Dipseudopsidae); 4.36, Polycentropus sp. (Polycentropodidae). 4.32 and 4.33 originals; all other figures © R.W. Holzenthal 2006.

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12(5’) 12’

Abdominal segment I without either dorsal or lateral humps (Fig. 4.37); metanotal sa1 absent (Fig. 4.41); constructing portable case usually of plant material but occasionally of sand or silk, either tubular or 4-sided ............................... BRACHYCENTRIDAE, 24 Abdominal segment I with at least lateral humps (Fig. 4.38, 4.40), sometimes collaped and evidenced by only crease or fold; metanotum sa1 usually represented by anterior, submesal pair of sclerites with one to several setae (Fig. 4.42); constructing more-or-less tubular portable case (Figs. 4.43–4.46) ................................... 13

13(12’) 13’

Mesepisterna extended anterad as prominent, pointed processes (similar to Fig. 4.47); constructing tubular case of rock fragments (Fig. 4.45) usually with larger lateral ballast stones (Fig. 4.46) ...................................................................................... GOERIDAE, 56 Mesepisterna not extended anterad as prominent, pointed processes (Fig. 4.48) ................... 14

14(13’) 14’

Labrum with transverse row of about 16 large setae across central area (Fig. 4.49); constructing depressed case of 2 dead leaves or gouging piece of wood or pith of stick to serve as case ...................................................................... CALAMOCERATIDAE, 45 Labrum with transverse row of 6 setae across central area (Fig. 4.50) .................................. 15

15(14’)

Antennae situated near anterior margins of eyes (Fig. 4.51); abdominal segment I without median dorsal hump (Fig. 4.38); constructing variously shaped case of dead plant material or small sand grains, but most commonly 4-sided with quadrate leaf panels (Fig. 4.39) .......................................................................... LEPIDOSTOMATIDAE, 136 Antennae situated at anterior edge of head capsule (Figs. 4.52, 4.53) or about halfway between eyes and anterior edge of head capsule (Fig. 4.54); abdominal segment I with lateral and dorsal humps (Fig. 4.40); case variable, but never as in Fig. 4.39 .................. 16

15’

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Fig. 4.37

Fig. 4.41

Fig. 4.38

Fig. 4.39

Fig. 4.42 Figs. 4.43

Fig. 4.47

Fig. 4.40

Fig. 4.48

Fig. 4.51

Fig. 4.49

4.44

4.45

4.46

Fig. 4.50

Fig. 4.53 Fig. 4.52

Fig. 4.54

Figures 4.37–4.54, Brachycentridae, Lepidostomatidae, and Limnephilidae, genn. spp. 4.37–4.38, metathorax and abdominal segments I & II, left lateral: 4.37, Brachycentridae gen. sp.; 4.38, Lepidostomatidae gen. sp. 4.39, Lepidostoma (Lepidostomatidae), case. 4.40, Psychoglypha sp. (Limnephilidae), abdominal segment I, left lateral. 4.41–4.42, Brachycentridae and Limnephilidae genn. spp., metanota, dorsal: 4.41, Brachycentrus sp. (Brachycentridae); 4.42, Pycnopsyche sp. (Limnephilidae). 4.43–4.46, Limnephilidae and Goeridae, genn. spp., cases: 4.43, Anabolia bimaculata (Limnephilidae); 4.44, Ironoquia sp. (Limnephilidae); 4.45, Goerita semata (Goeridae); 4.46, Goera fuscula (Goeridae), case. 4.47–4.48, Goeridae and Limnephilidae genn. spp., pronota and mesonota, dorsal: 4.47, Goerita semata (Goeridae); 4.48, Anabolia bimaculata (Limnephilidae). 4.49–4.50, Calamoceratidae and Limnephilidae genn. spp., labra, dorsal: 4.49, Calamoceratidae gen. sp.; 4.50, Limnephilidae gen. sp.; 4.51, Lepidostomatidae gen. sp. head, right lateral. 4.52, Oligostomis ocelligera (Phryganeidae), right anterolateral corner of head, dorsal. 4.53, Agarodes libalis (Sericostomatidae), head, left lateral. 4.54, Limnephilus sp. (Limnephilidae), head and prosternal horn, left lateral. 4,37, 4.38. 4.41, 4.49, and 4.50 originals; 4.39 redrawn from Wiggins (1996); all other figures © R.W. Holzenthal 2006.

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16(15’) Antennae situated about halfway between eyes and anterior edge of head capsule (Fig. 4.54); prosternal horn usually present (Fig. 4.54) but may be short (similar to Fig. 4.55); chloride epithelia usually present on at least some abdominal segments (Figs. 4.56, 4.57) ............................................................................................................... 17 16’ Antennae situated at anterior edge of head capsule (Figs. 4.52, 4.53); prosternal horn and chloride epithelia absent ................................................................................................... 19 17(16) 17’

Mesonotum with anteromesal emargination (Fig. 4.58); constructing stout case of rock fragments, usually with small ballast stones laterally (similar to Fig. 4.64) ..................................................................................THREMMATIDAE, Neophylax, 304 Mesonotum not notched anteromesally .................................................................................. 18

18(17’) 18’

Mandibles usually without teeth (Fig. 4.59); basal seta of each tarsal claw as long as claw (Fig. 4.61); constructing tapered and curved case of rock fragments (similar to Fig. 4.65), rarely with small twigs and leaf fragments trailing from top and sides ............................................................................................................. APATANIIDAE, 22 Mandibles usually with teeth (similar to Fig. 4.60); basal seta of each tarsal claw much shorter than claw (similar to Fig. 4.62); constructing variously shaped case of plant (Fig. 4.66) and/or mineral materials (Fig. 4.68) ......................... LIMNEPHILIDAE, 211

19(16’) 19’

Tarsal claws of hind legs conspicuously different from those of middle and forelegs, much shorter than tarsi (Fig. 4.63); constructing case of sand grains with lateral flanges and anterodorsal hood (Fig. 4.67) .......................................... MOLANNIDAE, Molanna, 234 Tarsal claws of hind legs similar to those of other legs, nearly as long as tarsi (similar to Fig. 4.69) ........................................................................................................................... 20

20(19’)

Pronotum with transverse ridge extended onto rounded anterolateral lobes (Figs. 4.70, 4.71), constructing tubular, slightly tapered and curved case of fine sand grains (similar to 4.68) ; probably collectors-gatherers; sprawlers in seeps; see also Hamilton (1985); GA ..................................................................................... BERAEIDAE, Beraea gorteba Pronotum without transverse ridge and with anterolateral corners acute (Figs. 4.72, 4.73) ..... 21

20’

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Fig. 4.55

Fig. 4.56

Fig. 4.57 Fig. 4.58

Fig. 4.59

Fig. 4.60

Fig. 4.61 Fig. 4.62 Fig. 4.63

Fig. 4.69

Fig. 4.70

Figs. 4.64

Fig. 4.71

4.65

4.66

Fig. 4.72

4.67

4.68

Fig. 4.73

Figures 4.55–4.73, Phryganeidae, Limnephilidae, Thremmatidae, Apataniidae, Molannidae, Goeridae, and Molannidae, genn. spp. 4.55, Agrypnia vestita (Phryganeidae), head and prothorax, ventral. 4.56, Halesochila taylori (Limnephilidae), abdomen, with insets of dorsal chloride epithelia, left lateral. 4.57, Dicosmoecus sp. (Limnephilidae), abdominal sternum II, ventral. 4.58, Neophylax sp. (Thremmatidae), pronotum and mesonotum, dorsal. 4.59–4.60, Apataniidae genn. spp., left mandibles, ventral: 4.59, Manophylax annulatus; 4.60, Moselyana comosa. 4.61–4.62, Apataniidae genn. spp., left metathoracic tarsal claws, left lateral: 4.61, Manophylax annulatus; 4.62, Moselyana comosa. 4.63, Molanna tryphena (Molannidae), right metathoracic tarsus, right lateral. 4.64–4.68, Apataniidae, Limnephilidae, and Molannidae genn. spp., cases: 4.64, Goera fuscula (Goeridae); 4.65, Pedomoecus sierra (Apataniidae); 4.66, Anabolia bimaculata (Limnephilidae); 4.67, Molanna sp. (Molannidae); 4.68, Ecclisocosmoecus scylla (Limnephilidae). 4.69, Psilotreta sp. (Odontoceridae), left prothoracic leg, left lateral. 4.70, Beraea gorteba (Beraeidae), pronotum, dorsal. 4.71–4.73, pronota, left lateral: 4.71, Beraea gorteba (Beraeidae); 4.72, Psilotreta sp. (Odontoceridae); 4.73, Agarodes libalis (Sericostomatidae). All figures © R.W. Holzenthal 2006.

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21(20’) 21’

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Anal proleg with cluster of about 30+ setae dorsally, lateral sclerites relatively small (Fig. 4.74); foretrochantin large and hooked (Fig. 4.76); constructing tubular, slightly tapered and curved case of fine sand grains (Fig. 4.78) ..... SERICOSTOMATIDAE, 295 Anal proleg with 3–5 setae dorsally, lateral sclerites large (Fig. 4.75); foretrochantin small and triangular (Fig. 4.76); constructing case of rock fragments (Figs. 4.79, 4.80) ....................................................................................................ODONTOCERIDAE, 236

GENERA AND SPECIES

(adapted from the keys of Morse & Holzenthal, 2008, unless otherwise indicated)

APATANIIDAE, key to genera and species 22(18) 22’

Metanotal sa1 sclerites present, about same size as sa2 sclerites (Fig. 4.81); case of rock fragments, somewhat depressed, tapered and slightly curved, with plant pieces dorsally and laterally; scrapers; clingers in madicolous mountain habitats ................................... ................................................................................................ Manophylax (Madeophylax), 23 Metanotal sa1 sclerites absent, sa1 setae forming transverse row (Fig. 4.82); case of rocks, strongly tapered, anterior opening usually oblique with dorsal edge extending beyond ventral edge for final instar; scrapers and collectors-gatherers; clingers, climbers, and sprawlers in cold mountain streams (Chen, 1993); QC to NH and south to NC, SC, TN ................................................................................................................... Apatania incerta

Manophylax (subgenus Madeophylax), key to species (from Schuster, 1997)

23(22) Pebbling on frontoclypeus generally covering entire sclerite (Fig. 4.83); abdominal sternum I with transverse sclerite linear (Fig. 4.85); dry to moist rock walls 1500 m elevation in NC, WV ....................................................................................... Manophylax (Madeophylax) altus

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Fig. 4.75

Fig. 4.76

Fig. 4.74

Fig. 4.81

Fig. 4.77 Fig. 4.78

Fig. 4.79

Fig. 4.80

Fig. 4.82

Fig. 4.85

Fig. 4.83

Fig. 4.84

Fig. 4.86

Figures 4.74–4.86, Sericostomatidae Odontoceridae, Apataniidae, and Brachycentridae genn. spp. 4.74, Sericostomatidae gen. sp., abdominal segments IX and X and anal prolegs, dorsal. 4.75, Odontoceridae gen. sp., abdominal segments IX and X and anal prolegs, dorsal. 4.76, Sericostomatidae gen. sp., foretrochantin, right lateral. 4.77, Odontoceridae gen. sp., foretrochantin, right lateral. 4.78, Sericostomatidae gen. sp., case. 4.79, Pseudogooera singularis (Odontoceridae), case. 4.80, Psilotreta sp. (Odontoceridae), case. 4.81–4.82, Apataniidae genn. spp., metanota, dorsal: 4.81, Manophylax annulatus; 4.82, Apatania arizona. 4.83–4.84, Manophylax spp., head, dorsal: 4.83, M. butleri; 4.84, M. altus. 4,85–4.86, Manophylax spp., abdominal sterna I, ventral: 4.85, M. butleri; 4.86, M. altus. 4.74–4.78 original; 4.79–4.82, © R.W. Holzenthal 2006; 4.83–4.86 redrawn from Schuster (1997).

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

BRACHYCENTRIDAE, key to genera and species 24(12)

24’

Meso- and metathoracic legs long, their femora about as long as head capsule, their tibiae each produced distally into prominent process from which stout spur arises (Fig. 4.87); case usually square in cross section, composed of small pieces of plant materials fastened transversely (Fig. 4.89a), although case sometimes cylindrical and largely of silken secretion, or occasionally of small rock fragments (Figs. 4.89b, 4.93); collectors-filterers, collectors-gatherers, shredders-herbivores; clingers in streams ............ Brachycentrus, 26 Meso- and metathoracic legs shorter, their femora much shorter than head capsule (Fig. 4.88), each tibia not produced distally into prominent process, although spur arising from about same point on unmodified tibia (Fig. 4.88) ........................................ 25

25(24’) Ventral apotome of head longer than wide, narrowed somewhat posteriorly, rudimentary prosternal horn present on anterior part of prosternum (Fig. 4.90); case 4-sided and tapered, composed of short pieces of plant material placed crosswise with loose ends often protruding (Fig. 4.94); probably shredders-herbivores; clingers and climbers in stream moss; CT, PA, MD, NY, VA, WV ...................................... Adicrophleps hitchcocki 25’ Ventral apotome of head usually wider than long (Fig. 4.91), sometimes squarish (Fig. 4.92) or rectangular and longer than broad; prosternal horn absent; case cylindrical, tapered, straight or curved, composed of lengths of plant material wound around circumference (Fig. 4.95) or of silk or silk and rock material (Fig. 4.96); shredders-herbivores and collectors-gatherers; clingers and sprawlers on dead or living plant material in streams, including mosses .......................................................................................... Micrasema, 37

Brachycentrus, key to subgenera and species (from Flint, 1984; Harrington & Morse, 2004)

26(24) 26’

Mid- and hind femora each with 2 major setae on ventral margin (Fig. 4.97); ventral filtering fringe of each mid- and hind femur in fan-like arrays (Fig. 4.97); see also Lloyd, 1915c, 1921 as B. nigrisoma; Appalachian Mtns. and upper Coastal Plain from ME to AL, GA, KY, NC, SC, TN .............. Subgenus Brachycentrus, Brachycentrus (B.) nigrosoma Mid- and hind femora each with 1 major seta on ventral margin (Fig. 4.98); ventral filtering fringe of each femur nearly uniform in length (Fig. 4.98) ..... Subgenus Sphinctogaster, 27

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Fig. 4.87

Fig. 4.88

Fig. 4.90

Fig. 4.91

Fig. 4.93

Fig. 4.97

Fig. 4.94

Figs. 4.89a

4.89b

Fig. 4.92

Fig. 4.95

Fig. 4.96

Fig. 4.98

Figures 4.87–4.98, Brachycentridae genn. spp. 4,87–4.88, Brachycentridae, genn. spp., right metathoracic legs, right lateral: 4.87, Brachycentrus sp.; 4.88, Micrasema wataga. 4.89, Brachycentrus spp. (Brachycentridae), cases. 4.90, Adicrophleps hitchcocki (Brachycentridae), head and prosternum, ventral. 4.91–4.92, heads, ventral: 4.91, Amiocentrus aspilus; 4.92, Micrasema wataga. 4.93–4.96, cases: 4.93, Brachycentrus etowahensis; 4.94, Adicrophleps hitchcocki; 4.95, Micrasema wataga; 4.96, Micrasema rusticum; 4.97–4.98, left hind legs, left lateral: 4.97, B. nigrosoma; 4.98, B. appalachia. 4.93 from Wallace (1971); 4.97–4.98 from Flint (1984); all other figures © R.W. Holzenthal 2006.

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27(26’) 27’

Mid- and hind tibiae each with 1 large basomesal seta (Figs. 4.98, 4.99) .............................. 28 Mid- and hind tibiae each with row of 3–5 mesal setae (Figs. 4.100, 4.101) ......................... 29

28(27) 28’

Mid- and hind tibiae fuscous for basal 1/3rd, pale apically (Fig. 4.98); tarsi fuscous (Fig. 4.98); northeastern and Appalachian Mtn. region from ON to NS and south to GA, NC, SC, TN .......................................... Brachycentrus (Sphinctogaster) appalachia Mid- and hind tibiae pale with narrow, fuscous dorsal stripe (Fig. 4.99); tarsi pale (Fig. 4.99); ON to ME and south to AR and KY, NC, SC, TN ............................................ ............................................................................ Brachycentrus (Sphinctogaster) lateralis

29(27’) 29’

Head uniformly dark brown or fuscous (Fig. 4.102), rarely paler over muscle scars (Fig. 4.103) ....................................................................................................................... 30 Head distinctly banded or spotted with fuscous and yellow marks (Figs. 4.104–4.106) ....... 31

30(29) Head uniformly brown to nearly black, in frontal aspect nearly circular in outline (Fig. 4.102), front evenly and finely granulate; southwestern NC, northeastern GA, northwestern SC .............. Brachycentrus (Sphinctogaster) spinae (“dark-headed” race) 30’ Head brownish fuscous, often with genae laterally and posteriorly paler than front; in frontal aspect elongate, sides subparallel (Fig. 4.103), front rugose; upper Coastal Plain of AL, FL, GA, NC, SC ................................... Brachycentrus (Sphinctogaster) chelatus 31(29’) 31’ 32(31’) 32’ 33(32’) 33’

Mid- and hind femora each with 2 or more major setae on each of anterior and posterior faces (Fig. 4.100); case of sand grains with projecting mineral pieces (Fig. 4.93); see also Wallace (1971); northern GA, southeastern ............................................................. .............................................................. TN Brachycentrus (Sphinctogaster) etowahensis Mid- and hind femora each with only 1 major seta on each of anterior and posterior faces (Figs. 4.101, 4.107, 4.108) ................................................................................................ 32 Mesal fuscous band of frontoclypeus separated from inner fuscous bands of genae, or at most merely touching them near posterior apex of frontoclypeus, so that head generally with 5 pale stripes (Fig. 4.104); WI to QC and south to NC ........................................... ............................................................................ Brachycentrus (Sphinctogaster) incanus Mesal fuscous band broadly confluent with inner fuscous bands of genae for posterior 1/3rd of frontoclypeus, with cluster of pale muscle scars or without conspicuous pale spot at posterior apex of frontoclypeus so that head has 2–4 pale stripes (Fig. 4.105) ................ 33 Frontoclypeus with cluster of pale muscle scars near posterior apex (Fig. 4.105); mid- and hind femora generally pale with fuscous dorsal and ventral margins (Fig. 4.101); MB to ME and south to CO, TX and AL, FL, KY, MS, NC, SC, TN .................................. ....................................................................... Brachycentrus (Sphinctogaster) numerosus Frontoclypeus without pale spot(s) at posterior apex (Fig. 4.106); mid- and hind femora unicolorous (Fig. 4.107) or more commonly with dorsal half broadly brown (Fig. 4.108) ........................................................................................................................34

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Fig. 4.99

Fig. 4.100

Fig. 4.102

Fig. 4.101

Fig. 4.103

Fig. 4.105

Fig. 4.107

Fig. 4.104

Fig. 4.106

Fig. 4.108

Figures 4.99–4.108, Brachycentrus spp. (Brachycentridae). 4.99–4.101, right hind legs, right lateral: 4.99, B. lateralis; 4.100, B. etowahensis; 4.101, B. numerosus. 4.102–4.106, heads, dorsal: 4.102, B. spinae “dark-headed” race; 4.103, B. chelatus; 4.104, B. incanus; 4.105, B. numerosus; 4.106, B. solomoni. 4.107–4.108, right hind legs, right lateral: 4.107, B. spinae “dark-headed” race; 4.108 B. solomoni. All figures from Flint (1984).

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34(33’) 34’

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species



Pale dorsal marks of frontoclypeus restricted to anterior half of frontoclypeus (Fig. 4.109); pale oval spot posteriorly on each gena completely surrounded by fuscous posteriorly (Fig. 4.107); NY, PA, QC, VA, VT, WV .......... Brachycentrus (Sphinctogaster) solomoni Pale dorsal marks of frontoclypeus either restricted to anterior half of frontoclypeus (Fig. 4.110) or more elongate and generally evident to posterior apex (Fig. 4.112); pale oval spot posteriorly on each gena either absent (Fig. 4.111) or faint and reaching from midlength of head nearly or completely to occiput (posterior region) (Fig. 4.112) ......... 35

35(34’) 35’

Pale markings of frontoclypeus and genae large and clearly defined (Fig. 4.110); eastern Blue Ridge Escarpment in NC and SC ............... Brachycentrus (Sphinctogaster) lunatus Pale markings of frontoclypeus small (Fig. 4.111) or large and somewhat obscure (Fig. 4.112); pale markings of genae absent (Fig. 4.111) or obscure (Fig. 4.112) ............ 36

36(35’) 36’

Pale oval spots of frontoclypeus small and of genae absent (Fig. 4.111); northeastern GA ......................................... Brachycentrus (Sphinctogaster) spinae (“intermediate” race) Pale oval spots of frontoclypeus and genae large and obscure (Fig. 4.112); Appalachian Mtns. of southwestern VA and of NC and TN border region north of 35.6°N latitude ................................................... Brachycentrus (Sphinctogaster) spinae (“typical” race)

Micrasema, key to species (from Chapin, 1978)

37(25’) 37’

Mesonotal sclerites partially or completely divided into 4 plates (Figs. 4.113–4.115); ventral apotome rectangular, broader than long (Fig. 4.117) or longer than broad (Fig. 4.118); east of Great Plains ............................................................. Micrasema rusticum Group, 38 Mesonotal sclerites entire, consisting of 2 plates (Fig. 4.116); ventral apotome trapezoidal (Fig. 4.119); small cold streams, typically in Fontinalis mats grazing on moss leaves and detritus; QC to NB and south to NC and SC ................................ Micrasema sprulesi

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Fig. 4.109

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Fig. 4.110

Fig. 4.111

Fig. 4.112

Fig. 4.114

Fig. 4.115

Fig. 4.116

Fig. 4.113

Figs. 4.117a

4.117b

Fig. 4.118

4.117c

Fig. 4.119

Figures 4.109–4.119 Brachycentridae genn. spp. 4.109–4.112, Brachycentrus spp., heads, dorsal: 4.109, B. solomoni; 4.110, B. lunatus; 4.111, B. spinae “intermediate” race; 4.112, B. spinae “typical” race. 4.113–4.116, Micrasema spp., mesonota, dorsal: 4.113, M. burksi; 4.114, M. wataga; 4.115, M. rusticum; 4.116, M. sprulesi. 4.117–4.119, Micrasema spp. ventral apotomes of heads, ventral: 4.117, M. spp., 3 ventral apotomes; 4.118, M. burksi; 4.119, M. sprulesi. 4.109, 4.112 from Flint (1984); 4.110–4.111 redrawn from Harrington & Morse (2004); 4.113–4.119 from Chapin (1978).

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38(37)

Ventral apotome longer than broad (Fig. 4.120); head dark brown, muscle scars not apparent (Fig. 4.127); case curved, typically of silk but may incorporate some sand (Fig. 4.122); springs and small cold streams of Appalachian Mountains of NY, VA, and GA, NC, SC, TN .................................................................................................... Micrasema burksi Ventral apotome broader than long (Fig. 4.121); head, case, and habitat variable ................. 39

38’ 39(38’) 39’

Head flat with pronounced carina (Figs. 4.128, 4.129); case of sand, curved, with posterior bevel (Fig. 4.123); large streams to cold rivers, typically at base of Podostemum and mosses rather than on plants, or on rocks with no vegetation in GA, NC, SC, TN ................................................................................................................ Micrasema rickeri Head rounded dorsally (Fig. 4.130); if case of sand, posterior bevel absent or inconspicuous (Fig. 4.124) ....................................................................................................................... 40

40(39’) 40’

Case of sand, curved (Fig. 4.124); head pale yellow to light brown, frontoclypeus with bold, regular muscle scar pattern (Figs. 4.131, 4.132); large streams and rivers ............. 41 Case of silk or plant matter, straight (Figs. 4.125, 4.126); frontoclypeus lacking pattern of bold, regular muscle scars (Figs. 4.133–4.136); habitat variable ..................................... 42

41(40)

Anterior frontoclypeus with 4 circular muscle scars and constriction of frontoclypeus with dark semicircular markings (Fig. 4.131); setae 2 and 3 on anterior margin of frontoclypeus flattened, much thicker than other head setae and having expanded apices (Fig. 4.131); at base of Podostemum mats in rivers of VA, WV, and GA, KY, NC, SC, TN .............................................................................................................. Micrasema bennetti Frontoclypeus lacking above markings (Fig. 4.132); setae 2 and 3 not conspicuously thicker than other head setae (Fig. 4.132); large streams to large rivers, at base of Podostemum or mosses, often with M. wataga, SK to NS and south to OK and all southeastern states except MS ............................................................................... Micrasema rusticum

41’ 42(40’) 42’

Head pattern with very irregular or mottled appearance and conspicuous white or yellow spot on posterior end of frontoclypeus (Fig. 4.133); case mostly of plant material (similar to Fig. 4.125); larger, often warmer streams and rivers, usually in Podostemum mats with M. rusticum, SK to NS and south to all southeastern states ............................................................................................................... Micrasema wataga Head pattern not irregular or mottled (Figs. 4.134–4.136); case of plant material or of silk only ........................................................................................................................................ 43

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Fig. 4.120

Figs. 4.121a

4.121b

4.121c

Figs. 4.122 4.123 4.124

4.125 4.126a 4.126b

Fig. 4.129 Fig. 4.127

Fig. 4.130

Fig. 4.128

Fig. 4.131

Fig. 4.134

Fig. 4.132

Fig. 4.135

Fig. 4.133

Fig. 4.136

Figures 4.120–4.136, Micrasema spp. (Brachycentridae). 4.120–4.121, ventral apotomes of heads, ventral: 4.120, M. burksi; 4.121, Micrasema 3 spp. 4.122–4.126, cases: 4.122, M. burksi; 4.123, M. rickeri; 4.124, M. bennetti; 4.125, M. charonis; 4.126, Micrasema 2 spp. 4.127–4.128, heads, dorsal: 4.127, M. burksi; 4.128, M. rickeri. 4.129–4.130, heads, left lateral: 4.129, M. rickeri; 4.130, M. bennetti. 4.131–4.136, heads, dorsal: 4.131, M. bennetti; 4.132, M. rusticum; 4.133, M. wataga; 4.134, M. florida; 4.135, M. charonis; 4.136, M. scotti. All figures from Chapin (1978).

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43(42’) 43’ 44(43’) 44’

Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Episterna of mesothorax and metathorax each with single seta (similar to Fig. 4.137); frontoclypeus without pale anterior markings (Fig. 4.139); case sometimes with short pieces of projecting vegetation (Fig. 4.142); streams and rivers of Coastal Plain of FL Panhandle and southern AL .................................................................... Micrasema florida Episterna of mesothorax and metathorax each with multiple setae (Fig. 4.138); frontoclypeus with or without pale markings anteriorly (Figs. 4.140, 4.141) ......................................... 44 Head light brown to dark brown, with distinct muscle scars laterally and anterolateral regions of frontoclypeus with pale areas (Fig. 4.140); case of plant material (Fig. 4.143); cold streams and tributaries of cold rivers, typically in long trailing mosses of genus Fontinalis; QC to NF and south to AL, KY, NC, SC, TN ..............................................................................................................Micrasema charonis Head dark brown, without distinct muscle scars and anterolateral regions of frontoclypeus sometimes with pale areas (Fig. 4.141); case typically of silk (Fig. 4.144); mosses of limestone springs and small streams in IN, VA, WV and AL, GA, KY, TN .................................................................................................................. Micrasema scotti

CALAMOCERATIDAE, key to genera and species 45(14)

Anterolateral corners of pronotum somewhat extended (Fig. 4.147), but much less than Fig. 4.146; gill filaments single (Fig. 4.149); case a hollowed twig (Fig. 4.145); shredders-detritivores, gougers; sprawlers in lotic-erosional and lotic-depositional habitats; see also Lloyd, 1915d, 1921 as Ganonema americanum; QC to ME and south to AL, FL, GA, KY, NC, SC, TN ........................................ Heteroplectron americanum 45’ Anterolateral corners of pronotum produced into prominent lobes (Fig. 4.146); gills with 2 or 3 branches (Fig. 4.148); case of 2 leaf pieces, dorsal piece overhanging ventral one; shredders-detritivores; sprawlers in lotic-depositional habitats; see also Wallace & Sherberger (1970); DE, NJ, VA, and all southeastern states ..... Anisocentropus pyraloides

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Fig. 4.138 Fig. 4.139

Fig. 4.140

Fig. 4.141 Fig. 4.142

Fig. 4.146

Fig. 4.147

Fig. 4.143

Fig. 4.148

Fig. 4.144

Fig. 4.145

Fig. 4.149

Figures 4.137–4.149, Brachycentridae and Calamoceratidae genn. spp. 4.137–4.138, Micrasema spp. (Brachycentridae), left episterna of meso- and metathoraces, left lateral: 4.137, M. florida; 4.138, M. scotti. 4.139–4.141, Micrasema spp. (Brachycentridae), heads, dorsal: 4.139, M. florida; 4.140, M. charonis; 4.141, M. scotti. 4.142–4.144, Micrasema spp. (Brachycentridae), cases: 4.142, M. florida; 4.143, M. charonis; 4.144, M. scotti. 4.145, Heteroplectron americanum (Calamoceratidae), case. 4.146–4.147, Calamoceratidae genn. spp., pronota, dorsal: 4.146, Anisocentropus pyraloides; 4.147, Heteroplectron americanum. 4.148–4.149, Calamoceratidae genn. spp., abdominal gills: 4.148, Anisocentropus pyraloides; 4.149, Heteroplectron americanum. 4.137–4.144 from Chapin (1978); 4.145–4.149 © R.W. Holzenthal 2006.

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DIPSEUDOPSIDAE Phylocentropus, key to species (from Sturkie & Morse, 1999)

Larvae of Phylocentropus auriceps (rare in mountains of GA, NC, SC, TN, and VA) and P. harrisi (rare in AL and TX) are unknown. 46(11) 46’

Head light yellowish brown, with posterior region of frontoclypeus darkened (Fig. 4.150); hind tibiae each with 2 stout spines and 3 long hairs in lateral view (Fig. 4.153); body length of mature larva 15–20 mm; ON, QC, ME and south to LA and all southeastern states ................................................................... Phylocentropus carolinus Head uniformly brown with light spots, posterior region of frontoclypeus not darkened (Fig. 4.151); hind tibiae each with 2 stout spines (Fig. 4.154) or 3 stout spines (Fig. 4.155); body length of mature larva variable ........................................................... 47

47(46’) 47’

Hind tibiae each with 2 stout spines (Fig. 4.154); body length no more than 15 mm; QC to NB and south to LA and all southeastern states .......................... Phylocentropus lucidus Hind tibiae each with 3 stout spines (Fig. 4.155); body length of mature larva 15–19 mm; MB to NB and south to TX and all southeastern states ............ Phylocentropus placidus

GLOSSOSOMATIDAE, key to genera and species 48(8) 48’



Mesonotum with 2 or 3 sclerites (Figs. 4.156, 4.157, sometimes same color as membrane and hard to see—look for shiny, slightly hardened surfaces); head with ventromesal margins of genae not thickened and posterior median ventral ecdysial line about 1.5 times as long as each anterior divergent branch (Figs. 4.159, 4.160); anal opening without dark, sclerotized line on each side ....................................................................... 49 Mesonotum without sclerites; head with ventromesal margins of genae thickened and posterior median ventral ecdysial line about as long as each anterior divergent branch (Fig. 4.158); anal opening with dark, sclerotized line on each side (Fig. 4.152); constructing portable case, dome-like dorsally, composed of stones, each with transverse band of finer sand ventrally; scrapers; clingers in lotic-erosional habitats ..................................................................................................................... Glossosoma, 53

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Fig. 4.153

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Fig. 4.151

Fig. 4.154

Fig. 4.156

Fig. 4.158

Fig. 4.152

Fig. 4.155

Fig. 4.157

Fig. 4.159

Fig. 4.160

Figures 4.150–4.160, Dipseudopsidae and Glossosomatidae genn. spp. 4.150–4.151, Phylocentropus spp. (Dipseudopsidae), heads, dorsal: 4.150, P. carolinus; 4.151, P. placidus. 4.152, Glossosoma nigrior (Glossosomatidae), abdominal segments IX–X; caudal. 4.153–4.155, Phylocentropus spp. (Dipseudopsidae), right hind legs, right lateral: 4.153, P. carolinus; 4.154, P. lucidus. 4.155, P. placidus. 4.156–4.157, Glossosomatidae genn. spp., mesonota, dorsal: 4.156, Agapetus sp.; 4.157, Padunia jeanae. 4.158–4.160, Glossosomatidae genn. spp., heads, ventral: 4.158, Glossosoma nigrior; 4.159, Agapetus sp.; 4.160, Padunia jeanae; 4.150–4.151, 4.153–4.155 redrawn from Sturkie & Morse (1999); 4.152, 4.156–4.160 © R.W. Holzenthal 2006.

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49(48) 49’

Mesonotum with 3 sclerites and sa1 setae widely separated (Fig. 4.161); ventral apotome of head slender, parallel-sided sclerite (Fig. 4.163) .......................................................... 50 Mesonotum with 2 sclerites and distance between sa1 setae about same as distance between sa1 and sa2 setae (Fig. 4.162); ventral apotome of head not as slender, broader laterally (Fig. 4.164); constructing portable case of stones, dome-like dorsally with relatively large stones laterally and with transverse band of finer sand ventrally (similar to Fig. 4.169); scrapers, collectors-gatherers; clingers in lotic-erosional habitats ......................................................................................................................... Agapetus, 52

50(49) 50’

Each tarsal claw apparently trifid, with 3 points subequal in length (Fig. 4.166); see also Flint (1962b); scapers; clingers in lotic-erosional habitats; VA and AL, GA, KY, NC, SC, TN ........................................................................................................ Padunia jeanae Each tarsal claw with normal single point, basal seta and basal process much smaller (Figs. 4.167, 4.168) ........................................................................................................... 51

51(50’) Basal seta of each tarsal claw long, thin or stout, and arising from side of stout, long or short basal process (Fig. 4.167), sometimes causing the claw to appear trifid (similar to Fig. 4.166); constructing portable case, dome-like dorsally, composed of stones, with some relatively large stones laterally, each with a transverse band of finer sand ventrally (Fig. 4.169); scrapers; clingers in lotic-erosional habitats .............. Protoptila, 55 51’ Basal seta of each tarsal claw short and stout, larger than basal process (Fig. 4.168); case constructed of uniformly small stones (Fig. 4.170); scrapers; clingers in lotic-erosional habitats; MD, ME, VA and NC ..................................................... Culoptila plummerensis

Agapetus, species 52(49’)



Larvae have been described for Agapetus diacanthus (Edwards, 1956; TN), A. illini (Ross, 1944; ID to IL south to AR and KY, TN), A. jocassee (Craft & Morse, 1997; GA, NC, SC, TN), A. minutus (Sibley, 1926; ON to MA south to KY, NC, TN), and “very likely” A. walkeri (Wiggins, 1996, according to Etnier et al., 2010; QC to NB south to AL, GA, NC, SC, TN). Larvae of 25 southeastern Agapetus species were described/ redescribed by Etnier et al. (2010) without illustrations or key. The genus is known from eastern Canada, most of USA including all southeastern states except FL.

Glossosoma, key to species (from Wymer & Morse, 2000)

53(48’) 53’

Pronotum with small black spot above each forecoxa (Fig. 4.165) ........................................ 54 Pronotum without small black spots above forecoxae; see also Lloyd (1921) as G. americanum; QC to ME and south doubtfully to NC? .................. Glossosoma lividum

54(53) 54’

Intersegmental fold between thorax and abdomen posterior to each metathoracic epimeron with small sclerite (Fig. 4.171, before arrow); see also Neubauer & Robertson (1985); MN to NF and south to AL, GA, KY, MS, NC, SC, TN ........................... Glossosoma nigrior Intersegmental fold between thorax and abdomen posterior to each metathoracic epimeron without small sclerite; BC to NF and south to KY ..................... Glossosoma intermedium

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Fig. 4.161

Fig. 4.162

Fig. 4.163

Fig. 4.164

Fig. 4.165

Fig. 4.166

Fig. 4.167

Fig. 4.168

Fig. 4.169

Fig. 4.170

Fig. 4.171

Figures 4.161–4.171, Glossosomatidae genn. spp. 4.161–4.162, mesonota, dorsal: 4.161, Padunia jeanae; 4.162, Agapetus sp. 4.163–4.164, heads, ventral: 4.163, Padunia jeanae; 4.164, Agapetus sp. 4.165, Glossosoma nigrior, pronotum and base of left forecoxa, left lateral. 4.166–4.168, left mesothoracic tarsal claws, left lateral: 4.166, Padunia jeanae; 4.167, Protoptila sp.; 4.168, Culoptila moselyi. 4.169–4.170, cases: 4.169, Protoptila sp.; 4.170, Culoptila moselyi. 4.171, Glossosoma nigrior, meso- and metathorax and abdominal segment I, left lateral. 4.165, 4.171 redrawn from Wymer & Morse (2000); all other figures © R.W. Holzenthal 2006.

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Protoptila, species 55(53)

The larva of Protoptila maculata (MB to QC south to TX and AL, NC, TN) was described by Ross (1944). Larvae of no other southeastern species of Protoptila have been described. The genus has been reported from BC to QC south to CA, AZ, NM, TX and all southeastern states except MS.

GOERIDAE, key to genera 56(13) 56’

Gills present, mostly 3-branched (Fig. 4.172), generally occurring on abdominal segments II–VII; case portable, tubular, made of small rocks, usually with 2 larger stones along each side (Fig. 4.173) ........................................................................................... Goera, 57 Gills absent; case portable, tapered, slightly curved, smooth, made only of small sand grains (Fig. 4.174) .............................................................................................. Goerita, 59

Goera, key to species (from Flint, 1960)

The larva of Goera townesi (AL, GA, NC, SC) is unknown. 57(56)



Metanotum with 3 pairs of sclerites (Fig. 4.175); spicules on central hump and anteriorly on pronotum conspicuously longer than those on remainder of pronotum; frontoclypeal setae 2 and 3 subequal in length (Fig. 4.177); sternal thoracic plates indistinct; see also Lloyd (1921); MN to NS and south to AL, GA, KY, NC, SC, TN .......... Goera calcarata Metanotum with 4 pairs of sclerites (Fig. 4.176); spicules on central hump and anteriorly on pronotum not conspicuously longer than those on remainder of pronotum; frontoclypeal setae 2 twice as long as setae 3 (Figs. 4.178, 4.179); sternal thoracic plates conspicuous ............................................................................................................ 58

58(57’) 58’

Front of head with central area smooth; posteriorly with sharp, high carina (Fig. 4.178); QC to NB and south to GA, KY, NC, SC, TN ................................................... Goera fuscula Front of head covered entirely with spicules; posterior ridge low and broad (Fig. 4.179); MN to NH and south to AL, KY, NC, TN ..................................................... Goera stylata

57’

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Fig. 4.172

Fig. 4.173

Fig. 4.175

Fig. 4.177

Fig. 4.174

Fig. 4.176

Fig. 4.178

Fig. 4.179

Figures 4.172–4.179, Goeridae genn. spp. 4.172, Goera fuscula, gill. 4.173–4.174, cases: 4.173, Goera fuscula; 4.174, Goerita semata. 4.175–4.176, Goera spp., metanota, dorsal: 4.175, G, calcarata; 4.176, G, fuscula. 4.177–4.179, Goera spp., heads, dorsal: 4.177, G, calcarata; 4.178, G, fuscula; 4.179, G, stylata. 4.172–4.174 © R.W. Holzenthal 2006; 4.175–4.179 redrawn from Flint (1960).

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Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species

Goerita, key to species

(from Parker, 1999; see also Wiggins, 1973) 59(56’)’ Pronotal median ridge evenly convex laterally (Fig. 4.180); OH, PA, VA, WV, and KY, NC, TN ........................................................................................................ Goerita betteni 59’ Pronotal median ridge with bulbous swellings posterolaterally (Figs. 4.181, 4.180) ............. 60 60(59) 60’

Pronotal median ridge with deep thumb-like impression posteriorly (Fig. 4.181); MT and mountains of NC, TN, VA ........................................................................... Goerita semata Pronotal median ridge without impression (Fig. 4.182); mountains of NC, TN ......................................................................................................................... Goerita flinti

HELICOPSYCHIDAE Helicopsyche, key to species (based on Floyd, 1995b)

61 (1) 61’

Head with pale muscle scars (Fig. 4.184); lateral sclerite of each anal proleg pale (Fig. 4.186), length of mature larva 7–8 mm (see also Lloyd, 1921; Elkins, 1936); AB to NF and south to CA, AZ, NM, TX and all southeastern states ..... Helicopsyche borealis Head lacking muscle scars (Fig. 4.185); lateral sclerite of each anal proleg dark (Fig. 4.187); length of mature larva