Neotropical Ceratopogonidae Diptera Insecta AquaticBiodiversity in Latin America

FOR MANY YEARS, STUDENTS FROM NUMEROUS COUNTRIES HAVE COMPLAINEDABOUT THE LACK OF “TOOLS” TO IDENTIFY AQUATIC INVERTEBRATES FROMLATIN AMERICA. KEYS FOUND IN ACCEPTED TEXTBOOKS ARE MOSTLY LIMITED,SUPERFICIAL OR TOO POPULAR-SCIENCE. ON THE OTHER HAND, MORE ADVAN-CED KEYS IN ACADEMIC BOOKS ON AQUATIC INVERTEBRATES FAIL TO COVERNEOTROPICAL REPRESENTATIVES IN DUE DETAIL. THE FEW THAT DO, HOWEVER,ALSO SHOW TOO MANY DEFECTS IN SCOPE AND PRESENTATION TO BE CONSID-ERED UP-TO-DATE.AN INTERNATIONAL TEAM OF EDITORS COMBINED THEIR EFFORTS WITH PENSOFT PUBLISHERS TO LAUNCH A NEW MAJOR SERIES ON THE AQUATIC BIODI-VERSITY IN LATIN AMERICA. ABOUT 15 MONOGRAPHS, WRITTEN BY SCIENTISTSFROM VARIOUS COUNTRIES, WILL OFFER A NEW, UNRIVALLED VIEW OF THE MAR-VELLOUS AQUATIC WORLD OF SOUTH AMERICA!THE SERIES IS ADDRESSED TO ZOOLOGISTS, ECOLOGISTS, HYDROBIOLOGISTS,BIOGEOGRAPHERS, CONSERVATIONISTS AND STUDENTS INTERESTED IN AQUATICBIODIVERSITY. THE SERIES WILL BE AN IMMINENT TOOL FOR ANY BIOLOGICALLIBRARY.

Volume 4This book brings together, for the first time, all known information on the 51 genera and 1095 described Neotropical species of biting midges. Anoverview of the group includes a discussion of bionomics, adaptations of thevarious life stages, habitats, their ecological roles (including disease transmis-sion), collecting and preparation techniques, zoogeography and classification. Akey to adults (including males) provides the first key to all the genera of thisregion. Limited keys to eggs and larvae are presented. Keys are written in bothEnglish and Spanish. A table of all previous descriptions of immatures and acatalog of all Neotropical species is included.

This book will be of interest to systematists, ecologists, medical entomolo-gists and students wishing to undertake innovative research in a remarkablyinteresting and instructive group of insects.

ABLA

Vol. 4

Neotropical Ceratopogonidae (D

iptera:Insecta)A

rt Borkent & G

ustavo R.Spinelli

Aquatic Biodiversity in Latin AmericaBiodiversidad Acuática en América Latina

ABLA Vol. 4 – Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae 1

Aquatic Biodiversity in Latin America

Biodiversidad Acuática en América Latina

Volume 4Neotropical Ceratopogonidae (Diptera: Insecta)

Volumen 4Neotropical Ceratopogonidae (Diptera: Insecta)

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

BIODIVERSIDAD ACUÁTICA EN AMÉRICA LATINA

Volumen 4

CERATOPOGONIDAENEOTROPICALES

(DIPTERA: INSECTA)por

Art Borkent & Gustavo R. Spinelli

Lector Científico a los Editores:William L. Grogan

Editores de la Serie:Joachim Adis, Jorge R. Arias,

Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow2007


AQUATIC BIODIVERSITY IN LATIN AMERICA

Volume 4

NEOTROPICALCERATOPOGONIDAE(DIPTERA: INSECTA)

byArt Borkent & Gustavo R. Spinelli

Scientific Reader to the Editors:William L. Grogan

Series Editors:Joachim Adis, Jorge R. Arias,

Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow2007


Front cover:

Female Forcipomyia (Microhelea) sp. feeding on phasmid antenna (Chlorosphasma sp.)in Costa Rica; Amazon River near Manaus

(photos: P. Naskrecki, WJ. Junk; design: Zheko Aleksiev & Elke Bustorf).

© PENSOFT Publishers

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system ortransmitted in any form by any means, electronic, mechanical, photocopying, recording or

otherwise, without the prior written permission of the copyright owner.

Aquatic Biodiversity in Latin America Vol. 4ISSN: 1312-7276

First published 2007ISBN-13: 978-954-642-301-6

Pensoft PublishersGeo Milev Str. 13a, Sofia 1111, Bulgaria

[email protected]

Printed in Bulgaria, June 2007



Series Editors/Editores de la Serie:Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

The book should be cited as follows:

Art Borkent & Gustavo R. Spinelli. 2007. Neotropical Ceratopogonidae (Diptera: Insecta).In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.):

Aquatic Biodiversity in Latin America (ABLA). Vol. 4. Pensoft, Sofia-Moscow, 198 pp.

VOLUMEN 4.CERATOPOGONIDAE NEOTROPICALES

(DIPTERA: INSECTA)por

Art Borkent & Gustavo R. SpinelliLector Científico a los Editores:

William L. Grogan

VOLUME 4.NEOTROPICAL CERATOPOGONIDAE

(DIPTERA: INSECTA)by

Art Borkent & Gustavo R. SpinelliScientific Reader to the Editors:

William L. Grogan


Habitus of female Culicoides variipennis COQUILLETT

(Nearctic Region; from McALPINE et al., 1981).




Series Editors: Joachim Adis, Jorge R. Arias,Guillermo Rueda-Delgado & Karl Matthias Wantzen

Biodiversity is a key word in science and global management schemes; however very fewpeople are able to identify the species and their ecology that make up “biodiversity”. Formany years, researchers and students from numerous countries complain about the lack of“tools” to identify aquatic invertebrates from Latin America. Keys found in acceptedentomological textbooks are mostly highly limited, superficial and rarely cover Neotropicalbiota in sufficient detail. On the other hand, specialized information on taxonomy or ecologyis scattered throughout the literature in many single publications.

An international team of editors have combined their efforts with Pensoft Publishersto launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Theirgoal was to find experts who combine the current state of knowledge in taxonomy andecology, in order to produce a concise and affordable handbook for each group. About 15separate monographs, written by reference scientists from various countries will offer anew, unrivalled view on the aquatic fauna of South America. Information on the ecologyand status of the taxa (written in English) is combined with illustrated identification keys tofamilies and genera, in both English and Spanish. The series is aimed at zoologists, ecologists,hydrobiologists, biogeographers, conservationists and students interested in aquaticbiodiversity. The series will be an essential tool for any biological library.

Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher.Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642-258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings andphotos (15 plates in color); publication date: March 2006.

Volume 2: Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri,Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto.Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-259-2, ISBN-13: 978-954-642-259-0, 165x240 mm, 646 pp., including 234 plates of figures in line drawings andphotos (16 plates in color); publication date: May 2006.


Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarón & Cecilia L.Coscarón Arias.Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-293-4, 165x240 mm, 685pp., including 135 plates of figures in line drawings and 19 maps of distribution;publication date: March 2007.

Volume 4: Neotropical Ceratopogonidae (Diptera: Insecta) by Art Borkent & GustavoR. Spinelli.Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-301-6, 165x240 mm, 198 pp.,including 26 plates of figures in line drawings and photos; publication date: June 2007.

Ceratopogonidae are found in virtually every habitat with even a small amount of sustainedmoisture, are notorious as severe pests of man and domestic animals, and are one of themost common fly families in traps. Yet they are by far the most poorly understood of allthe biting flies. In part, this is due to their remarkably diversity, both in numbers of speciesand in their various adaptations. Larvae range from being detritivores and scavengers topredators and adult females of various species feed on a wide array of vertebrates andinvertebrates. Some females draw blood from conspecific males while they are mating.Adults are also important pollinators of a wide array of plants.

This book brings together, for the first time, all known information on the 51 generaand 1095 described Neotropical species of biting midges. An overview of the group includesa discussion of bionomics, adaptations of the various life stages, habitats, their ecologicalroles (including disease transmission), collecting and preparation techniques, zoogeographyand classification. A key to adults (including males) provides the first key to all the generaof this region. Limited keys to eggs and larvae are presented. Keys are written in bothEnglish and Spanish. A table of all previous descriptions of immatures and a catalog of allNeotropical species is included.

This book will be of interest to systematists, ecologists, medical entomologists andstudents wishing to undertake innovative research in a remarkably interesting and instructivegroup of insects.


Authors:DR. ART BORKENT

691-8th Avenue SESalmon Arm, British Columbia V1E 2C2Canadae-mail: [email protected]

DR. GUSTAVO R. SPINELLI

Divisíon EntomologiaMuseo de La PlataPaseo Del Bosque s/n1900 La Plata, Argentinae-mail: [email protected]

Series Editors:PROF. DR. JOACHIM ADIS

Max-Planck-Institute for EvolutionaryBiologyTropical Ecology Working GroupPostfach 16524302 Plön, Germanye-mail: [email protected]

DR. JORGE R. ARIAS

5870 Colfax AvenueAlexandria, Virginia 22311, USAe-mail: [email protected]

ADDRESSES

M.Sc. GUILLERMO RUEDA-DELGADO

Grupo de Investigación en Cuencas yHumedales Tropicales GICHTUNIMAGADALENAUniversidad de Bogotá Jorge Tadeo LozanoLaboratorio de LimnologíaCarrera 4 No. 22-61Bogotá, D.C., Colombiae-mail: [email protected]

DR. KARL MATTHIAS WANTZEN

Institute of LimnologyUniversity of KonstanzPostfach M 65978457 Konstanz, Germanye-mail: [email protected]

Scientific Reader to the Editors:DR. WILLIAM L. GROGAN

Department of Biological SciencesSalisbury UniversitySalisbury, Maryland 21801-6837e-mail: [email protected]


FOREWORD

Neotropical Ceratopogonidae (Diptera: Insecta)by Art Borkent & Gustavo R. Spinelli

The Ceratopogonidae are a large and diverse family of small, mostly crepuscular fliescommonly known as biting midges. Though biting midge species are known as viciousbiters of humans and domestic animals in many areas, some of the most vivid descriptionsof these encounters have come from the Neotropical region. Even the intrepid Victoriannaturalist Alfred Russell Wallace, writing near the headwaters of the Rio Negro on theBrazil-Venezuelan border, found them to be intolerable:

“I was…exposed to the pest of the sand-flies, which, every afternoon, from fourto six, swarm in millions, causing by their bites on the face, ears and hands, themost painful irritation.”

The sheer numbers of biting midges in many areas continue to circumvent the advanceof agriculture and tourism in the Neotropics. In addition, some ceratopogonid species areknown to vector important viruses (e.g., Oropouche) and filarial nematodes (e.g., Dipetalonemaspp.) to humans. Though it comes as little consolation to those who have experienced anonslaught by biting midges, most species in the Neotropics feed exclusively on the bloodof other insects. Nevertheless, to their credit, some ceratopogonids (species in the genusForcipomyia) serve as an important cacao pollinator, making a direct contribution to theenjoyment of chocolate worldwide.

Although there have been many taxonomic papers published on NeotropicalCeratopogonidae from the nineteenth century to the present, this volume represents the firsttruly comprehensive work for the region because it includes all described taxa. The significanceof this work can not be overstated since approximately 20 percent of the World’s describedceratopogonid species are treated in its 198 pages. Both authors have collected extensively inLatin America and have authored or coauthored taxonomic and ecological papers aboutceratopogonid species from the region, many of which included descriptions of new species.I have had the distinct pleasure of meeting Art Borkent and Gustavo Spinelli. Both areprofessional systematists with an infectious enthusiasm for the study of ceratopogonids. Bothhave extensive contacts in many countries of Central and South America and have trainedparataxonomists to assist them (e.g., through the National Biodiversity Institute (INBio) inCosta Rica). Both are advocates of continued biodiversity studies in the Neotropics towardsthe conservation and sustainable use of the numerous species which inhabit the region.

In addition to their comprehensive, well-illustrated keys to the adults, eggs and larvaeof Neotropical genera, the authors have included important chapters on the bionomics of


species in this region, methods of collection and rearing ceratopogonids, phylogeneticrelationships among taxa including references to the fossil record and a preliminarydiscussion of the zoogeography of selected species. A synopsis of the medical importanceof ceratopogonids in Latin America is also included as are useful syntheses of informationabout Neotropical ceratopogonids (e.g., Table 1 includes references for all species whoseimmature stages have been described). The color photographs in the bionomics chapter ofbiting midges feeding on other insect species are particularly striking. Another useful featurefound in the Catalog of the Ceratopogonidae of the Neotropical Region is the designation ofspecies in the genera Culicoides and Leptoconops which are known to bite humans. Borkent andSpinelli acknowledge more than once that many more ceratopogonid species will be describedfrom Latin America following expeditions to poorly collected areas. Their foundational workwill most certainly facilitate future taxonomic work. The authors have provided an invaluableresource for scientists who investigate aquatic biodiversity in the Neotropics.

STEVE MURPHREEEditor, Ceratopogonidae Information ExchangeBelmont UniversityNashville, Tennessee, USA


PREFACE

The Ceratopogonidae are commonly known as biting midges, no-see-ums or punkies, inSpanish as manta blanca, polvorines, or jejenes (which also may refer to Simuliidae), and inPortuguese as maruim or mosquito pólvora. They have a bad reputation as being nastybiters that pester humans and domestic animals and, in some instances, transmit harmfuldiseases. Because of their small size, the females of some species can pass through screenand mesh that keeps other biting pests outside and these can make life insufferable. Bitingmidges may occur in such huge numbers that in some areas people are driven indoors (orcomplain loudly and then suffer numerous itchy bites!).

Few people, however, realize that this group of flies provides important services in awide array of ecosystems. Some species are important pollinators of such plants as cacao(without them we wouldn’t enjoy chocolate!) and rubber trees, and the larvae of many areimportant predators of other organisms in semiaquatic and aquatic habitats. The adults ofmost biting midges actually suck blood from other insects and may even be importantvectors of viruses that kill caterpillars.

The biting midges are a diverse group in both numbers of species and in their habitsand there are some intriguing stories associated with the way they obtain their food, mateand the kinds of different habitats they occupy. This contribution provides the basis forunderstanding the family in the Neotropical Region, allows for their identification to thegeneric level, gives a synopsis of each genus, details some aspects of their behaviour andecology, discusses various collecting and preparation techniques, and provides a catalog ofthe Neotropical species.

The first author thanks his wife and dearest friend Annette BORKENT for providingmoral and financial support throughout the writing of this contribution and for her steadfastfaith in his work. Pablo DELLAPÉ and Maria M. RONDEROS provide substantial advice andassistance in preparing the plates and we express our gratitude to them. A number of thefigures were skillfully produced by Ana BRENES (INBio) and we appreciate permissionfrom the Instituto Nacional de Biodiversidad in Costa Rica to reproduce these here. NelidaCALIGARIS redrew some of the illustrations of previously published figures. The SeriesEditors and the Scientific Reader are thanked for valuable suggestions, improving the contentand the final shape of the book. Many of the figures reproduced here are from Chapter 28of Volume 1 of the Manual of Nearctic Diptera (MCALPINE et al. (eds), Agriculture CanadaMonograph 27, Agriculture & Agri-Food Canada, 1981) and we are grateful for permissionto do so from the Minister of Public Works and Government Services Canada (2006).

Annia PICADO (INBio) has slide mounted tens of thousands of Ceratopogonidae fromCosta Rica with tremendous skill and this has helped our understanding of Neotropicalbiting midges immeasurably. Heron HUERTA provided a careful review of this work and wethank him for his detailed critique.


We gratefully acknowledge permission to use photographs from several colleagueswho are noted in the figure captions.

THE AUTHORS


1 BIONOMICS 15References to species of Neotropical Ceratopogonidae

described as eggs, larvae and/or pupae (Table 1) 25Number of named species with described immatures for

genera of Neotropical Ceratopogonidae (Table 2) 32Organisms transmitted by Neotropical

Ceratopogonidae (Table 3) 33

2 CAPTURE AND STUDY OF CERATOPOGONIDAE 34

3 REARING CERATOPOGONIDAE 38

4PHYLOGENETIC RELATIONSHIPS AND FOSSIL

RECORD OF CERATOPOGONIDAE 39

5 ZOOGEOGRAPHY 40

6 CLASSIFICATION 42Catalog of the Ceratopogonidae of the Neotropical

Region (Table 4) 43

7 IDENTIFICATION 100Numbers of named valid species in each genus of

Neotropical Ceratopogonidae (Table 5) 101Diagnosis of the Ceratopogonidae 102

Table of Contents


Key to the genera of adult Ceratopogonidae in theNeotropical Region 103

Key to the genera of eggs of Ceratopogonidae in theNeotropical Region 129

Key to the genera of larvae of Ceratopogonidae in theNeotropical Region 129

Clave para los géneros de adultos de Ceratopogonidaede la Región Neotropical 134

Clave para los huevos de géneros de Ceratopogonidaede la Región Neotropical 146

Clave para las larvas de géneros de Ceratopogonidaede la Región Neotropical 146

8SYNOPSIS OF THE CERATOPOGONIDAE OF THE

NEOTROPICAL REGION 148

9 REFERENCES 154

10 SUBJECT INDEX 182


1BIONOMICS

Ceratopogonidae are one of the common of fly families in the Neotropical Region (BROWN,2005). Adults may be found from the coast (some are intertidal) to the highest elevations. Wehave records of Forcipomyia MEIGEN and Dasyhelea KIEFFER from 3700 to 4000 meters innorthwestern Argentina and Forcipomyia and Atrichopogon KIEFFER from 3819 meters in CostaRica (which is that country’s highest point). They are present in nearly all habitats where thereis even a little long-lasting moisture. Adults of a few species are vicious biters of humans anddomestic animals and some act as vectors of various diseases or parasites (BORKENT, 2004).

There is little information on the specific life cycle of Ceratopogonidae in the Neotro-pical Region. Further to this, our survey of the literature (Tables 1, 2) indicates that fewerthan 15 percent of the species named from this area are described in even one immaturestage. Nineteen genera are entirely unknown. Some descriptions, especially those that areolder, are likely based on misidentified species and these immatures need to be restudied.Aware that many species remain to be described even as adults, there is a rich field availablefor investigation. Undoubtedly, many of the larvae of Ceratopogonidae play an importantecological role that is nearly entirely uninvestigated. Rearings of larvae will certainly provideimportant information regarding the structure of tropical ecosystems but clearly, the firststep is to have these described and to have keys made available. At the present time there areno reliable keys to the larvae or pupae to even the generic level although these stages arepresently being studied at a worldwide level (BORKENT, in prep.).

All members of the family have an egg, four larval instars, a pupal and an adult stage.Eggs may be laid on moist substrate or, in the case of fully aquatic species, in water. Thefact that adult ceratopogonids, as a group, are present throughout the year in tropical areassuggests that many species have multiple life cycles, without the third or fourth instarexperiencing a diapause as do nearly all species in northern temperate and arctic regions. Insome tropical areas, it appears that adult Ceratopogonidae may be somewhat more diverseduring the rainy season. It is unknown how tropical Ceratopogonidae respond in areas witha pronounced dry season. The life cycle of Ceratopogonidae in the southern temperatezone of South America is entirely unstudied but the presence of both numbers of speciesand adults in late spring in this area indicates that many species likely overwinter in the latelarval stage, as they do in the northern temperate region.

Larvae are important detritivores or predators in semiaquatic and aquatic systems ofall sizes, from tiny tree holes and the water held by tropical flowers, to large rivers and lakes,where they may be an important source of food for other invertebrates and fish. In tropicalareas of Latin America, the immatures of Ceratopogonidae form an important compo-


nent of arboreal aquatic systems (such as in bromeliads and tree holes). The most diversegroup, both in terms of numbers of taxa and structurally, are those in the subfamily Cer-atopogoninae. Members of this group have predaceous larvae which are like small snakes,working their way slowly through the wet substrate and, depending on the instar and/ortaxon, feeding on microorganisms or lashing out at prey. Some prey are swallowed whole(e.g. nematodes) but in the case of larger prey, which are mostly other insect larvae, thebiting midge larvae of some taxa burrow through the cuticle of the host and eat theirinsides. It seems certain that in many habitats, biting midge larvae play an important role inthe biological control of other insects, but this has not been studied in much detail.

The immatures of each of the four subfamilies differ substantially.The Leptoconopinae includes just one genus, Leptoconops SKUSE in the Neotropical

Region. The group is an ancient one and fossil Leptoconops are known from Lebanese am-ber, 121 million years old. Their larvae live in wet sand (especially on marine beaches) or incracked alkaline, clay soils. The sluggish larvae burrow through wet sand or loose clay soiland feed on microorganisms by raking their heads over the substrate.

Larvae of Forcipomyiinae are aquatic, semiaquatic or terrestrial. When they are inaquatic habitats, they are generally restricted to lentic (standing) waters although a few areknown from streams and small rivers and are associated with mosses and algae. They retainthe primitive anterior and posterior prolegs and therefore can walk on surfaces beneath orabove water. As they move, they graze the substrate surface for microorganisms, lookingsomewhat like cattle eating grass. Some species feed on decaying vegetation and a fewspecies may be found associated with manure. The larvae of most species have specializedsetae along the length of their bodies that secrete a sticky fluid that repels ants. When thelarvae pupate, most retain the larval exoskeleton on the posterior end of their bodies.

Larvae of Dasyheleinae are aquatic and semiaquatic and tend to be sluggish. Theyhave posterior hooks on their abdomens and move slowly over the substrate in which theyare found. Most species live in small water bodies such as those in epiphytes, tree holes,broken or damaged bamboo, rotten banana stems, fruit husks and small rock pools on themargins of streams and rivers. A few species are common in mangrove swamps and saltmarshes. Larvae may also be found in wet vegetation. One species in Costa Rica is a leaf-miner of the floating leaves of Salvinia and another is found in the wet refuse piles of someants (BORKENT, pers. obs.). Larvae feed on detritus, algae, and, in a few species, on carrion(dead insects). There are some species whose adults are associated with rocky areas in theupper tidal zone of coastal Costa Rica but the immatures of these are not known.

Larvae of the Ceratopogoninae, the most species-rich group of biting midges, are prog-nathous, lack prolegs and hooks (at least in instars 2-4), and their abdominal segments are notsecondarily divided. Larvae of most species swim in a characteristic, very rapid, serpentinemotion through the water column. When on or in substrate they assume a slower motion,working their way through wet substrate, eating microorganisms or lashing out at larger prey,looking very much like small snakes. Some smaller invertebrates are swallowed whole but inthe case of larger prey, which are mostly other insect larvae, the biting midge larva penetratesthe cuticle of the prey and devours the internal contents. The earlier phylogenetic lineagestend to feed on microorganisms and/or smaller invertebrates, often ingesting these whole.The more derived lineages, especially in the tribes Heteromyiini, Sphaeromiini, and Palpomyiini,


feed on larger invertebrates such as chironomid larvae and sometimes burrow into the bodyof their host. In many habitats, biting midge larvae probably exert an important biologicalcontrol on other insects. Larvae of the species-rich genus Culicoides LATREILLE may be foundin decaying vegetation, fruits, and manure, epiphytes, tree holes, wet mud, mangrove swampsand a large array of other aquatic habitats. In streams and rivers, Ceratopogoninae larvae maybe an important source of food for other invertebrates and fish. In general, the most primi-tive lineages of the subfamily are in smaller habitats, whereas those in the Heteromyiini,Sphaeromiini and Palpomyiini may be found in larger bodies of water: ponds, springs, andlarger lotic and lentic habitats such as rivers and lakes.

Pupae are generally sluggish and move their abdomens slowly in more or less circularmovements until they find an appropriate resting position in the midst of the substrate orat the water surface where their paired respiratory organs can obtain air. The pupae ofsome Sphaeromiini move above the water line and have specialized membranous pads onthe venter of the abdomens which they use to stick to the dry substrate. BORKENT & CRAIG

(2001) and CAZORLA & MARINO (2004) described unusual pupae of Stilobezzia KIEFFER fromCosta Rica and Peru respectively, which use their respiratory organs to obtain oxygen bypiercing the submerged leaves of aquatic plants.

Adult females of Leptoconops, Culicoides and Forcipomyia (only in the subgenus LasioheleaKIEFFER) are haematophagous on vertebrates, needing the blood to develop their eggs.Female Leptoconops and Culicoides (Fig. 1B, frontispiece) may occur in large numbers in plac-es where there is appropriate and abundant habitat (Table 4). Their bite (sometimes unno-ticed at the time) can produce burning and itching welts completely out of proportion totheir small size. In some areas of the Neotropical Region (northern South America, Cen-tral America and tropical Mexico) species of Leptoconops are generally restricted to marinebeaches and adult females are daytime biters, generally on the lower legs of humans. Theadults of some species (including L. bequaerti (KIEFFER) in Central America) have an inter-esting behaviour of resting under a thin layer of sand (LINLEY, 1968).

Adult Culicoides occur, in varying numbers, in virtually every region of the NeotropicalRegion, from Mexico to southern Chile (Puerto Natales, at almost 52º S), from the coast tohigh elevations (at least 3320 m in the Colombian Andes (WIRTH & LEE, 1967)). Althoughthe females of a few species may bite during the day and may occur in huge numbers, mostare crepuscular or nocturnal feeders.

Of the 266 species of Culicoides known from the Neotropical Region, 70 (26%) havebeen recorded feeding on humans (Table 4) but of these only the following occur in suffi-cient numbers to be considered serious pests of humans: Culicoides furens (POEY) and C. phle-botomus (WILLISTON) in coastal Central and northern South America, C. maruim LUTZ in coastalBrazil, C. insinuatus ORTÍZ & LEON, C. puracensis WIRTH & LEE at high elevations in the Andesof Colombia, C. pseudodiabolicus FOX and C. paraensis (GOELDI) in the Amazon basin, C. lahillei(ICHES) and C. debilipalpis LUTZ in Paraguay and northeastern Argentina, C. caridei (BRÈTHES) inUruguay and the province of Buenos Aires, Argentina, and C. patagoniensis RONDEROS & SPINELLI

in Argentinean and Chilean Patagonia. Further pestiferous species will likely be discovered inthe largely unknown Andes. Females of some other species feed on domestic animals. Forexample, females of C. debilipalpis feed on both humans and horses and C. arubae FOX &HOFFMAN, C. foxi ORTÍZ and C. barbosai WIRTH & BLANTON have been collected from mules or


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

horses. Culicoides insignis LUTZ is typically associated with cattle. Culicoides phlebotomus, a wellknown species in the Neotropics biting humans and dogs on marine beaches, is also known tosuck blood from leatherback turtles that are on the beach laying their eggs (BORKENT, 1996).The hosts of the remaining species of Culicoides are largely unknown.

The biting habits of female adult Forcipomyiinae, including only two genera, Forcipomyiaand Atrichopogon, are remarkably diverse. In general, female adults act as ectoparasites of largeinsects, with some sucking blood from their hosts for extended periods of time. In particular,those Forcipomyia in the ixodoides species group of the subgenus Microhelea KIEFFER attackphasmids (Fig. 2B) and katydids (Figs. 2A, 4A) and when replete and developing their eggs,look like small ticks (CLASTRIER & WIRTH, 1995). One female of this species group we exam-ined in Costa Rica had developed 2778 eggs. Other species of Forcipomyia suck hemolymph

A

B

Culicoides furensDet. G.R. Spinelli

2 km N. Montezeuma,Playa Grande, Costa Rica,21-XI-2000, A. BorkentBiting humansCD5099

‡


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

A

B


from caterpillars (Fig. 3B), sawfly larvae (WIRTH, 1972), spiders, crane flies, the wing veins ofdragonflies and damselflies (Fig. 3A), lacewings and butterflies (WIRTH, 1956a; WIRTH &MESSERSMITH, 1971; MARINO & VON ELLENRIEDER, 1999). The feeding habits of female Atri-chopogon are more poorly known but those of at least one subgenus feed on the blood of adultMeloidae beetles (BORKENT & ROCHA-FILHO, 2006). Species in the subgenus Forcipomyia (Lasio-helea) are the only Forcipomyiinae that are vertebrate feeders. Their hosts are poorly knownand females have been recorded feeding only on frogs and lizards in the New World (SPINELLI

et al., 2002). The males and females of many adult Forcipomyia and Atrichopogon feed on nectarto fuel their flight and are often common on flowers, particularly those that are small andwhite or green. One unusual example of consuming sugars is Forcipomyia adults from thebacks of pentastomid bugs in Costa Rica (Fig. 4B).

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

A

B

A


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

A

B

A


Adult females of Dasyhelea have reduced mouthparts and do not suck blood fromeither vertebrates or invertebrates. Some species are common at flowers where they, likemany other biting midges, obtain nectar from flowers.

Adult females in the subfamily Ceratopogoninae, other than those of Culicoides, arepredators and feed on other flying insects that are approximately the same size as them-selves. In general, a female flies into a swarm of male non-biting midges (Chironomidae),grabs a male, and either settles to the ground or surrounding vegetation. The female injectsan enzyme that dissolves the internal organs and tissues of the prey, sucking up the lique-fied contents as food. In some members of the Heteromyiini, Sphaeromiini and Palpomyi-ini the female enters a male swarm of her own species and, while mating occurs, pierces thebody of the male and sucks out its contents (DOWNES, 1978). Such females may later befound with the dried male genitalia still attached to their own abdomen, after the bulk ofthe dried male has broken off. The males of these species contribute not only their spermand body tissues but also an effective mating plug to the female with which they mate.

BORKENT (2004) provided a list of all diseases and parasites known to be transmittedby biting midges worldwide. The females of Leptoconops, Forcipomyia (Lasiohelea) and Culi-coides are the only genera which include species which bite vertebrates (from frogs to birdsto mammals) in the Neotropical Region. Worldwide, Ceratopogonidae are known to trans-mit 66 viruses, 15 species of protozoa and 26 species of filarial nematodes. It is highlylikely that many more remain undetected. This strongly suggests that Ceratopogonidae willbe found to transmit many more organisms in the Neotropical Region than the six virusesand eight nematodes currently known (Table 3). For example, it would be quite remarkableif there were no protozoan parasites of birds in the entire Neotropical Region when atleast nine species of protozoans are known from North America. The adults of thosebiting midges that suck hemolymph from large insects (some Atrichopogon and Forcipomyia)suggest the possibility that they may be involved in transmitting viruses from one insecthost to another, particularly among caterpillars.

Bluetongue is a viral disease of cattle, sheep, goats, and wild ruminants. It is particu-larly damaging in sheep, where up to half of infected animals may die. In cattle and goatsthe disease primarily affects the animal’s reproductive capabilities. Although it was knownthat the virus occurred in Central America (MO et al., 1994) and antibodies were detected incattle in the late 1980s in southern Brazil (CUNHA, 1990), there were no published data ofanimal illness in the Neotropics until recently. The Office International des Epizooties,World Health Organization (OIE, 2001) reported the presence of the disease in the stateof Paraná, Brazil, in which eight sheep and one goat showed classical symptoms, includingthe death of four of these animals, and 70 cattle showed susceptibility as well. More re-cently, the virus was isolated and characterized from asymptomatic bovines from north-eastern Argentina (GORCH et al., 2002). Bluetongue occurs virtually worldwide and hasdifferent, phylogenetically unrelated vectors in different regions of the globe, although allare in the genus Culicoides. In the Neotropical Region the primary vectors are likely C.insignis, C. pusillus LUTZ (WIRTH & DYCE, 1985; SÁENZ & GREINER, 1994; GORCH et al., 2002)and possibly C. filarifer HOFFMAN (SÁENZ & GREINER, 1994).

There are other diseases of humans and domestic animals that occur elsewhere in theNeotropical Region. One of the more significant is Oropouche fever, a viral infection pro-


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

A

B


ducing severe flu-like symptoms, mostly in Brazil. It has been responsible for at least 27epidemic outbreaks in humans, including one in the state of Pará, Brazil in 1978-1980 whichafflicted at least 130,000 people. The disease is primarily transmitted by species of Culicoides(and particularly by C. paraensis) but has also been isolated (at low levels) from some mosqui-toes. Outbreaks occur primarily in urban areas, where populations of C. paraensis may occur atgreat densities (HOCH et al., 1990; MELLOR et al., 2000). MERCER et al. (2003) reported decayingplatano (plantain: Musa parasidiaca) stems, stumps, flowers, fruits and debris beneath platanotrees, as well as soil beneath a fruiting mamey (Pouteria sapota) tree and organic-rich mud alonga lake shoreline, as larval substrates of C. paraensis in Iquitos, Peru, a city which has sufferedseveral outbreaks of Oropouche fever. LEDUC & PINHEIRO (1989) provide an excellent over-view of Oropouche fever. The disease has also been studied in Peru (CHAVES et al., 1992;WATTS et al.,1997; PINHEIRO et al.,1998) and Panama (PINHEIRO et al.,1998).

Ceratopogonids provide some important services in ecosystems, although often thoughtof in negative terms. The adults of many genera, primarily Forcipomyia and Dasyhelea, areimportant pollinators of some commercial trees such as cacao (Theobroma cacao) and rubber(Hevea brasiliensis). Species of these ceratopogonid genera, and those of Atrichopogon, Culi-coides, and Stilobezzia, are likely to be pollinators for a number of other plants in tropicalAmerica (possibly including mango (Mangifera indica) and a number of palms). Furtherinformation about pollination by ceratopogonids may be found in a series of papers byWARMKE (1951, 1952), SAUNDERS (1959), SAUNDERS & Bowman (1956), WINDER (1977a, b),WIRTH (1956c) and YOUNG (1982, 1983, 1986a, b). Although it is unpleasant for many, thevicious attacks on humans and domestic animals by the females of some species of Culi-coides serve to restrict human encroachment in some important and threatened habitats,helping to preserve these for the future.

Ceratopogonidae larvae, pupae and adults harbour a variety of their own parasitesand diseases, including viruses, bacteria, fungi, protozoa, nematodes, mites, and even para-sitic wasps (WIRTH, 1977; FRANA et al., 2001). Although nearly all reports are from else-where, VILLALOBOS & RONDEROS (2003) recently experimentally infected immatures stagesof Dasyhelea necrophila SPINELLI & RODRÍGUEZ from Argentina with larvae of Nematomor-pha, causing a high level of parasitization and death of the host.


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

Taxon Reference Stages Described

Leptoconops bequaerti (as Leptoconops sp.) PAINTER, 1927 L PMAYER, 1934 PLINLEY, 1965a ELINLEY, 1968 L

Atrichopogon bifidus EWEN & SAUNDERS, 1958 L PA. caribbeanus EWEN & SAUNDERS, 1958 L PA. fusculus MALLOCH, 1915a L P

THOMSEN, 1937 L P(as polydactylus) NIELSEN, 1951 L P(as polydactylus) EWEN & SAUNDERS, 1958 L P(as polydactylus) DOWNES & WIRTH, 1981 L

WIRTH, 1994a L PA. fusculus or rostratus

(as Kempia haesitans) LENZ, 1934 L PA. incultus EWEN & SAUNDERS, 1958 L PA. longicornis EWEN & SAUNDERS, 1958 L PA. obscurus EWEN & SAUNDERS, 1958 L PA. remigatus EWEN & SAUNDERS, 1958 L PA. saundersi EWEN & SAUNDERS, 1958 L PA. tuberculatus EWEN & SAUNDERS, 1958 L P

Forcipomyia (Caloforcipomyia) caerulea SAUNDERS, 1957 L PF. (C.) glauca (as monilis) KRIVOSHEINA, 1968 L PF. (C.) varicolor SAUNDERS, 1957 L P

Forcipomyia (Euprojoannisia) blantoni SORIA & BYSTRAK, 1975 L PBYSTRAK & WIRTH, 1978 L P

F. (E.) bromeliae SAUNDERS, 1957 L PF. (E.) calcarata WIRTH, 1952a L PF. (E.) falcifera SAUNDERS, 1957 L PF. (E.) longispina SAUNDERS, 1957 L PF. (E.) mortuifolii SAUNDERS, 1957 L P

BYSTRAK & WIRTH, 1978 L PF. (E.) setigera SAUNDERS, 1957 L PF. (E.) spatulifera SAUNDERS, 1957 L P

F. (Forcipomyia) argenteola LANE, 1947c L PF. (F.) genualis (as raleighi) SAUNDERS, 1957 L P

(as raleighi) GUTSEVICH & GLUKHOVA, 1970 L PF. (F.) harpegonata WIRTH & SORIA, 1975 L P


F. (F.) pictoni WIRTH, 1976 L PF. (F.) rioplatensis SPINELLI et al., 2005a L P

F. (Lasiohelea) attenuata SAUNDERS, 1964 L PF. (L.) cornuta SAUNDERS, 1964 L P

KRIVOSHEINA, 1968 LF. (L.) intermedia SAUNDERS, 1964 L PF. (L.) quasicornuta SAUNDERS, 1964 L PF. (L.) stylifer LANE, 1947c L P

F. (Lepidohelea) brasiliensis WIRTH & SPINELLI, 1993a PF. (L.) edmistoni WIRTH & SPINELLI, 1993b L PF. (L.) herediae WIRTH & SPINELLI, 1993a L PF. (L.) luteigenua WIRTH & SPINELLI, 1992a L PF. (L.) seminole WIRTH & SPINELLI, 1992a L P

F. (Metaforcipomyia) cerifera SAUNDERS, 1957 L P

F. (Microhelea) fuliginosa(as inornatipennis spp.ornaticrus) LANE, 1947c L P

F. (Phytohelea) antiguensis SAUNDERS, 1957 L PF. (P.) bromelicola (as a Apelma) SAUNDERS, 1925 L PF. (P.) caribbeana SAUNDERS, 1957 L PF. (P.) dominicana DE MEILLON & WIRTH, 1979 L PF. (P.) edwardsi (as a Apelma) SAUNDERS, 1925 L PF. (P.) jocosa SAUNDERS, 1957 L PF. (P.) keilini (as a Apelma) SAUNDERS, 1925 L PF. (P.) magna (as a Apelma) SAUNDERS, 1925 L PF. (P.) musae CLASTRIER & DELÉCOLLE, 1994 PF. (P.) oligarthra SAUNDERS, 1957 L P

TOKUNAGA, 1961 L P

F. (Thyridomyia) nodosa SAUNDERS, 1957 L P

F. (Trichohelea) trinidadensis SAUNDERS, 1964 P

F. (Warmkea) aeria SAUNDERS, 1957 L PWIRTH & SORIA, 1980 L P

F. (W.) lesliei (as bicolor) SAUNDERS, 1957 L PWIRTH & SORIA, 1980 L P

F. (W.) spinosa SAUNDERS, 1957 L PWIRTH & SORIA, 1980 L P

F. (W.) terrestris SAUNDERS, 1964 L PWIRTH & SORIA, 1980 L P

Table 1. Continued.



F. (W.) tuberculata SAUNDERS, 1957 L PWIRTH & SORIA, 1980 L P

F. sp. (as an Apelma sp.) SAUNDERS, 1925 L PF. sp. (as species A) SAUNDERS, 1957 LF. sp. (as species A) SAUNDERS, 1964 LF. sp. (as species B) SAUNDERS, 1964 L

Dasyhelea atlantis WAUGH & WIRTH, 1976 PBORKENT, 1991 P

D. bahamensis RONDEROS et al., 2003 L PD. calvescens WILLIAMS, 1944 E L PD. cincta (as penthesileae) SPINELLI, 1983b PD. correntina RONDEROS et al., 2004a PD. filibranchia (as a Ceratopogon) LUTZ, 1914 PD. flavicauda SPINELLI & RONDEROS, 1987 PD. lacustris SPINELLI & RONDEROS, 1987 PD. mediomunda MINAYA, 1978 PD. mutabilis THOMSEN, 1937 L P

WIRTH, 1952a PWAUGH & WIRTH, 1976 P

D. necrophila SPINELLI & RODRÍGUEZ, 1999 L PRONDEROS et al., 2003 L PRONDEROS et al., 2006 E L

D. paracincta BORKENT, 1991 L PD. paulistana FORATTINI & RABELLO, 1957 PD. pusilla FORATTINI & RABELLO, 1957 P

Culicoides annettae SPINELLI & BORKENT, 2004a PC. annuliductus VITALE et al., 1981 PC. arubae (as sp. 1) FOX, 1942 P

FORATTINI, 1957 PMURPHREE & MULLEN, 1991 L

C. bambusicola LUTZ, 1913 L(as sp. 2) FOX, 1942 P

LANE, 1947c L PBARBOSA, 1952, 1953 L PFORATTINI, 1957 PRONDEROS & SPINELLI, 2000 L PRONDEROS et al., 2000 L P

C. barbosai LINLEY, 1965b PLINLEY & DAVIES, 1971 EBLANTON & WIRTH, 1979 PMURPHREE & MULLEN, 1991 L

C. bayano VITALE et al., 1981 PC. bermudensis JAMNBACK, 1965 L P

Table 1. Continued.



BLANTON & WIRTH, 1979 L PMURPHREE & MULLEN, 1991 L

C. borinqueni LINLEY, 1965b PC. cancer HOGUE & WIRTH, 1968 L PC. charruus SPINELLI et al., 1993 PC. chaverrii SPINELLI & BORKENT, 2004a L PC. cochisensis MURPHREE & MULLEN, 1991 LC. crepuscularis THOMSEN, 1937 P

FOX, 1942 PJAMNBACK, 1965 L PBLANTON & WIRTH, 1979 L PMURPHREE & MULLEN, 1991 L

C. debilipalpis BARBOSA, 1952, 1953 PFORATTINI, 1957 P

C. dicrourus WIRTH & SORIA, 1981 L PC. edeni (as “possibly haematopotus”) JONES, 1961 P

BLANTON & WIRTH, 1979 PC. fililductus VITALE et al., 1981 PC. floridensis LINLEY, 1970 P

BLANTON & WIRTH, 1979 PC. furens PAINTER, 1927 L P

(as dovei) DOVE et al., 1932 E L PFOX, 1942 PWIRTH, 1952d L PFORATTINI, 1957 L PJONES, 1961 PLINLEY & KETTLE, 1964 L PJAMNBACK, 1965 L PLINLEY & DAVIES, 1971 EBLANTON & WIRTH, 1979 L PHRIBAR & MULLEN, 1991 LMURPHREE & MULLEN, 1991 L

C. guyanensis FORATTINI & RABELLO, 1956 L PFORATTINI, 1957 L P

C. haematopotus THOMSEN, 1937 PJONES, 1961 PJAMNBACK, 1965 L PATCHLEY & WIRTH, 1979 L PBLANTON & WIRTH, 1979 L PHRIBAR & MULLEN, 1991 LMURPHREE & MULLEN, 1991 L

C. hoffmani LINLEY & KETTLE, 1964 L PC. hylas (as sp. 3) FOX, 1942 P

FORATTINI, 1957 P

Table 1. Continued.



C. insignis FORATTINI et al., 1956 L PFORATTINI, 1957 L PLINLEY, 1965b PBLANTON & WIRTH, 1979 PSPINELLI et al., 1993 P

C. lahillei LAMBERSON et al., 1992 PC. luteovenus WIRTH, 1952a L PC. macieli WIRTH & SORIA, 1981 L PC. maruim LUTZ, 1913 E P

BARBOSA, 1952, 1953 PFORATTINI, 1957 PSPINELLI et al., 1993 P

C. melleus WIRTH, 1952d L PJAMNBACK et al., 1958 L PJONES, 1961 PJAMNBACK, 1965 L PBLANTON & WIRTH, 1979 L PHRIBAR & MULLEN, 1991 LMURPHREE & MULLEN, 1991 L

C. neopulicaris JONES, 1961 PC. oklahomensis MURPHREE & MULLEN, 1991 L

LAMBERSON et al., 1992 PC. paraensis MURPHREE & MULLEN, 1991 L

LAMBERSON et al., 1992 PC. phlebotomus PAINTER, 1927 L P

FOX, 1942 PFORATTINI, 1957 P

C. reticulatus LUTZ, 1913 LFORATTINI et al., 1960 L P

C. venezuelensis RONDEROS & SPINELLI, 1989 PDÍAZ et al., 2005 E L P

C. sp. PAINTER, 1927 L P

Alluaudomyia bella (as splendida) THOMSEN, 1937 L PWIRTH & GROGAN, 1981 L P

A. biestroi SPINELLI, 1988 PA. caribbeana SPINELLI & WIRTH, 1984c PA. distispinulosa SPINELLI, 1997b PA. schnacki SPINELLI, 1983b L P

Baeodasymyia christopheri BORKENT & CRAIG, 1999 E L PB. michaeli BORKENT & CRAIG, 1999 L P

Echinohelea lanei WIRTH, 1994b P

Table 1. Continued.



Parabezzia alexanderi GROGAN & WIRTH, 1977 PWIRTH & GROGAN, 1981 PSPINELLI & GROGAN, 1987 P

P. balseiroi SPINELLI & GROGAN, 1987 P

Schizonyxhelea forattinii BORKENT, 2000b L P

Stilobezzia antennalis THOMSEN, 1937 L PWIRTH & GROGAN, 1981 L PHRIBAR & MULLEN, 1991 L

S. bicolor FORATTINI & RABELLO, 1956 PS. chaconi LANE & FORATTINI, 1961 PS. coquilletti LANE & FORATTINI, 1961 P

WIRTH & GROGAN, 1981 PS. dorsofasciata (as a Palpomyia) LUTZ, 1914 ES. dubitans LANE et al., 1955 PS. fiebrigi FORATTINI & RABELLO, 1956 L P

CAZORLA et al., 2006 PS. glauca LANE et al., 1955 P

WIRTH & GROGAN, 1981 PS. panamensis FORATTINI et al., 1960 PS. rabelloi BORKENT & CRAIG, 2001 PS. wygodzinskyi LANE et al., 1955 P

Clinohelea horacioi SPINELLI & DURET, 1993 PC. nigripes SPINELLI & DURET, 1993 P

Heteromyia clavata WIRTH & GROGAN, 1977 PH. wokei WIRTH & GROGAN, 1977 P

Mallochohelea termophila SPINELLI, 1984 P

Neobezzia amnicola MAYER, 1959 P

Nilobezzia schwarzii WIRTH, 1962 PPALCHICK, 1981 P

Amerohelea sordidipes LANE et al., 1955 P

Bezzia bivittata HRIBAR & MULLEN, 1991 LB. blantoni SPINELLI & WIRTH, 1989b PB. brevicornis SPINELLI, 1983c P

SPINELLI & WIRTH, 1989b PB. bromeliae SPINELLI & WIRTH, 1989b PB. glabra THOMSEN, 1937 E L P

Table 1. Continued.



WIRTH, 1952a L PCHAN & LEROUX, 1967 EPALCHICK, 1981 PWIRTH, 1983a L P

B. nobilis (as setulosa) JOHANNSEN, 1905 L P(as setulosa) MALLOCH, 1915a L P(as chrysocoma) THIENEMANN, 1928 L

MAYER, 1934 L P(as chrysocoma) MAYER, 1934 P(as setulosa) LENZ, 1934 P(as chrysocoma) LENZ, 1934 P(as setulosa) THOMSEN, 1937 L P

GLUKHOVA, 1979 L(as setulosa) PALCHICK, 1981 P

WIRTH, 1983b L P(as acanthodes) SPINELLI, 1983c P

KNAUSENBERGER, 1987 LHRIBAR & MULLEN, 1991 LHRIBAR, 1993 L

B. pulchripes MAYER, 1959 PSPINELLI & WIRTH, 1989b P

B. roldani SPINELLI & WIRTH, 1981, 1989b L PB. snowi SPINELLI & WIRTH, 1991 P

Clastrieromyia dycei SPINELLI & GROGAN, 1986 P

Pachyhelea pachymera SPINELLI, 1983c P

Palpomyia guarani RONDEROS et al., 2004b L PP. lacustris LANE et al., 1955 PP. subaspera (as essigi) WIRTH, 1952a P

GROGAN & WIRTH, 1979c PP. wirthi LANE et al., 1955 P

Paryphoconus angustipennis MAYER, 1952 EP. flavidus (as lanei) MAYER, 1959 PP. mayeri MAYER, 1959 PP. oliveirai WIRTH & RATANAWORABHAN, 1972b P

Stenoxenus johnsoni WIRTH & RATANAWORABHAN, 1972b E

Table 1. Continued.



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

Genus Egg Larva Pupa Total # Percentage ofSpecies Neotropical fauna

Leptoconops 1 1 1 1 8.3Forcipomyia 0 42 45 45 21.3Atrichopogon 0 10 10 10 10.0Dasyhelea 2 5 14 14 24.6Culicoides 4 22 34 35 13.2Paradasyhelea 0 0 0 0 0Allohelea 0 0 0 0 0Alluaudomyia 0 2 5 5 26.3Austrohelea 0 0 0 0 0Baeodasymyia 1** 2 2 2 40.0Brachypogon 0 0 0 0 0Echinohelea 0 0 1 1 9.1Monohelea 0 0 0 0 0Parabezzia 0 0 2 2 8.3Schizonyxhelea 0 1 1 1 50.0Stilobezzia 1 2 10 11 17.2Clinohelea 0 0 2 2 14.3Heteromyia 0 0 2 2 18.2Pellucidomyia 0 0 0 0 0Johannsenomyia 0 0* 0 0 0Mallochohelea 0 0 1 1 16.7Neobezzia 0 0 1 1 12.5Nilobezzia 0 0 1 1 33.3Amerohelea 0 0 1 1 9.1Bezzia 1 4 8 9 19.6Clastrieromyia 0 0 1 1 25.0Pachyhelea 0 0 1 1 50.0Palpomyia 0 1 4 4 8.7Phaenobezzia 0** 0 0 0 0Paryphoconus 1** 0 3 4 10.0Stenoxenus 1** 0 0 1 6.3

Total 12 92 150 155 14.2

* The first instars of Johannsenomyia are superficially described by JOHANNSEN 1905: 108 and THOMSEN

1937: 73.** Eggs extracted from or observed within adult.


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

Taxon Vector Vertebrate Hosts Location

VIRUSESBunyavirusAnanindeua C. paraensis Birds, marsupials, Brazil

rodents, primates

Oropouche C. paraensis Sloths, humans; antibodies Trinidad, Panama,in other mammals Brazil, Peru

Utinga Ceratopogonidae Sloths; antibodies in South and Centralother mammals America

OrbivirusBluetongue C. insignis,C. filarifer, Wild and domestic New World, Southern

C. pusillus ruminants Europe, Africa, Asia,Australia

VesiculovirusVesicular stomatitis – C. spp. Horses, cattle, pigs, humans New WorldNew Jersey

Unclassified ArbovirusesBuritirana C. spp. Unknown Brazil

FILARIOIDEADipetalonema caudispina C. hollensis (in lab) Monkeys (Cebidae) South America

Dipetalonema gracile C. hollensis (in lab) Monkeys Mexico, Central and(Cebidae, Callithricidae) South America

Dipetalonema marmosetae C. furens Monkeys South America(Cebidae, Callithricidae)

Dipetalonema ozzardi C. lahillei, C. paraensis Humans, other primates? Central and SouthL. bequaerti, C. furens AmericaC. phlebotomus

Dipetalonema perstans C. spp. Humans, other primates Central and SouthAmerica, Africa

Icosiella neglecta F. spp. Frogs South America,Europe, Asia, Africa

Onchocerca cervicalis C. spp. Equidae Worldwide

Onchocerca gutturosa C. spp. Bovidae Worldwide


2CAPTURE AND STUDY OF

CERATOPOGONIDAEAdult biting midges may be collected by any one of the following techniques, with varyingresults.

1. Light trap. Although many specialized light traps exist that work very well, a goodsample may even be obtained at a lit window pane when there is good surrounding habitat.A simple and inexpensive technique is to place a funnel, approximately 20 cm in diameterabove a jar of alcohol, close to a source of light. Another productive method is to lay a UVlight on metal mesh (1 cm square, to keep out larger insects) which is lain over a shallowpan with soapy water; the water must be screened with fine mesh in the morning and thespecimens preserved in alcohol. Ultraviolet light works well to attract adult midges butregular florescent or incandescent light may also bring in large numbers. An advantage ofthis technique is that it collects both males and females of a given species so that they canbe associated.

2. CO2 trap. These traps collect females of those species that feed on vertebrate hosts(generally only those of species of Culicoides). A serious limitation of this technique is thatmales are not collected and therefore cannot be studied.

3. Sweeping vegetation, especially in areas with wet habitats, and aspirating the adultsfrom the net. This provides a good method to sample diverse habitats and often results inspecies which are not discovered otherwise. This method also results in samples of bothmales and females.

4. Malaise traps. These work very well to sample males and females of many species.Larvae and pupae are obviously restricted to specific habitats and may be sampled

directly from those aquatic, semiaquatic, or moist terrestrial habitats where they are found.The variety of habitats in which biting midge larvae and pupae may be found is describedabove. Sampling methods include the following:

1. Careful examination of wet substrate with a large magnifying glass or under a dis-secting microscope.

2. Samples of substrate from aquatic habitats (e.g. mud, silt, detritus, leaves) placed in apan with additional water. Most Ceratopogoninae larvae swim with a rapid serpentine motionand these may be removed with an eyedropper. Pupae almost always float at the water sur-face. Detritus removed from the margins of ponds, streams and lakes and placed in a whitebucket allows for manual removal of pupae as they float. Some workers add salt or sugar tofloat larvae and pupae from sample substrate and this is the only practical way to collect thosefew species in which the larvae do not swim or the pupae do not float at the surface.


3. Berlese funnels are a good method for extracting larvae from semiaquatic or merelymoist material (e.g. mosses growing on trees in cloud forest).

Once specimens are preserved in 70% alcohol (or some other preservative), mostneed to be placed on microscope slides (Fig. 1A) to be identified beyond the generic level.Methods vary but the following gives excellent results for slide mounting adult specimensstored in ethyl alcohol and which are not too old (3-4 years or, if stored under dark andvery cold conditions (below 0 °C), over 10 years). This method works particularly well formaterial to be used for taxonomic studies. A more rapid method is given further below.

Prepare for slide mounting by putting the following solutions into a series of fourstender dishes: 15% acetic acid, 2-propanol, 2-propanol layered over clove oil, clove oil(each dish about 3/4 full). Each specimen needs to go through this sequence of solutions.Preparation of five specimens at a time is convenient and manageable, so that 20 dishes arein use at a time. Placing all the dishes on a thin board so that they can be moved around asa group on a table top makes it easier to look at specimens while they are in the solutionsand still put all the dishes out of harm’s way while they are soaking.

A series of slides needs to be ready, by cleaning these well with a cloth or paper toweland placing a label on each slide. These slides should be placed in a slide tray. In making thefollowing preparations, round, 10 mm in diameter coverslips work very well.

1. Place the specimen into a dish of ethyl alcohol under the microscope. Remove thewings by grabbing the very base of the wing with fine forceps. Place both wings in the dishwith 15% acetic acid.

2. For most specimens, separate head and abdomen from thorax. For very small spec-imens, just tear the membrane between the head and thorax and between the abdomen andthorax so that all the parts remain attached to each other. Place parts, or torn specimen,into a three dram vial filled about 1/4 to 1/3 full with 8% KOH. Prepare a total of fivespecimens in this way. Each vial needs to be numbered with a grease pencil so that it can beassociated with the wings that are in the acetic acid.

3. Place vials of KOH into a beaker partially filled with water. Place beaker on hotplate and heat water to boiling point. Specimens will clear after 2-5 minutes, depending onthe size of the specimens. It is important that specimens of a similar size are “cooked” atthe same time so that they all clear at the same rate.

4. The following step requires some speed because specimens are damaged by re-maining in hot KOH for too long. Remove the specimen from the vial into a dish withsome 8% KOH in it and put under the microscope to see if it is properly cleared. Themuscles should have dissolved and freely flow out of the specimen if squeezed very, verygently with the forceps. If the specimens are over-cleared they will make for a poor micro-scope preparation. If cleared sufficiently, place specimen in acetic acid along with the asso-ciated wings. Leave for 15 minutes (or longer if necessary).

5. Move specimen to 2-propanol for 15 minutes (or longer if necessary).6. Move specimen to 2-propanol layered over clove oil. Leave until parts have sunk to

the bottom of the dish which generally takes about 20-60 minutes, depending on the sizeof the specimen (specimens can be left longer if necessary). After a day or two, the 2-propanol will mix with the clove oil and some more 2-propanol should be added so thatthere is always a distinct layer between the 2-propanol and the clove oil. If the clove oil has


too much 2-propanol mixed into it, leave the dish without a lid for a few hours in a wellventilated area and the 2-propanol will evaporate from the mixture.

7. Move specimen to pure clove oil and leave for at least 30 minutes (or longer ifnecessary).

8. Place small drop of Canada Balsam on the microscope slide; place the wings in thedrop and cover with a clean coverslip. If Canada Balsam has already hardened too much tomove the wings around easily, add a small drop of xylene, which will make the CanadaBalsam more fluid. Use xylene for the following procedures as well; it is always neededbecause Canada Balsam dries faster than the time it takes for a good slide to be made.

9. Place three small drops of Canada Balsam on the microscope slide and put thehead, thorax and abdomen into each. If the head, thorax and abdomen are still partiallyattached (small specimens), put specimen into the drop which will contain just the thoraxand remove the head and abdomen; put the head and abdomen into their proper places.

10. Remove the right legs and place all these under a separate coverslip.11. Arrange the thorax so that the left side is facing upward. Orient the head so that

the front of the head is oriented upward and the antennae are splayed outward. Do not adda coverslip.

12. Details of the genitalia are often very important in species identification, so thatthe following step is a very important one. Arrange the abdomen so that the ventral part ofthe genitalia is facing upward. In some males, the genitalia may be only partially turned inrelation to the rest of the abdomen and in this case, sometimes the genitalia will need to beremoved from the abdomen (by tearing with the forceps and a pin) so that the separatedgenitalia lies with the ventral side pointed up. The rest of the abdomen should be straight,with the tergites dorsal, if possible. Do not add a coverslip.

13. Put the slide into the drying oven at 40-50 °C. After 1-2 days, the Canada Balsamwith the head, abdomen and thorax will have dried somewhat. Add some more CanadaBalsam to each of these and very gently put a coverslip over each, so they are not crushed.The partially dried Canada Balsam should keep these parts from being damaged. While theslide is drying, some of the Canada Balsam may leave a small air bubble under the coverslip.If so, add a tiny bit of Canada Balsam. Don’t worry if the added Canada Balsam traps abubble under the coverslip; the bubble nearly always moves out by itself (after some time).

14. Put the slide back into the drying oven. Check for air bubbles for the next severaldays and add Canada Balsam as needed. Remove the slide after 1-2 weeks and put into aslide box. Keep the slide box upright, so the slides are still horizontal.

Larvae and pupae may be treated with the same solutions and should be mounteddorsal side up. Larvae require no dissection but the operculum of the pupa as well as oneof the respiratory organs should be removed and placed under a separate coverslip (leavingthe Canada Balsam to dry for a day or two and then adding the coverslips). If the larva orpupa is represented by an exuvia, the specimens obviously do not require clearing and maybe placed directly into 2-propanol and processed from there.

A simpler method for mounting specimens, which is particularly useful for mass mount-ing material which is likely to include previously described species, is as follows: removespecimens from 70% ethanol (many can be done as one lot), very briefly blot excess alco-hol, transfer to 100% ethanol saturated with phenol, leave overnight or longer, and mount


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


3REARING CERATOPOGONIDAE

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


4PHYLOGENETIC RELATIONSHIPS AND

FOSSIL RECORD OF CERATOPOGONIDAEThe phylogenetic and fossil record of the Ceratopogonidae is one of the best understoodamong insects and provides a good example of combining cladistic analysis of extant taxawith interpretation of a detailed fossil record (BORKENT, 2000a; BORKENT & CRAIG, 2004).

The biting midges are an exemplary group for studying evolutionary relationshipsbecause of three outstanding features. First, they are diverse in numbers of species. Gen-erally, when a group is species-rich, there are intermediate species or groups present whichprovide good clues on the direction of evolutionary change of a given structure. Second,the biting midges have three very different stages (larva, pupa, adult) each of which havediversified structurally. Unlike many other groups, the immatures of biting midges haveevolved in many different morphological directions. Therefore, it is possible to compareconclusions about relationships based on one stage against the conclusions based on adifferent stage. Third, the biting midges have an incredible fossil record in amber that hasbeen investigated in some detail. We now have 11 major deposits ranging from 17 to 121million years of age, with thousands of specimens studied. The fossil biting midges arelargely in excellent condition and most taxonomic features can be studied in detail. Thesefossils provide an excellent gauge as to the accuracy of our interpretation of evolutionaryrelationships based on living species.

General conclusions show that the Leptoconopinae are the sister group of all remain-ing Ceratopogonidae, the Forcipomyiinae and Dasyheleinae are sister groups, and that thepreceding subfamilies form the sister group of the Ceratopogoninae. Within the Cerato-pogoninae, the Culicoidini are the sister group of the remaining taxa (BORKENT & CRAIG,2004). Significant research remains to be completed to further interpret the relationshipsbetween most of the remaining genera of the Ceratopogoninae.


5ZOOGEOGRAPHY

Our understanding of the zoogeography of Ceratopogonidae is rather limited. This is fortwo important reasons. One is that the phylogenetic relationships between many taxa areunknown and it is therefore impossible to interpret the significance of their distributions inlight of those relationships. Secondly, in the Neotropical Region, large areas have not beensampled, including much of the Amazon Basin and nearly all of the Andes. Without doubt,there remain a great number of species to be discovered.

Nevertheless, there are some broad geographic patterns which are apparent. Mostnotable is the distinctive fauna of Patagonia. The following genera are endemic to the area,with most of these present in forested habitats: Borkenthelea SPINELLI & GROGAN (threeforest species), Diaphanobezzia INGRAM & MACFIE (four species, three from forest, one fromsteppe), Isthmohelea INGRAM & MACFIE (monotypic, from forest), Notiohelea GROGAN & WIRTH

(two forest species), Physohelea GROGAN & WIRTH (two forest species), AustrosphaeromiasSPINELLI (two forest species). In addition, the following are restricted within the Neotropi-cal Region to Patagonia but are also present in other, mostly southern hemisphere areas:Austrohelea WIRTH & GROGAN (four species (three undescribed) from Patagonian forest; sixothers from Australia or New Zealand), Macrurohelea INGRAM & MACFIE (13 species (twoundescribed) from Patagonian forest, except one species that extends northward to aridzones of Chile and Argentina, one undescribed species in the Somuncura Plateau in thePatagonian steppe; three described and two unnamed from Australia), and ParadasyheleaMACFIE (three Patagonian species, two from forest and one from steppe; seven others fromNew Caledonia, Australia, New Zealand; one Nearctic species). The latter three generamay have southern hemisphere connections that reflect patterns of continental drift buttheir phylogenetic relationships await analysis to test this possibility.

Study of other ceratopogonids in Patagonia indicates that most species are endemicin this region including species of Forcipomyia, Dasyhelea, and Stilobezzia (all of which aremembers of the subgenus Acanthohelea KIEFFER and all restricted to Nothofagus forest).

In addition to those endemic Patagonian genera discussed above, several genera orsubgenera of Ceratopogonidae are restricted or nearly restricted to the Neotropical Re-gion: Forcipomyia (Rhynchoforcipomyia WIRTH & DOW) (seven species), Culicoides (AnilomyiaVARGAS) (17 species), C. (Cotocripus BRÈTHES) (five species), C. (Macfiella FOX) (two species),C. (Mataemyia VARGAS) (15 species), C. (Psychophaena PHILIPPI) (two species), BaeodasymyiaCLASTRIER & RACCURT (five species), Baeohelea WIRTH & BLANTON (one species), BahiaheleaWIRTH (one species), Cacaohelea WIRTH & GROGAN (two species), Leptohelea WIRTH & BLAN-TON (one species), Parastilobezzia WIRTH & BLANTON (one species), Echinohelea MACFIE (Echi-


noideshelea WIRTH) (one species), Fittkauhelea WIRTH & BLANTON (one species), SchizonyxheleaCLASTRIER (two species), Clinohelea KIEFFER (Ceratobezzia KIEFFER) (one species), GroganheleaSPINELLI & DIPPOLITO (one species), Lanehelea WIRTH & BLANTON (two species), NeobezziaWIRTH & RATANAWORABHAN (eight species), Sphaerohelea SPINELLI & FELIPPE-BAUER (onespecies), Clastrieromyia SPINELLI & GROGAN (four species), Pachyhelea WIRTH (one of the twospecies extends north into Texas), Amerohelea GROGAN & WIRTH (one of the 11 knownspecies extends north into Texas), Paryphoconus ENDERLEIN (one of 40 species is Nearctic).Several of these groups are clearly monophyletic and this indicates that they have speciatedwithin the Neotropical Region (i.e. Baeodasymyia + Baeohelea, Clastrieromyia, Paryphoconus).

A few of these genera, each with only 1-2 species, are endemic to Central Americaand northern South America: Cacaohelea (Costa Rica), Leptohelea (Costa Rica, Colombia),Parastilobezzia (Costa Rica, Colombia).

In Central America, the genus Ceratoculicoides WIRTH & RATANAWORABHAN is present intropical Mexico (one described species) and Costa Rica (one undescribed species) and thisis likely a southern extension of an otherwise Holarctic genus.

Within Costa Rica we recognize a pattern, common in many groups of organisms,that lowland species tend to be broadly distributed in the Neotropical Region and thosethat occur higher than 1500 meters in elevation tend to be more restricted in distributionand are often endemic to Costa Rica or Costa Rica and Panama. This general pattern willalmost certainly be true in other high elevation areas in the Neotropics once they are inves-tigated more thoroughly.


6CLASSIFICATION

Ceratopogonidae belong to the superfamily Chironomoidea and are the sister group of theChironomidae. Worldwide, extant Ceratopogonidae are divided into four well establishedsubfamilies, based on cladistic relationships (BORKENT, 2000a; BORKENT & CRAIG, 2004):Leptoconopinae (2 genera), Forcipomyiinae (2 genera), Dasyheleinae (1 genus) and Cer-atopogoninae (104 genera). All subfamilies are known from the Neotropical Region (Table4). REMM (1975) proposed recognizing the Forcipomyiinae and Dasyheleinae as tribes inthe subfamily Forcipomyiinae and, as sister groups, this is conceivable but has not yet beenwidely accepted.

The Ceratopogoninae, which make up the bulk of the family both generically and atthe species level, is divided into 6 tribes: Culicoidini (3 genera), Ceratopogonini (57 gen-era), Heteromyiini (8 genera), Sphaeromiini (28 genera), Palpomyiini (6 genera) and theStenoxenini (2 genera). The relationships among these tribes are poorly understood, al-though it is clear that the Culicoidini are the sister group of the remaining tribes, theCeratopogonini are paraphyletic in relation to the remaining tribes and that the Palpomyi-ini and Stenoxenini form a monophyletic group. The relationships among most of thegenera of Ceratopogonini and the relationships among and within the genera of Heter-omyiini and Sphaeromiini are particularly in need of cladistic analysis.

Although 5925 extant species have been described worldwide (as of May, 2007) it isclear that many more remain unnamed. It is certainly reasonable to suggest that at least15,000 species exist on our planet. Within the Neotropical Region, the Andes are virtuallyuncollected, in spite of the fact that Ceratopogonidae are abundant and diverse there(BORKENT, pers. obs.). Many areas of Brazil, Peru, northern Chile and northwestern Argen-tina are also particularly poorly known. Limited portions of the tropical rain forests ofCentral America and Amazonia were extensively collected during the 1960s and 1970s,especially by F.S. BLANTON and E. FITTKAU, respectively, providing the basis for numeroustaxonomic papers mainly by W.W. WIRTH and his coworkers. Nevertheless, recent collec-tions of biting midges in Costa Rica and Peruvian Amazonia reveal the presence of manymore undescribed species (BORKENT & SPINELLI, pers. obs.), strongly suggesting that fur-ther collecting will be richly rewarded even in areas which have been previously sampled.Revisions are needed for virtually all genera; even Culicoides, which has drawn the mosttaxonomic attention, has many undescribed species and its subgeneric classification is ingreat need of analysis.


SUBFAMILY LEPTOCONOPINAE NOÈ, 1907: 143

Genus LEPTOCONOPS SKUSE

LEPTOCONOPS SKUSE, 1889: 288. Type species: Leptoconops stygius SKUSE, by monotypy.TERSESTHES TOWNSEND, 1893: 370. Type species: Tersesthes torrens TOWNSEND, by original

designation.MYCTEROTYPUS NOÈ, 1905: 114. Type species: Mycterotypus bezzii NOÈ, designation by

CARTER, 1921: 3.PROTERSESTHES KIEFFER, 1921c: 107. Type species: Tersesthes brasiliensis LUTZ, by original

designation.

REFERENCES: RONDEROS, 1988 (key to females from Argentina); RONDEROS & SPINELLI,1992 (key to females); WIRTH & ATCHLEY, 1973 (key to subgenera of females, otherthan Proleptoconops; key to Nearctic females).

Subgenus BRACHYCONOPS WIRTH & ATCHLEY

BRACHYCONOPS WIRTH & ATCHLEY, 1973: 15 (as subgenus of Leptoconops). Type species:Leptoconops californiensis WIRTH & ATCHLEY, by original designation.

patagoniensis RONDEROS, 1990b: 423 (female; Argentina). Distr.- Southern Argentina.

Subgenus HOLOCONOPS KIEFFER

HOLOCONOPS KIEFFER, 1918: 135. Type species: Leptoconops kerteszi KIEFFER, by originaldesignation.

MICROCONOPS KIEFFER, 1921c: 108. Type species: Microconops vexans KIEFFER, by originaldesignation.

REFERENCE: CLASTRIER & WIRTH, 1978 (key to Nearctic males and females).

bassoi RONDEROS & SPINELLI, 1993: 305 (male; Chile). Distr.- Northern Chile.bequaerti (KIEFFER), 1925: 405 (Holoconops, as becquaerti; female; Honduras). Distr.- USA

(Texas) to Panama and Trinidad.hondurensis HOFFMAN, 1926: 135. Honduras.

knowltoni CLASTRIER & WIRTH, 1978: 26 (male, female; USA, California). Distr.- USA(western) south to Mexico (Sinaloa), Brazil (Santa Catarina).

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


Subgenus LEPTOCONOPS SKUSE

brasiliensis (LUTZ), 1913: 66 (Tersesthes; female; Brazil). Distr.- Argentina (Salta), Brazil(Amazonas), Uruguay.

casali CAVALIERI & CHIOSSONE, 1966: 45 (female; Argentina). Distr.- Argentina (Salta, SanLuis), Uruguay.

chilensis FORATTINI, 1958: 38 (female; Chile). Distr.- Bolivia, Chile, Argentina (Jujuy, Salta,La Pampa).

panamensis RONDEROS & SPINELLI, 1993: 308 (male; Panama). Distr.- Panama.petrocchiae SHANNON & DEL PONTE, 1927: 734 (female; Argentina). Distr.- Northwestern

Argentina.ricardoi RONDEROS & SPINELLI, 1992: 43 (female; Argentina). Distr.- Southwestern Argentina.venezuelensis ORTÍZ, 1952a: 165 (female; Venezuela). Distr.- Venezuela.

Subgenus MEGACONOPS WIRTH & ATCHLEY

MEGACONOPS WIRTH & ATCHLEY, 1973: 18 (as subgenus of Leptoconops). Type species:Leptoconops floridensis WIRTH, by original designation.

floridensis WIRTH, 1951a: 282 (female; USA, Florida). Distr.- USA (Florida), Jamaica, Colombia.

SUBFAMILY FORCIPOMYIINAE LENZ, 1934: 96

Genus ATRICHOPOGON KIEFFER

ATRICHOPOGON KIEFFER, 1906a: 53 (as subgenus of Ceratopogon). Type species:Ceratopogon exilis COQUILLETT (= Ceratopogon levis COQUILLETT), designation by COQUILLETT,1910: 512.

REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE, 1931(key males and females from Patagonia and southern Chile); MACFIE, 1939b (key tomales and females from the Neotropical Region).

Subgenus ATRICHOPOGON KIEFFER

adamsoni MACFIE, 1937a: 4 (female; Trinidad). Distr.- Mexico (Chiapas), Trinidad.albinensis INGRAM & MACFIE, 1931: 228 (female; Argentina). Distr.- Argentina (Buenos Aires).altivolans MACFIE, 1949: 113 (female; Mexico). Distr.- Mexico (Chiapas).aridus SPINELLI & MARINO, in SPINELLI et al., 2006: 304 (male, female; Argentina). Distr.-

Argentina (Mendoza, Rio Negro).asuturus BORKENT & PICADO, 2004: 29 (male; Costa Rica). Distr.- Costa Rica.auricoma KIEFFER, 1917b: 300 (male, female; Colombia). Distr.- Colombia.balseiroi SPINELLI, 1982: 206 (male, female; Argentina). Distr.- Argentina (Buenos Aires).barbatus BORKENT & PICADO, 2004: 23 (male; Costa Rica). Distr.- Costa Rica.


beccus BORKENT & PICADO, 2004: 31 (male; Costa Rica). Distr.- Costa Rica.bicuspis BORKENT & PICADO, 2004: 9 (male, female; Costa Rica). Distr.- Costa Rica.bifidus EWEN, in EWEN & SAUNDERS, 1958: 694 (larva, pupa, male, female; Brazil). Distr.-

Brazil (Rio de Janeiro).brasiliensis MACFIE, 1939b: 184 (female; Brazil). Distr.- Brazil (Santa Catarina).brevipalpis MACFIE, 1944: 298 (male, female; Trinidad). Distr.- Trinidad.caribbeanus EWEN, in EWEN & SAUNDERS, 1958: 683 (larva, pupa, male, female; Tobago).

Distr.- Tobago.carnatus BORKENT & PICADO, 2004: 13 (male; Costa Rica). Distr.- Costa Rica.carpinteroi MARINO & SPINELLI, 2004c: 157 (male, female; Argentina). Distr.- Northern Argentina.casali CAVALIERI & CHIOSSONE, 1973: 153 (male, female; Argentina). Distr.- Northeastern

Argentina.colossus BORKENT & PICADO, 2004: 10 (male, female; Costa Rica). Distr.- Costa Rica.columbianus KIEFFER, 1917b: 303 (female; Colombia). Distr.- Colombia, Brazil (Santa Catarina).comechingon SPINELLI & MARINO, in SPINELLI et al., 2006: 306 (male, female; Argentina).

Distr.- Argentina (Córdoba, Rio Negro).costalis MACFIE, 1939b: 179 (female; Brazil). Distr.- Brazil (Santa Catarina).costaricae MACFIE, 1953: 98 (female; Costa Rica). Distr.- Costa Rica.cryptogamus MACFIE, 1939b: 182 (female; Brazil). Distr.- Brazil (Santa Catarina).delpontei CAVALIERI & CHIOSSONE, 1972: 121 (male, female; Argentina). Distr.- Northeastern

Argentina.depilis MACFIE, 1939b: 178 (female; Brazil). Distr.- Brazil (Santa Catarina).didymothecae MACFIE, 1953: 99 (female; Costa Rica). Distr.- Costa Rica.domizii SPINELLI, 1982: 201 (male, female; Argentina). Distr.- Argentina (Buenos Aires).echinodes MACFIE, 1939b: 194 (female; Brazil). Distr.- Brazil (Santa Catarina).endemicus SPINELLI & MARINO, in SPINELLI et al., 2006: 304 (male, female; Argentina).

Distr.- Argentina (Rio Negro).eucnemus MACFIE, 1939b: 185 (male, female; Brazil). Distr.- Brazil (Santa Catarina).fiebrigi KIEFFER, 1917b: 302 (male; Paraguay). Distr.- Paraguay.fimbriatus MACFIE, 1939b: 194 (as variety of gordoni; female). Distr.- Brazil (Santa Catarina).flavicaudae MACFIE, 1939b: 181 (female; Brazil). Distr.- Brazil (Santa Catarina).

pilosior MACFIE, 1939b: 182 (as variety of flavicaudae). Brazil.flavipes LUTZ, 1914: 90 (female; Brazil). Distr.- Brazil (type locality not stated).flumineus MACFIE, 1935a: 52 (female; Brazil). Distr.- Brazil (Maranhão, Santa Catarina).gamboai BORKENT & PICADO, 2004: 26 (male; Costa Rica). Distr.- Costa Rica.glaber MACFIE, 1935a: 50 (female; Brazil). Distr.- Costa Rica, Trinidad, Brazil (Pará,

Maranhão).globulifer MACFIE, 1939b: 191 (female; Brazil). Distr.- Brazil (Santa Catarina).gordoni MACFIE, 1938: 164 (female; Trinidad). Distr.- Costa Rica, Trinidad, Brazil (Santa

Catarina).granditergitus BORKENT & PICADO, 2004: 27 (male; Costa Rica). Distr.- Costa Rica.granditibialis BORKENT & PICADO, 2004: 20 (male, female; Costa Rica). Distr.- Costa Rica.guianensis MACFIE, 1940b: 184 (female; Guyana). Distr.- Guyana.harrisi MACFIE, 1938: 163 (female; Trinidad). Distr.- Trinidad.


homofacies SPINELLI, in SPINELLI et al., 1989: 734 (male, female; Argentina). Distr.- Argentina(Buenos Aires).

inacayali SPINELLI & MARINO, in SPINELLI et al., 2006: 309 (male, female; Argentina). Distr.-Argentina (Chuvut, Rio Negro).

incultus EWEN, in EWEN & SAUNDERS, 1958: 699 (larva, pupa, male, female; Costa Rica).Distr.- Costa Rica.

insignipalpis MACFIE, 1940a: 73 (male, female; Brazil). Distr.- Northeastern Brazil.insigniventris MACFIE, 1935a: 51 (female; Brazil). Distr.- Brazil (Maranhão), Trinidad.lacajae MACFIE, 1953: 100 (male; Costa Rica). Distr.- Costa Rica.lituratus (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- St. Vincent.lobatus BORKENT & PICADO, 2004: 14 (male, female; Costa Rica). Distr.- Costa Rica.longicornis EWEN, in EWEN & SAUNDERS, 1958: 700 (larva, male, female; Costa Rica). Distr.-

Costa Rica.maculipennis CLASTRIER, 1968: 89 (male, female; French Guiana). Distr.- French Guiana.magnus BORKENT & PICADO, 2004: 16 (male, female; Costa Rica). Distr.- Costa Rica.mendozae INGRAM & MACFIE, 1931: 229 (female; Argentina). Distr.- Argentina (Mendoza).mexicanus HUERTA, 2001: 373 (male, female; Mexico). Distr.- Mexico (Chiapas).nanus MACFIE, 1940b: 184 (male, female; Guyana). Distr.- Guyana.nebulosus MACFIE, 1939b: 192 (female; Brazil). Distr.- Brazil (Santa Catarina).novaeteutoniae MACFIE, 1939b: 189 (female; Brazil). Distr.- Brazil (Santa Catarina).nubeculosus MACFIE, 1949: 113 (female; Mexico). Distr.- Mexico (Chiapas).obfuscatus INGRAM & MACFIE, 1931: 173 (male, female; Argentina). Distr.- Southern Chile,

southwestern Argentina south to Tierra del Fuego.obnubilus INGRAM & MACFIE, 1931: 175 (male, female; Argentina). Distr.- Southern Chile,

southwestern Argentina.chilensis INGRAM & MACFIE, 1931: 175. Argentina.assimilis INGRAM & MACFIE, 1931: 176. Argentina.

obscurus EWEN, in EWEN & SAUNDERS, 1958: 685 (larva, pupa, female; Puerto Rico). Distr.-Puerto Rico.

ocumare (ORTÍZ), 1952b: 254 (Monohelea; male; Venezuela). Distr.- Venezuela.ornatipennis CLASTRIER, 1987: 271 (male; French Guiana). Distr.- French Guiana.pachycnemus MACFIE, 1953: 101 (female; Costa Rica). Distr.- Costa Rica.pallidipes KIEFFER, 1917b: 301 (female; Paraguay). Distr.- Mexico (Chiapas), Costa Rica,

Paraguay.palpalis MACFIE, 1939b: 196 (male, female; Brazil). Distr.- Brazil (Santa Catarina).pectinatus MACFIE, 1939b: 195 (female; Brazil). Distr.- Brazil (Santa Catarina).penicillatus DELÉCOLLE & RIEB, 1994: 274 (female; Guadeloupe). Distr.- Guadeloupe.peruvianus KIEFFER, 1917b: 302 (male; Peru). Distr.- Peru.piceiventris KIEFFER, 1917b: 301 (female; Paraguay). Distr.- Paraguay.pictipennis CLASTRIER, 1979: 30 (male; French Guiana). Distr.- French Guiana.quartibrunneus BORKENT & PICADO, 2004: 30 (male; Costa Rica). Distr.- Costa Rica.redactus BORKENT & PICADO, 2004: 32 (male; Costa Rica). Distr.- Costa Rica.remigatus EWEN, in EWEN & SAUNDERS, 1958: 687 (larva, pupa, male, female; Brazil). Distr.-

Brazil (Rio de Janeiro).


rusticus MACFIE, 1939b: 190 (female; Brazil). Distr.- Brazil (Santa Catarina).sanctaeclarae MACFIE, 1949: 114 (male, female; Mexico). Distr.- Mexico (Chiapas).sanctilaurentii KIEFFER, 1917b: 303 (female; Paraguay). Distr.- Paraguay.saundersi EWEN, in EWEN & SAUNDERS, 1958: 691 (larva, pupa, male, female; Puerto Rico).

Distr.- Puerto Rico.sequax (WILLISTON), 1896: 282 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.setosilateralis BORKENT & PICADO, 2004: 33 (male; Costa Rica). Distr.- Costa Rica.seudoobfuscatus SPINELLI, 1982: 208 (male, female; Argentina). Distr.- Argentina (Buenos Aires).similis SPINELLI & MARINO, in SPINELLI et al., 2006: 316 (male, female; Argentina). Distr.-

Southern Chile, southwestern Argentina.spinosus BORKENT & PICADO, 2004: 12 (male; Costa Rica). Distr.- Costa Rica.taeniatus MACFIE, 1939b: 186 (female; Brazil). Distr.- Brazil (Santa Catarina).talarum SPINELLI, 1982: 204 (male, female; Argentina). Distr.- Argentina (Buenos Aires).tapantiensis BORKENT & PICADO, 2004: 34 (male; Costa Rica). Distr.- Costa Rica.tirzae BORKENT & PICADO, 2004: 28 (male; Costa Rica). Distr.- Costa Rica.thersites (WILLISTON), 1896: 280 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.trinidadensis MACFIE, 1937a: 5 (male; Trinidad). Distr.- Trinidad.tuberculatus EWEN, in EWEN & SAUNDERS, 1958: 709 (larva, pupa, female, male; Trinidad).

Distr.- Trinidad.umbratilis MACFIE, 1935a: 51 (female; Brazil). Distr.- Guyana, Brazil (Maranhão).utricularis MACFIE, 1953: 102 (female; Costa Rica). Distr.- Costa Rica.warmkei WIRTH, 1956c: 243 (male, female; Puerto Rico). Distr.- USA (Florida, Puerto Rico).woodfordi MACFIE, 1938: 162 (female; Trinidad). Distr.- Trinidad.yolancae BORKENT & PICADO, 2004: 24 (male; Costa Rica). Distr.- Costa Rica.

Subgenus LOPHOMYIDIUM CORDERO

LOPHOMYIDIUM CORDERO, 1929: 94. Type species: Lophomyidium uruguayense CORDERO,by original designation.

REFERENCE: WIRTH 1994a (key to Nearctic males and females).

archboldi WIRTH, 1994a: 27 (male, female; USA, Florida). Distr.- USA (California to Florida)to Argentina.

fusculus (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, New Jersey). Distr.- NorthernEurasia, North America, Central America and Caribbean to Bolivia and Brazil (Pará).

uruguayensis (CORDERO), 1929: 95 (Lophomyidium; male; Uruguay). Distr.- Paraguay, easternArgentina, Uruguay.

Genus FORCIPOMYIA MEIGEN

FORCIPOMYIA MEIGEN, 1818: 73, 75. Type species: Tipula bipunctata Linnaeus, designationby WESTWOOD, 1840: 126. Generic name first published in synonymy with Ceratopogonbut available under ICZN Code Article 11(e).


TETRAPHORA PHILIPPI, 1865: 630. Type species: Tetraphora fusca PHILIPPI, by monotypy.

REFERENCES: DEBENHAM, 1987 (key to males and females to subgenera from AustralasianRegion which includes all Neotropical subgenera other than Rhynchoforcipomyia andSaliohelea); INGRAM & MACFIE, 1931 (key to males and females from Patagonia andsouthern Chile); WIRTH & RATANAWORABHAN, 1978 (key to females to subgenera; theirgenus A near Lepidohelea = Pedilohelea).

Subgenus BLANTONIA WIRTH & DOW

BLANTONIA WIRTH & DOW, 1971: 289 (as subgenus of Forcipomyia). Type species:Forcipomyia caribbea WIRTH & DOW, by original designation.

caribbea WIRTH & DOW, 1971: 291 (male, female; Jamaica). Distr.- USA (Florida) to Trinidad,Mexico (Yucatán), Belize, Guyana.

Subgenus CALOFORCIPOMYIA SAUNDERS

CALOFORCIPOMYIA SAUNDERS, 1957: 680 (as subgenus of Forcipomyia). Type species:Forcipomyia caerulea SAUNDERS, by original designation.

REFERENCE: UTMAR & WIRTH, 1976 (key to New World males and females).

caerulea SAUNDERS, 1957: 681 (male, female; Brazil). Distr.- Panama, Colombia, Dominica,Brazil (Pará, Rio de Janeiro). .

copanensis UTMAR & WIRTH, 1976: 129 (male, female; Honduras). Distr.- Distr.- Honduras,Costa Rica, northern Argentina.

eukosma MACFIE, 1939b: 148 (male, female; Brazil). Distr.- Colombia, Dominica, Brazil(Pará, São Paulo, Rio de Janeiro, Santa Catarina).

furcifera MACFIE, 1940e: 920 (male; Brazil). Distr.- Brazil (Santa Catarina).glauca MACFIE, 1934: 144 (Great Britain). Distr.- Europe, North America (Alberta to Nova

Scotia to Arkansas and Florida) to Colombia and southern Brazil.hatoensis UTMAR & WIRTH, 1976: 131 (male, female; Panama). Distr.- Mexico (Sinaloa),

Costa Rica, Panama, Colombia.hermosa UTMAR & WIRTH, 1976: 117 (male, female; Brazil). Distr.- Costa Rica, Panama,

Colombia, Jamaica, Panama, Brazil (Amazonas, Pará).nigrescens MACFIE, 1939b: 156 (male, female; Brazil). Distr.- Panama, Brazil (São Paulo,

Rio de Janeiro, Santa Catarina).remigera UTMAR & WIRTH, 1976: 121 (male, female; Colombia). Distr.- Panama, Colombia,

French Guiana, Brazil (Pará).sabalitensis UTMAR & WIRTH, 1976: 127 (male, female; Costa Rica). Distr.- Honduras to

Colombia, Brazil (Paraná, Santa Catarina), northeastern Argentina.varicolor SAUNDERS, 1957: 683 (larva, pupa, male; Brazil). Distr.- Brazil (Rio de Janeiro).


Subgenus EUPROJOANNISIA BRÈTHES

EUPROJOANNISIA BRÈTHES, 1914: 155. Type species: Euprojoannisia platensis BRÈTHES,by original designation.

REFERENCES: BYSTRAK & WIRTH, 1978 (key to Nearctic larvae, pupae, males, females);HUERTA, 1996 (key to males and females from Mexico).

blantoni SORIA & BYSTRAK, 1975: 3 (larva, pupa, male, female; Brazil). Distr.- Eastern USAto Ecuador, Brazil (Bahia).

bromeliae SAUNDERS, 1957: 665 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro).calcarata (COQUILLETT), 1905: 64 (Ceratopogon; male; Mexico). Distr.- Eastern USA, Mexico

(Tabasco, Quintana Roo).dowi BYSTRAK & WIRTH, 1978: 21 (male, female; USA, Florida). Distr.- USA (Florida),

Mexico (Quintana Roo).esteparia MARINO & SPINELLI, 2001b: 12 (male, female; Argentina). Distr.- Southern Argentina.falcifera SAUNDERS, 1959: 39 (larva, pupa, male, female; Trinidad). Distr.- Venezuela, Trinidad.galliarii MARINO & SPINELLI, 1999a: 5 (female; Argentina). Distr.-. Argentina (Misiones).longispina SAUNDERS, 1957: 669 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro).lota (WILLISTON), 1896: 282 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.mortuifolii SAUNDERS, 1959: 35 (larva, pupa, male, female; Trinidad). Distr.- USA (Florida)

to St. Lucia.navaiae BYSTRAK & WIRTH, 1978: 31 (male, female; USA, Florida). Distr.- USA (Florida),

Mexico (Yucatán).platensis (BRÈTHES), 1914: 156 (Euprojoannisia; female; Argentina). Distr.- Argentina (Buenos Aires).quasiingrami MACFIE, 1939b: 164 (male; Brazil). Distr.- USA (Florida) south through

Caribbean to Brazil (Bahia, Santa Catarina).setigera SAUNDERS, 1959: 38 (larva, pupa, male, female; Trinidad). Distr.- Trinidad.setosicrus (KIEFFER), 1906b: 357 (Ceratopogon; male; Chile). Distr.- Southern Argentina and Chile.spatulifera SAUNDERS, 1957: 667 (larva, pupa, female; Brazil). Distr.- Brazil (Bahia, São

Paulo), Venezuela.unica BYSTRAK & WIRTH, 1978: 44 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas.

Subgenus FORCIPOMYIA MEIGEN

REFERENCES: HUERTA, 1996 (key to females from Mexico); MARINO & SPINELLI, 1999b (keyto species groups); MARINO & SPINELLI, 2001a (key to males and females from Patagonia);MARINO & SPINELLI, 2002 (key to squamitibia group); WIRTH, 1982a (key to argenteola group).

argenteola Species Group

argenteola MACFIE, 1939b: 146 (female; Brazil). Distr.- Brazil (Rio de Janeiro, Santa Catarina).calatheae WIRTH, 1982a: 573 (larva, pupa, male, female; Dominica). Distr.- Colombia,

Dominica, Brazil (Bahia, Santa Catarina). .


marini SPINELLI & DIPPOLITO, 1995: 155 (male, female; Venezuela). Distr.- Venezuela.quatei WIRTH, 1952a: 142 (male, female; USA, California). Distr.- USA (California to Florida)

to Brazil (Bahia).uramaensis SPINELLI & DIPPOLITO, 1995: 156 (female; Venezuela). Distr.- Venezuela.youngi WIRTH, 1982a: 579 (larva, pupa, male, female; Panama). Distr.- Costa Rica, Panama,

Ecuador.

fusca Species Group

caliginosella WIRTH, 1974: 5. New name for caliginosa. Distr.- Southwestern Argentina.caliginosa INGRAM & MACFIE, 1931: 167 (preoccupied by Forcipomyia caliginosa (INGRAM &MACFIE), 1924). Argentina.

fusca (PHILIPPI), 1865: 630 (Tetraphora; female; Chile). Distr.- Central and southern Chilesouth to Deceit Island, southwestern Argentina south to Tierra del Fuego.

patagonica INGRAM & MACFIE, 1931: 165. Argentina.piroskyi CAVALIERI, 1961b: 172 (female; Argentina). Distr.- Argentina (Tierra del Fuego).wygodzinskyi CAVALIERI, 1961a: 17 (female; Argentina). Distr.- Argentina (Tierra del Fuego).

delpontei CAVALIERI, 1961b: 169. Argentina.

genualis Species Group

genualis (LOEW), 1866: 128 (Ceratopogon; male; Cuba). Distr.- USA (Louisiana to Florida),Bahamas, Mexico (Yucatán, Chiapas) to Colombia, Galápagos Islands, Cayman Islands,Cuba to Argentina (Buenos Aires province); Sao Tomé.

propinqua (WILLISTON), 1896: 279 (Ceratopogon). St. Vincent.raleighi MACFIE, 1938: 160. Trinidad.

harpegonata WIRTH & SORIA, 1975: 19 (larva, pupa, male, female; Puerto Rico). Distr.-Costa Rica, Bahamas, Jamaica, Puerto Rico, Trinidad to Brazil (Amazonas, Pará, Bahia).

taragui MARINO, SPINELLI & CAZORLA, 2002: 7. New name for guarani. Argentina. Distr.-Paraguay, Uruguay, Argentina (Corrientes, Misiones).

guarani MARINO & SPINELLI, 1999: 448 (preoccupied by Forcipomyia guarani RONDEROS &SPINELLI, 1999). Argentina.

poulaineae Species Group

edwardsiana WIRTH, 1974: 5. New name for edwardsi. Distr.- Southwestern Argentina.edwardsi INGRAM & MACFIE, 1931: 160 (preoccupied by Forcipomyia edwardsi (SAUNDERS),

1925). Argentina.minitheca MARINO & SPINELLI, 2001a: 111 (male, female; Chile). Distr.- Southern Chile

south to Deceit Island, southwestern Argentina.multipicta INGRAM & MACFIE, 1931: 157 (male, female; Argentina). Distr.- Southern Chile,

southwestern Argentina.muzoni MARINO & SPINELLI, 2004a: 150 (male, female; Argentina). Distr.- Argentina (Rio Negro).nuncupata MACFIE, 1949: 110 (male, female; Mexico). Distr.- Mexico (Chiapas).


ornatipennis MACFIE, 1939b: 151 (male, female; Brazil). Distr.- Costa Rica, Brazil (SantaCatarina).

poulaineae INGRAM & MACFIE, 1931: 227 (male; Argentina). Distr.- Brazil (Bahia, SantaCatarina), Argentina (Buenos Aires).

somuncurensis MARINO & SPINELLI, 2001a: 116 (male, female; Argentina). Distr.- Argentina(Rio Negro).

townsendi KNAB, 1915: 111 (male, female; Peru). Distr.- Peru.utae KNAB, 1915: 109 (female; Peru). Distr.- Peru.yamana MARINO & SPINELLI, 2001a: 118 (female; Chile). Distr.- Chile (Deceit Island).zonogaster INGRAM & MACFIE, 1931: 168 (male; Chile). Distr.- Southern Chile.

squamitibia Species Group

catarinensis MARINO & SPINELLI, 2002: 309 (male, female; Brazil). Distr.- Mexico, CostaRica, Colombia, Brazil.

chilensis (PHILIPPI), 1865: 601 (Ceratopogon; male; Chile). Distr.- Central and southern Chile,southwestern Argentina south to Tierra del Fuego.

pinamarensis SPINELLI, 1983a: 121 (male, female; Argentina). Distr.- Costa Rica, Panama,Venezuela, Brazil (Pará, Bahia, Rio de Janeiro), Uruguay, Argentina (Buenos Aires).

quechua MARINO & SPINELLI, 2002: 312 (male, female; Argentina). Distr.- Argentina (Salta).rioplatensis MARINO & SPINELLI, 2002: 314 (male, female; Argentina). Distr.- Argentina

(Buenos Aires), Uruguay.sexvittata WIRTH, 1956c: 248 (male, female; Costa Rica). Distr.- Costa Rica.spatuligera MACFIE, 1949: 111 (female; Mexico). Distr.- Mexico (Chiapas).squamitibia LUTZ, 1914: 88 (female; Brazil). Distr.- Colombia, Brazil (Rio de Janeiro, Santa

Catarina), Puerto Rico, Trinidad.soriai WIRTH, 1991b: 168. Brazil.

tenuisquamipes WIRTH, 1952b: 89 (female; Juan Fernández Islands). Distr.- Juan FernándezIslands.

Unplaced to Species Group

flava (WILLISTON), 1896: 280 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.pictoni MACFIE, 1938: 161 (male; Trinidad). Distr.- USA (Louisiana to Florida) to Brazil

(Amazonas, Pará, Bahia, São Paulo).

Subgenus LASIOHELEA KIEFFER

CENTRORHYNCHUS LUTZ, 1913: 62 (preoccupied by Centrorhynchus STEVEN or FISHER

WALDHEIM, 1829). Type species: Centrorhynchus stylifer LUTZ, by original designation.LASIOHELEA KIEFFER, 1921c: 115. Type species: Atrichopogon pilosipennis KIEFFER

(= Ceratopogon velox WINNERTZ), by original designation.


REFERENCES: AITKEN et al., 1975 (species in Trinidad and Tobago); RONDEROS & SPINELLI,1999 (revision of Neotropical species).

anitae HUERTA & IBAÑEZ-BERNAL, 1996: 350 (male; Mexico). Distr.- Mexico (San Luis Potosi,Chiapas), Costa Rica, Brazil (Bahia, Itabuna), northeastern Argentina.

attenuata SAUNDERS, 1964: 469 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica,Bolivia, Trinidad.

cacaophila RONDEROS & SPINELLI, 1999: 152 (male; Venezuela). Distr.- Costa Rica, Venezuela.cornuta SAUNDERS, 1964: 464 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica,

Trinidad, Brazil (São Paulo).guarani RONDEROS & SPINELLI, 1999: 153 (female; Paraguay). Distr.- Paraguay.insigniforceps MACFIE, 1939b: 165 (male; Brazil). Distr.- Brazil (Santa Catarina).intermedia SAUNDERS, 1964: 468 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica.multidentata RONDEROS & SPINELLI, 1999: 153 (female; Brazil). Brazil (Bahia, Itaguipe,

Fazenda Almirante).quasicornuta SAUNDERS, 1964: 467 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica.saltensis (CAVALIERI), 1962: 360 (Lasiohelea; female; Argentina). Distr.- Northwestern Argentina.stylifer (LUTZ), 1913: 63 (Centrorhynchus; female; Brazil). Distr.- Belize to Ecuador, Trinidad,

Venezuela to northeastern Argentina.

Subgenus LEPIDOHELEA KIEFFER

LEPIDOHELEA KIEFFER, 1917b: 364. Type species: Ceratopogon chrysolophus KIEFFER, byoriginal designation.

REFERENCES: WIRTH, 1991c (key to species groups; key to males and females of bicolorgroup); WIRTH & SPINELLI, 1992a (males and females of seminole subgroup); WIRTH &SPINELLI, 1993a (key to males and some females of annulatipes group).

abercrombyi MACFIE, 1938: 161 (male, female; Trinidad). Distr.- Trinidad.annulatipes MACFIE, 1939b: 154 (male; Brazil). Distr.- Colombia, Brazil (Pará, Bahia, Santa Catarina).bahiensis WIRTH & SPINELLI, 1993a: 113 (male, female; Brazil). Distr.- Trinidad, Brazil (Bahia).basifemoralis WIRTH & SPINELLI, 1993a: 113 (male, female; Jamaica). Distr.- USA (Florida)

to Trinidad, Panama.bicolor LUTZ, 1914: 89 (male; Brazil). Distr.- Costa Rica, Ecuador, Brazil (Rio de Janeiro,

Santa Catarina).discoloripes MACFIE, 1939b: 159. Brazil.

bifida WIRTH & SPINELLI, 1993a: 115 (male, female; Jamaica). Distr.- Jamaica.brasiliensis MACFIE, 1939b: 153 (male; Brazil). Distr.- Ecuador, Trinidad, Brazil (Santa Catarina).convexipenis WIRTH & SPINELLI, 1993a: 116 (male, female; Colombia). Distr.- Colombia.dubia MACFIE, 1939b: 162 (male; Brazil). Distr.- Brazil (Santa Catarina).edmistoni WIRTH & SPINELLI, 1993b: 624 (larva, pupa, male, female; USA, Maryland).

Distr.- Eastern USA, Argentina (Buenos Aires).euthystyla WIRTH & SPINELLI, 1993a: 117 (male, female; Colombia). Distr.- Panama, Colombia.


flavifemoris MACFIE, 1940d: 24 (male, female; Guyana). Distr.- Guyana.herediae WIRTH & SPINELLI, 1993a: 119 (male, female; Costa Rica). Distr.- Costa Rica.hobbsi WIRTH & SPINELLI, 1993a: 121 (male, female; Dominica). Distr.- Mexico (Tabasco)

to Colombia, Jamaica to Grenada.kuanoskeles MACFIE, 1939b: 150 (male; Brazil). Distr.- Brazil (Santa Catarina).lacrimatorii MACFIE, 1939b: 161 (male, female; Brazil). Distr.- Brazil (Santa Catarina).luteigenua WIRTH & SPINELLI, 1992a: 353 (male, female; Costa Rica). Distr.- USA (Florida)

to Colombia and Brazil (Bahia).seminole WIRTH, 1976: 81 (male, female; USA, Florida). Distr.- USA (Florida) to Brazil

(Rio de Janeiro).squamithorax CLASTRIER, 1972: 170 (male, female; French Guiana). Distr.- French Guiana.varipennis WIRTH & WILLIAMS, 1957: 8 (female; Bermuda). Distr.- USA (Florida and Puerto

Rico), Bermuda, Paraguay, northeastern Argentina.weemsi WIRTH & SPINELLI, 1993a: 123 (male, female; USA, Florida). Distr.- USA (Maryland

to Florida, Arizona), Mexico (Baja California).winderi WIRTH, 1991b: 171 (male, female; Brazil). Distr.- Ecuador, Brazil (Bahia).

Subgenus METAFORCIPOMYIA SAUNDERS

METAFORCIPOMYIA SAUNDERS, 1957: 685 (as subgenus of Forcipomyia). Type species:Forcipomyia cerifera SAUNDERS, by original designation.

REFERENCE: MARINO & SPINELLI, 2003 (key to males and females of New World species).

cerifera SAUNDERS, 1957: 685 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro),northeastern Argentina.

darwini MARINO & SPINELLI, 2003: 24 (male, female; Chile). Distr.- Chile (Deceit Island).maculosa INGRAM & MACFIE, 1931: 159 (female; Argentina). Distr.- Southwestern Argentina.mapuche MARINO & SPINELLI, 2003: 26 (male; Argentina). Distr.- Southwestern Argentina,

southern Chile.morenoi MARINO & SPINELLI, 2003: 26 (male; Argentina). Distr.- Southwestern Argentina.williamsi MARINO & SPINELLI, 1999a: 5 (male, female; Argentina). Distr.- Argentina (Buenos

Aires, Martin Garcia Island).

Subgenus MICROHELEA KIEFFER

MICROHELEA KIEFFER, 1917b: 364. Type species: Atrichopogon microtomus KIEFFER,designation by KIEFFER, 1921a: 7, with explanatory notation on erroneous previoustype designation).

PHASMIDOHELEA MAYER, 1937: 233. Type species: Phasmidohelea crudelis MAYER

(= Ceratopogon fuliginosus MEIGEN), by original designation.

REFERENCES: CLASTRIER & WIRTH, 1995 (key to females in ixodoides group); HUERTA, 1996(key to males and females from Mexico); WIRTH, 1972 (key to females in fuliginosa group).


alleni CLASTRIER & WIRTH, 1995: 117 (female; Costa Rica). Distr.- Costa Rica.amazonica WIRTH, 1971: 241 (female; Brazil). Distr.- Brazil (Amazonas).antioquiae CLASTRIER & WIRTH, 1995: 142 (female; Colombia). Distr.- Colombia.belemensis CLASTRIER & WIRTH, 1995: 140 (female; Brazil). Distr.- Colombia, Brazil (Pará).brasiliana CLASTRIER & WIRTH, 1995: 116 (female; Peru). Distr.- Peru.breelandi CLASTRIER & WIRTH, 1995: 114 (female; Panama). Distr.- Panama.brevilabellata CLASTRIER & WIRTH, 1995: 136 (female; French Guiana). Distr.- French Guiana.broadheadi CLASTRIER & WIRTH, 1995: 129 (female; Panama). Distr.- Panama.castneri CLASTRIER & WIRTH, 1995: 111 (female; Peru). Distr.- Peru, Brazil (Pará).catarina CLASTRIER & WIRTH, 1995: 131 (female; Brazil). Distr.- Brazil (Santa Catarina).coheni CLASTRIER & WIRTH, 1995: 132 (female; Ecuador). Distr.- Ecuador.colombiana CLASTRIER & WIRTH, 1995: 102 (female; Colombia). Distr.- Costa Rica, Colombia.cristata CLASTRIER & WIRTH, 1995: 135 (female; Brazil). Distr.- Brazil (Santa Catarina).desutterae CLASTRIER & WIRTH, 1995: 100 (female; French Guiana). Distr.- French Guiana,

Brazil (Pará).donskoffi CLASTRIER & WIRTH, 1995: 125 (female; French Guiana). Distr.- French Guiana.dunklei CLASTRIER & WIRTH, 1995: 130 (female; Peru). Distr.- Peru.eriophora (WILLISTON), 1896: 279 (Ceratopogon; male; St. Vincent). Distr.- USA (Florida),

Caribbean, Belize to Brazil (Bahia).felippebauerae CLASTRIER & WIRTH, 1995: 122 (female; Peru). Distr.- Peru.grandcolasi CLASTRIER & WIRTH, 1995: 112 (female; French Guiana). Distr.- French Guiana.franklini CLASTRIER & WIRTH, 1995: 141 (female; Costa Rica). Distr.- Costa Rica, Panama.fuliginosa (MEIGEN), 1818: 86 (Ceratopogon; Germany). Distr.- Widespread in the Palaearctic,

Oriental, Afrotropical, Nearctic and Neotropical regions. In the Neotropics south toArgentina and Uruguay.

crudelis KNAB, 1914: 66 (preoccupied by Forcipomyia crudelis (KARSCH), 1886). Mexico.tropica (KIEFFER), 1917b: 297 (Ceratopogon). Costa Rica.

galapagensis (COQUILLETT), 1901b: 372 (Ceratopogon; male; Galápagos Islands). Distr.-Galápagos Islands.

guyana CLASTRIER & WIRTH, 1995: 124 (female; French Guiana). Distr.- French Guiana.insignipalpis MACFIE, 1949: 109 (female; Mexico). Distr.- Mexico (Chiapas), Costa Rica,

Panama, Colombia, Brazil (Pará).iquitosensis CLASTRIER & WIRTH, 1995: 139 (female; Peru). Distr.- Peru.ixodoides (FIEBRIG-GERTZ), 1928: 290 (Ceratopogon; female; Paraguay). Distr.- Paraguay.kawensis CLASTRIER & WIRTH, 1995: 120 (female; French Guiana). Distr.- French Guiana, Peru.luteisquamosa WIRTH, 1972: 575 (female; Brazil). Distr.- Brazil (Santa Catarina).mayeri FORATTINI & LANE, 1955: 4. New name for crudelis. Distr.- Costa Rica.

crudelis (MAYER), 1937: 233 (Phasmidohelea; female) (preoccupied by Forcipomyia crudelis(KARSCH), 1886). Costa Rica.

menzeli CLASTRIER & WIRTH, 1995: 106 (female; Ecuador). Distr.- Ecuador.microtoma (KIEFFER), 1917b: 299 (Atrichopogon; female; Paraguay). Distr.- Belize and

Dominica to Brazil (Amazonas, Santa Catarina), Paraguay.minisquamosa WIRTH, 1972: 570 (male, female; Belize). Distr.- Belize to Costa Rica, Brazil

(Amazonas, Pará).


moorei CLASTRIER & WIRTH, 1995: 133 (female; Brazil). Distr.- Brazil (Santa Catarina).neotropica CLASTRIER & WIRTH, 1995: 126 (female; Peru). Distr.- Peru.nigrimaxillata CLASTRIER & WIRTH, 1995: 118 (female; Peru). Distr.- Peru.novaeteutoniae CLASTRIER & WIRTH, 1995: 107 (female; Brazil). Distr.- Brazil (Santa Catarina).obesa COSTA LIMA, 1928: 85 (female; Brazil). Distr.- Brazil (Amazonas).penultimata WIRTH, 1972: 573 (male, female; Brazil). Distr.- Belize and Jamaica to Colombia,

Brazil (Santa Catarina).peruviana CLASTRIER & WIRTH, 1995: 123 (female; Peru). Distr.- Peru.plaumanni CLASTRIER & WIRTH, 1995: 127 (female; Brazil). Distr.- Brazil (Santa Catarina).raposoensis CLASTRIER & WIRTH, 1995: 144 (female; Colombia). Distr.- Colombia.rettenmeyerorum CLASTRIER & WIRTH, 1995: 104 (female; Panama). Distr.- Panama.squamosa LUTZ, 1914: 87 (male, female; Brazil). Distr.- Colombia, Bolivia, Brazil (Amazonas,

Bahia, Rio de Janeiro, Santa Catarina), Argentina (Mendoza).tettigonaris WIRTH & CASTNER, 1990: 159 (female; Peru). Distr.- Costa Rica, Dominican

Republic, Peru.thomasi CLASTRIER & WIRTH, 1995: 115 (female; Brazil). Distr.- Brazil (Pará).valleensis CLASTRIER & WIRTH, 1995: 145 (female; Colombia). Distr.- Colombia.vernoni CLASTRIER & WIRTH, 1995: 138 (female; Colombia). Distr.- Colombia.wagneri (SÉGUY), 1941: 85 (Phasmidohelea; female; Brazil). Distr.- Brazil (Rio de Janeiro).willistoni WIRTH, 1971: 242 (male, female; Puerto Rico). Distr.- Jamaica, Puerto Rico,

Colombia, Brazil (Santa Catarina).

Subgenus PEDILOHELEA DE MEILLON & WIRTH

PEDILOHELEA DE MEILLON & WIRTH, 1980: 9 (as subgenus of Forcipomyia). Type species:Forcipomyia clastrieri DESSART, by original designation.

REFERENCE: DE MEILLON & WIRTH, 1980 (key to males, some females).

aitkeni DE MEILLON & WIRTH, 1980: 12 (male, female; Brazil). Distr.- Colombia, Brazil (Pará).archboldi DE MEILLON & WIRTH, 1980: 13 (male, female; Dominica). Distr.- Jamaica, Puerto

Rico, Brazil (Pará).raposoi DE MEILLON & WIRTH, 1980: 21 (male; Colombia). Distr.- Colombia.spangleri DE MEILLON & WIRTH, 1980: 21 (male, female; Guatemala). Distr.- Mexico

(Yucatán), Guatemala, Panama, Colombia, Ecuador.spilmani DE MEILLON & WIRTH, 1980: 22 (male, female; Dominica). Distr.- Panama,

Colombia, Dominica, Trinidad, Brazil (Santa Catarina).

Subgenus PHYTOHELEA REMM

PHYTOHELEA REMM, 1971: 189 (as subgenus of Forcipomyia). Type species: Ceratopogonbromelicola LUTZ, by original designation.

REFERENCE: DE MEILLON & WIRTH, 1979 (key to females).


antiguensis SAUNDERS, 1957: 700 (larva, pupa, male, female; Antigua). Distr.- Antigua.bromelicola (LUTZ), 1914: 84 (Ceratopogon; male, female; Brazil). Distr.- USA (Florida),

Trinidad, French Guiana, Brazil (Rio de Janeiro).caribbeana SAUNDERS, 1957: 696 (larva, pupa, male, female; Trinidad and Tobago). Distr.-

Trinidad and Tobago, Guyana.dominicana DE MEILLON & WIRTH, 1979: 195 (larva, pupa, male, female; Dominica). Distr.-

Puerto Rico, Dominica.edwardsi (SAUNDERS), 1925: 260 (Apelma; larva, pupa, male, female; Brazil). Distr.- Trinidad,

Brazil (Pernambuco, Rio de Janeiro).jocosa SAUNDERS, 1957: 701 (male, female; Trinidad). Distr.- Trinidad.keilini (SAUNDERS), 1925: 265 (Apelma; larva, pupa, male; Brazil). Distr.- Brazil (Pernambuco).magna (SAUNDERS), 1925: 266 (Apelma; larva, pupa, male, female; Brazil). Distr.- Brazil

(Pernambuco).musae CLASTRIER & DELÉCOLLE, 1994: 51 (male, female; French Guiana). Distr.- French Guiana.oligarthra SAUNDERS, 1957: 698 (larva, pupa, male, female; Puerto Rico). Distr.- USA (Florida,

Puerto Rico), Cuba, Dominica, Mexico, Guyana, also from Singapore and Micronesia.

Subgenus PTEROBOSCA MACFIE

PTEROBOSCA MACFIE, 1932: 266. Unavailable name; proposed after 1930 without typespecies designation.

PTEROBOSCA MACFIE, 1940f: 16. Type species: Ceratopogon aeschnosuga DE MEIJERE, byoriginal designation.

farri WIRTH, 1966: 29 (female; Jamaica). Distr.- Jamaica.fusicornis (COQUILLETT), 1905: 63 (Ceratopogon; female; USA, Florida). Distr.- USA (Louisiana

to Florida) and Caribbean to Brazil (Rio de Janeiro).fur (JOHNSON), 1913: 444 (Ceratopogon). Bermuda.macfiei (COSTA LIMA), 1937b: 616 (Pterobosca). Brazil.

incubans (MACFIE), 1937b: 111 (Pterobosca; female; Belize). Distr.- Mexico (San Luis Potosi,Veracruz), Puerto Rico to Argentina (Buenos Aires).

Subgenus RHYNCHOFORCIPOMYIA WIRTH & DOW

RHYNCHOFORCIPOMYIA WIRTH & DOW, 1972: 863 (as subgenus of Forcipomyia).Type species: Forcipomyia messersmithi WIRTH & DOW, by original designation.

REFERENCE: WIRTH & DOW, 1972 (key to females).

brachyrhyncha WIRTH & DOW, 1972: 867 (male, female; Colombia). Distr.- Colombia,Trinidad.

cylindrica WIRTH & DOW, 1972: 865 (male, female; El Salvador). Distr.- El Salvador.dorsalis WIRTH & DOW, 1972: 870 (male, female; Mexico). Distr.- Mexico (Sinaloa, Veracruz),

Honduras, El Salvador.


guamai WIRTH & DOW, 1972: 869 (female; Brazil). Distr.- Panama, Brazil (Pará).messersmithi WIRTH & DOW, 1972: 864 (male, female; El Salvador). Distr.- Mexico

(Veracruz), Central America to Colombia, Jamaica.puracensis WIRTH & DOW, 1972: 867 (female; Colombia). Distr.- Colombia.zeteki WIRTH & DOW, 1972: 870 (female; Panama). Distr.- Panama, Colombia.

Subgenus SALIOHELEA WIRTH & RATANAWORABHAN

SALIOHELEA WIRTH & RATANAWORABHAN, 1978: 494 (as subgenus of Forcipomyia). Typespecies: Forcipomyia leei WIRTH & RATANAWORABHAN, by original designation.

leei WIRTH & RATANAWORABHAN, 1978: 498 (male, female; Colombia). Distr.- Eastern USAand Caribbean to southern Brazil (Amazonas, Pará, Bahia, Rio de Janeiro).

Subgenus SCHIZOFORCIPOMYIA CHAN & LEROUX

SCHIZOFORCIPOMYIA CHAN & LEROUX, 1971: 271 (as subgenus of Forcipomyia). Typespecies: Forcipomyia petersoni CHAN & LEROUX ( = Forcipomyia borbonica CLASTRIER), byoriginal designation.

harpa SPINELLI & BORKENT, 2004b: 3 (male, female; Costa Rica). Distr.- Costa Rica.

Subgenus SYNTHYRIDOMYIA SAUNDERS

SYNTHYRIDOMYIA SAUNDERS, 1957: 688 (as subgenus of Forcipomyia). Type species:Lasiohelea acidicola TOKUNAGA, by original designation.

floridensis DOW & WIRTH, 1972: 195 (male, female; USA, Florida). Distr.- USA (Florida),Mexico (Yucatán).

sanctaeclarae WIRTH, 1952b: 90 (male, female; Juan Fernández Islands). Distr.- JuanFernández Islands, central Chile, northwestern Argentina.

soibelzoni MARINO & SPINELLI, 2001c: 14 (female; Argentina). Distr.- Argentina (Rio Negro).tenuiforceps MACFIE, 1939b: 167 (male; Brazil). Distr.- USA (California, New Mexico) to

Brazil (Santa Catarina).

Subgenus THYRIDOMYIA SAUNDERS

THYRIDOMYIA SAUNDERS, 1925: 268. Type species: Thyridomyia palustris SAUNDERS

(= Ceratopogon monilicornis COQUILLETT), by original designation.

REFERENCE: WIRTH, 1970 (key to males and females).

calchaqui SPINELLI & MARINO, 1997: 188 (female; Argentina). Distr.- NorthwesternArgentina.


colombiae WIRTH, 1970: 435 (male, female; Colombia). Distr.- Mexico (Veracruz), CostaRica, Panama, Trinidad, Colombia.

jamaicensis WIRTH, 1970: 436 (male, female; Jamaica). Distr.- Jamaica, Puerto Rico.jipajapae WIRTH, 1970: 437 (male, female; Panama). Distr.- Costa Rica to Ecuador, Trinidad.nana (MACFIE), 1939b: 171 (Lasiohelea; female; Brazil). Distr.- Mexico (Veracruz), Jamaica

through Caribbean to southern Brazil (Santa Catarina).nodosa SAUNDERS, 1959: 43 (larva, pupa, male, female; Costa Rica). Distr.- USA (Arizona,

Kansas, Mississippi), Mexico (Baja California, Sonora, Sinaloa, San Luis Potosi), CostaRica, Colombia.

riojana SPINELLI & MARINO, 1997: 188 (male; Argentina). Distr.- Western and southern Argentina.sinuosa DOW & WIRTH, 1972: 189 (male, female; Mexico). Distr.- Mexico (Sinaloa) (possibly

Iowa in USA).univesicula MACFIE, 1939b: 170 (female; Brazil). Distr.- Brazil (Santa Catarina).

Subgenus TRICHOHELEA GOETGHEBUER

APELMA KIEFFER, 1919a: 64 (preoccupied by BILLBERG, 1820). Type species: Apelma auronitensKIEFFER (= Trichohelea tonnoiri GOETGHEBUER), designation by MACFIE, 1940f: 16.

TRICHOHELEA GOETGHEBUER, 1920: 39. Type species: Trichohelea tonnoiri GOETGHEBUER,by original designation.

REFERENCE: MARINO & SPINELLI, 2004b (key to males and females).

aeronautica MACFIE, 1935b: 265 (female; Guyana). Distr.- Venezuela to French Guiana.baueri WIRTH, 1956a: 361 (female; USA, Arizona). Distr.- USA (Florida, Maryland, Arizona),

Mexico (Baja California, Sonora, Sinaloa, Veracruz).danaisi (FLOCH & ABONNENC), 1949: 2 (1950a: 72) (Lasiohelea; female; Venezuela). Distr.-

Venezuela.intrepida MACFIE, 1936: 228 (female; Peru). Distr.- Peru.limnetis INGRAM & MACFIE, 1931: 169 (male; Argentina). Distr.- Southwestern Argentina.

shannoni (INGRAM & MACFIE), 1931: 171 (Lasiohelea). Argentina.macheti CLASTRIER & LEGRAND, 1990: 168 (female; French Guiana). Distr.- French Guiana.opilionivora (LANE), 1947c: 159 (Lasiohelea; female; Brazil). Distr.- Brazil (São Paulo).roubaudi CLASTRIER & DELÉCOLLE, 1997: 379 (female; French Guiana). Distr.- French Guiana.sayhuequei MARINO & SPINELLI, 2004b: 2257 (female; Argentina). Distr.- Southwestern

Argentina, southern Chile.tehuelche MARINO & SPINELLI, 2004b: 2260 (male, female; Argentina). Distr.- Southwestern

Argentina.trinidadensis SAUNDERS, 1964: 476 (pupa, female; Trinidad). Distr.- Trinidad.

Subgenus WARMKEA SAUNDERS

WARMKEA SAUNDERS, 1957: 671 (as subgenus of Forcipomyia). Type species: Forcipomyiabicolor SAUNDERS (= Forcipomyia lesliei WIRTH), by original designation.


REFERENCE: WIRTH & SORIA, 1980 (key to males and females).

aeria SAUNDERS, 1957: 675 (larva, pupa, male, female; Puerto Rico). Distr.- Puerto Rico,Colombia, Argentina (Tucuman).

galindoi WIRTH & SORIA, 1980: 146 (male, female; Panama). Distr.- Panama, Colombia,Ecuador, Brazil (Pará).

lesliei WIRTH, 1974: 12. New name for bicolor. Distr.- Jamaica, Haiti to Venezuela, Colombia.bicolor SAUNDERS, 1957: 672 (preoccupied by Forcipomyia bicolor LUTZ, 1914). Puerto Rico.

louriei (MACFIE), 1935a: 49 (Lasiohelea; female; Brazil). Distr.- Colombia, Bolivia, Trinidad,Brazil (Pará, Maranhão, Santa Catarina).

narthekophora MACFIE, 1939b: 166. Brazil.spinosa SAUNDERS, 1957: 676 (larva, pupa, male, female; Puerto Rico). Distr.- Puerto Rico,

Dominica, Trinidad to Colombia, Guyana, and Brazil (Pará, Bahia, Rio de Janeiro).terrestris SAUNDERS, 1964: 479 (larva, pupa, male, female; Trinidad). Distr.- Trinidad.tuberculata SAUNDERS, 1957: 677 (larva, pupa, male, female; Trinidad and Tobago). Distr.-

Costa Rica, Colombia, Trinidad and Tobago, Brazil (Amazonas, Pará), Argentina(Buenos Aires).

SUBFAMILY DASYHELEINAE LENZ, 1934: 96

Genus DASYHELEA KIEFFER

DASYHELEA KIEFFER, 1911c: 5. Type species: Dasyhelea halophila KIEFFER, by monotypy.PSEUDOCULICOIDES MALLOCH, 1915a: 309. Type species: Ceratopogon mutabilis

COQUILLETT, by original designation.

REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE,1931 (key males and females from Patagonia and southern Chile); SPINELLI & WIRTH,1984d (key to males and females of cincta group).

aegealitis SPINELLI & WIRTH, 1984d: 589 (male, female; Jamaica). Distr.- USA (Florida), Jamaica.albopicta INGRAM & MACFIE, 1931: 187 (male, female; Argentina). Distr.- Southern Chile,

southwestern Argentina.ancora (COQUILLETT), 1902: 87 (Ceratopogon; female; USA, Florida). Distr.- USA (California,

Arizona, New Mexico, Connecticut to Florida), Galápagos Islands.andensis INGRAM & MACFIE, 1931: 180 (male; Argentina). Distr.- Southwestern Argentina.argentinensis KIEFFER, 1925: 92 (male; Argentina). Distr.- Argentina (Cordoba).atlantis WIRTH & WILLIAMS, 1957: 11 (male, female; Bermuda). Distr.- Bermuda, USA

(New York, Florida), Grand Cayman, Galápagos Islands.australis WIRTH, 1952b: 92 (male, female; Juan Fernandez Islands). Distr.- Juan Fernandez Islands.bahamensis (JOHNSON), 1908: 71 (Ceratopogon; female; Bahamas). Distr.- USA (Florida,

Puerto Rico), Mexico (Yucatán), Bahamas, Cuba, Jamaica.bajensis WIRTH, 1978: 192 (male, female; Mexico). Distr.- USA (California), Mexico (Baja

California, Sinaloa).


bermudae WIRTH & WILLIAMS, 1957: 11 (male, female; Bermuda). Distr.- Bermuda.borgmeieri WIRTH & WAUGH, 1976: 225 (male, female; Trinidad). Distr.- Panama, Grenada,

Trinidad, Galápagos Islands, Brazil (Bahia).cacaoi WIRTH & WAUGH, 1976: 231 (male, female; Trinidad). Distr.- Panama, Trinidad.calvescens MACFIE, 1938: 157 (male, female; USA, Hawaii). Distr.- Mexico (Baja California,

Sonora), Panama, Hawaii.caribbeana SPINELLI & WIRTH, 1984d: 596 (female; Jamaica). Distr.- Mexico (Quintana

Roo, Chiapas), Panama, Jamaica, Haiti.chilensis INGRAM & MACFIE, 1931: 186 (female; Chile). Distr.- Southern Chilecincta (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Florida). Distr.- USA (California

to New York to Florida) to Brazil and Argentina.penthesileae MACFIE, 1935a: 55. Brazil.

columbiana (KIEFFER), 1917b: 304 (Culicoides; female; Colombia). Distr.- Colombia.correntina RONDEROS & DÍAZ, in RONDEROS et al., 2004a: 194 (male, female, pupa; Argentina).

Distr.- Northeastern Argentina.filibranchia (LUTZ), 1914: 85 (Ceratopogon; pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro).flavicauda MACFIE, 1939b: 201 (female; Brazil). Distr.- Northeastern Argentina, Brazil

(Santa Catarina).griseola WIRTH, 1978: 193 (male, female; Mexico). Distr.- Mexico (Baja California), Panama,

Trinidad.guadeloupensis DELÉCOLLE & RIEB, 1994: 272 (male, female; Guadeloupe). Distr.-

Guadeloupe.hippolytae MACFIE, 1935a: 54 (female; Brazil). Distr.- Brazil (Maranhão).hirtipes (KIEFFER), 1917b: 305 (Culicoides; male; Peru). Distr.- Peru.hondurensis SPINELLI & WIRTH, 1984d: 600 (male, female; Belize). Distr.- Belize.jamaicensis SPINELLI & WIRTH, 1984d: 602 (female; Jamaica). Distr.- Jamaica.koenigi DELÉCOLLE & RIEB, 1994: 267 (male, female; Guadeloupe). Distr.- Guadeloupe.lacustris INGRAM & MACFIE, 1931: 190 (female; Argentina). Distr.- Western Argentina.luteogrisea WIRTH & WILLIAMS, 1957: 10 (female; Bermuda). Distr.- USA (Florida to Texas),

Bermuda, Bahamas.maculata MACFIE, 1943: 119 (male, female; Bahamas). Distr.- USA (Florida), Mexico

(Quintana Roo) to Panama, Bahamas, Jamaica to Trinidad.mediomunda MINAYA, 1978: 79 (pupa, male, female; Peru). Distr.- Peru, Chile.monticola INGRAM & MACFIE, 1931: 188 (male, female; Chile). Distr.- Southern Chile.morrisoni Grogan and Wieners, 2006: 468 (male, female; Bahamas). Distr.- Bahamas.mutabilis (COQUILLETT), 1901a: 602 (Ceratopogon; male, female; USA, District of Columbia).

Distr.- North America (Alaska to Newfoundland, to California to Florida), GalápagosIslands.

necrophila SPINELLI & RODRÍGUEZ, 1999: 59 (male, female, pupa, larva; Argentina). Distr.-Mexico (Coahuila), Argentina (Buenos Aires).

paracincta WIRTH, 1969: 576 (male, female; Galápagos Islands). Distr.- Galápagos Islands.parahybae MACFIE, 1940a: 74 (male, female; Brazil). Distr.- Northeastern Brazil.patagonica INGRAM & MACFIE, 1931: 182 (male, female; Argentina). Distr.- Southwestern

Argentina.


paulistana FORATTINI & RABELLO, 1957: 245 (pupa, female; Brazil). Distr.- Brazil (São Paulo).philotherma MACFIE, 1953: 103 (male; Costa Rica). Distr.- Costa Rica.pollex BORKENT & FORSTER, 1986: 1286 (male; Bahamas). Distr.- USA (Florida), Bahamas,

Mexico (Baja California).pumila MACFIE, 1939b: 202 (male, female; Brazil). Distr.- Brazil (Santa Catarina).pusilla (LUTZ), 1913: 65 (Centrorhynchus; female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro).pygmaea (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- St. Vincent.pyrsonota MACFIE, 1953: 104 (male; Costa Rica). Distr.- Costa Rica.reynoldsi INGRAM & MACFIE, 1931: 185 (male, female; Chile). Distr.- Chile (Magallanes,

Tierra del Fuego).santaemarthae (KIEFFER), 1917b: 306 (Culicoides; female; Colombia). Distr.- Colombia.scissurae MACFIE, 1937a: 15 (male; Trinidad). Distr.- USA (Florida), Mexico (San Luis

Potosi, Guerrero) to Costa Rica, Bermuda, Trinidad to northeastern Argentina.shannoni INGRAM & MACFIE, 1931: 183 (male, female; Chile). Distr.- Southern Chile.sinclairi BORKENT, 1991: 110 (male, female; Galápagos Islands). Distr.- Galápagos Islands.soriai WIRTH & WAUGH, 1976: 229 (male, female; Brazil). Distr.- Brazil (Bahia).spathicerca WIRTH, 1969: 577 (male, female; Galápagos Islands). Distr.- Galápagos Islands.suarezi SPINELLI & RONDEROS, 1987: 11 (male, female; Argentina). Distr.- Argentina (Buenos

Aires).thalestris MACFIE, 1935a: 55 (female; Brazil). Distr.- Trinidad, Guyana, Brazil (Maranhão).villosipes (KIEFFER), 1917b: 305 (Culicoides; male; Paraguay). Distr.- Paraguay.viridans FORATTINI & RABELLO, 1957: 247 (male; Brazil). Distr.- Brazil (Rio de Janeiro).williamsi WIRTH & WAUGH, 1976: 228 (male, female; Trinidad). Distr.- Panama, Dominica,

Trinidad, Brazil (Bahia).winderi WIRTH & WAUGH, 1976: 233 (male, female; Brazil). Distr.- Brazil (Bahia).

SUBFAMILY CERATOPOGONINAE NEWMAN, 1834: 388

TRIBE CULICOIDINI KIEFFER, 1911c: 1, 1911b: 319

Genus CULICOIDES LATREILLE

CULICOIDES LATREILLE, 1809: 251. Type species: Culicoides punctatus LATREILLE

(= Ceratopogon punctatus MEIGEN), by monotypy.

PADROSIA RAFINESQUE, 1815: 130 (unnecessary replacement name for Culicoides LATREILLE).Type species: Culicoides punctatus LATREILLE (= Ceratopogon punctatus MEIGEN), automatic.

REFERENCES: AITKEN et al., 1975 (key to females from Trinidad and Tobago); BLANTON

& WIRTH, 1979 (key to larvae, males and females from Florida (USA)); FORATTINI,1957 (key to females of all species); HUERTA, 1996 (key to females from Mexico);ORTÍZ & LEÓN, 1955 (key to females from Ecuador); RONDEROS, 1988 (key to femalesfrom Argentina); RONDEROS & SPINELLI, 1998 (key to males and females from theYacyretá Dam, between Paraguay and Argentina); SPINELLI, RONDEROS, DÍAZ & MARINO,2005 (key to males and females from Argentina); SPINELLI & WIRTH, 1986b (key to


females south of the Amazon Basin); WIRTH & BLANTON, 1959 (key to females andsome males from Panama); WIRTH & BLANTON 1973 (key to females and some malesfrom Amazon Basin); WIRTH & BLANTON, 1974b (key to females and some malesfrom the Caribbean); WIRTH, DYCE & SPINELLI, 1988 (some meristic character statesand photographs of wings of all species).

There is no reliable key to the subgenera and species groups of Neotropical Culicoides. Forspecies outside of local areas for which revisions are available (as listed above), theeasiest guide to species identification is that of WIRTH, DYCE & SPINELLI (1988). Fromthere readers may refer to keys to subgenera or species groups referred to below.

Subgenus AMOSSOVIA GLUKHOVA

AMOSSOVIA GLUKHOVA, 1989: 226 (as subgenus of Culicoides). Type species: Ceratopogondendrophilus Amosova, by original designation.

REFERENCE: WIRTH & BLANTON, 1967 (key to males and females)

oklahomensis KHALAF, 1952: 355 (as subspecies of villosipennis; male; USA, Oklahoma).Distr.- USA (California to Mississippi) to Guatemala.

Subgenus ANILOMYIA VARGAS

ANILOMYIA VARGAS, 1960: 37 (as subgenus of Culicoides). Type species: Culicoides covagarciaiORTÍZ, by original designation.

REFERENCES: WIRTH & BLANTON, 1956b (key to males and females of covagarciai group);WIRTH & BLANTON, 1970d (key to key to males and females of nigrigenus group).

ameliae BROWNE, 1980: 543 (female; Colombia). Distr.- Colombia.chaverrii SPINELLI & BORKENT, 2004a: 373 (larva, pupa, male, female; Costa Rica). Distr.-

Costa Rica.chrysonotus WIRTH & BLANTON, 1956b: 226 (male, female; Panama). Distr.- El Salvador,

Costa Rica, Panama.covagarciai ORTÍZ, 1950b: 457 (male, female; Venezuela). Distr.- Honduras to Colombia,

Venezuela.beebei FOX, 1952: 366. Venezuela.

decor (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- Dominica, St. Lucia,St. Vincent.

dominicanus WIRTH & BLANTON, 1970d: 146 (male, female; Dominica). Distr.- Dominica.efferus FOX, 1952: 365 (female; Peru). Distr.- Guatemala to Peru and Bolivia.farri WIRTH & BLANTON, 1970d: 148 (male, female; Jamaica). Distr.- Jamaica.hayesi MATTA, 1967: 75 (larva, pupa, male, female; Honduras). Distr.- Mexico (Tamaulipas),

Honduras.


lutealaris WIRTH & BLANTON, 1956b: 225 (male, female; Panama). Distr.- Costa Rica, Panama.marshi WIRTH & BLANTON, 1956b: 220 (female; Panama). Distr.- Costa Rica, Panama, Colombia.metagonatus WIRTH & BLANTON, 1956b: 221 (male, female, Panama). Distr.- Nicaragua to Ecuador.monicae SPINELLI & BORKENT, 2004a: 372 (female; Costa Rica). Distr.- Costa Rica.nigrigenus WIRTH & BLANTON, 1956b: 222 (male, female; Panama). Distr.- Mexico

(Veracruz) to Colombia, Trinidad, Argentina (Salta).popayanensis WIRTH & LEE, 1967: 10 (male, female; Colombia). Distr.- Colombia.rostratus WIRTH & BLANTON, 1956b: 218 (male, female; Panama). Distr.- Panama.trapidoi WIRTH & BARRETO, 1978: 554 (male, female; Colombia). Distr.- Costa Rica,

Colombia, Brazil.

Subgenus AVARITIA FOX

AVARITIA FOX, 1955: 218 (as subgenus of Culicoides). Type species: Ceratopogon obsoletusMEIGEN, by original designation.

REFERENCES: RODRÍGUEZ & WIRTH, 1986 (revision of andicola group); WIRTH & MULLENS,1992 (key to males and females in pusillus group).

andicola WIRTH & LEE, 1967: 5 (female; Colombia). Distr.- Colombia.hermani SPINELLI & BORKENT, 2004a: 363 (male, female; Panama). Distri. - Costa Rica, Panama.impusilloides SPINELLI & WIRTH, 1984a: 178 (male, female; Brazil). Distr.- Brazil (Santa

Catarina), Uruguay.orjuelai WIRTH & LEE, 1967: 6 (female; Colombia). Distr.- Colombia.puracensis WIRTH & LEE, 1967: 7 (male, female; Colombia). Distr.- Colombia.pusilloides WIRTH & BLANTON, 1955a: 104 (male, female; Panama). Distr.- Guatemala and

Belize to Panama.pusillus LUTZ, 1913: 52 (male, female; Brazil). Distr.- USA (Florida), Mexico (Chiapas) to

northeastern Argentina.suarezi RODRÍGUEZ & WIRTH, 1986: 313 (female; Colombia). Distr.- Colombia.

Subgenus BELTRANMYIA VARGAS

BELTRANMYIA VARGAS, 1953a: 34 (as subgenus of Culicoides). Type species: Culicoidescrepuscularis MALLOCH, by original designation.

REFERENCE: BLANTON & WIRTH, 1979 (males and females included in key to speciesfrom Florida).

bermudensis WILLIAMS, 1956: 298 (female; Bermuda). Distr.- USA (New York to Floridaand Texas), Bermuda.

crepuscularis MALLOCH, 1915a: 303 (male, female; USA, Illinois). Distr.- Canada (BritishColumbia to Nova Scotia) to Costa Rica.

knowltoni BECK, 1956: 136 (male, female; USA, Florida). Distr.- Bahamas, USA (Florida).


Subgenus COTOCRIPUS BRÈTHES

COTOCRIPUS BRÈTHES, 1912: 451. Type species: Cotocripus caridei BRÈTHES, by monotypy.

bambusicola LUTZ, 1913: 62 (female; Brazil). Distr.- Colombia, Venezuela, Brazil (Bahia,Espírito Santo, Rio de Janeiro, São Paulo), Argentina (Misiones, Buenos Aires).

bahiensis BARBOSA, 1947: 11. Brazil.caridei (BRÈTHES), 1912: 451 (Cotocripus; female; Argentina). Distr.- Argentina (Buenos Aires

province), southeastern Brazil, Uruguay.setifer (LUTZ), 1913: 64 (Centrorhynchus). Brazil.

irwini SPINELLI & WIRTH, 1984a: 180 (female; Chile). Distr.- Chile, southwestern Argentina.patagoniensis RONDEROS & SPINELLI, 1997: 34 (male, female; Argentina). Distr.- Southern

Chile, southern Argentina.raposoensis WIRTH & BARRETO, 1978: 561 (male, female; Colombia). Distr.- Colombia.

Subgenus CULICOIDES LATREILLE

elutus MACFIE, 1948: 75 (female; Mexico). Distr.- Mexico (Chiapas, Oaxaca) to Panama.hondurensis SPINELLI & BORKENT, 2004a: 369 (male, female; Honduras). Distr.- El Salvador,

Honduras.luteovenus ROOT & HOFFMAN, 1937: 156 (male, female; Mexico). Distr.- USA (California,

Utah, Washington) to Panama.neopulicaris WIRTH, 1955: 355 (male, female; USA,Texas). Distr.- USA (Texas and

Louisiana) to Costa Rica.

Subgenus DIPHAOMYIA VARGAS

DIPHAOMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides baueriHOFFMAN, by original designation.

blantoni VARGAS & WIRTH, 1955: 33 (male, female; Mexico). Distr.- USA (Texas), Mexico(Sinaloa, Tamaulipas, Morelos).

edeni WIRTH & BLANTON, 1974a: 23 (male, female; notes, syn.; USA, Florida). Distr.-Southeastern USA to Bahamas.

evansi WIRTH & BLANTON, 1959: 342 (male, female; Panama). Distr.- Honduras, CostaRica, Panama.

freitasi WIRTH & BLANTON, 1973: 434 (female; Brazil). Distr.- Brazil (Pará).haematopotus MALLOCH, 1915a: 302 (male, female; USA, Illinois). Distr.- North America

(British Columbia to Nova Scotia, entire USA) to Honduras.iriartei FOX, 1952: 368 (female; Venezuela). Distr.- Guatemala to Colombia, Venezuela,

Tobagao, Brazil (Pará).vargasi WIRTH & BLANTON, 1953a: 74. Panama.

jurbergi FELIPPE-BAUER, in FELIPPE-BAUER et al., 2005: 51 (female; Peru). Distr.- Peru.marinkellei WIRTH & LEE, 1967: 13 (male, female; Colombia). Distr.- Colombia.


mckeeveri BRICKLE & HAGAN, 1999: 39 (female; Belize). Distr.- Belize.minasensis FELIPPE-BAUER, 1987: 147 (female; Brazil). Distr.- Brazil (Minas Gerais).mirsae ORTÍZ, 1953a: 801 (female; Venezuela). Distr.- Panama, Venezuela, Trinidad.ronderosae SPINELLI & BORKENT, 2004a: 377 (female; Costa Rica). Distr.- Costa Rica.tarapaca SPINELLI & WIRTH, 1984a: 184 (female; Chile). Distr.- Northern Chile.

Subgenus DRYMODESMYIA VARGAS

DRYMODESMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoidescopiosus ROOT & HOFFMAN, by original designation.

REFERENCE: WIRTH & HUBERT, 1960: 646 (key to females and some males; as copiosus group).

borinqueni FOX & HOFFMAN, 1944: 110 (male, female; Puerto Rico). Distr.- Jamaica to St. Croix.bredini WIRTH & BLANTON, 1970b: 41 (male, female; Dominica). Distr.- Dominica.chacoensis SPINELLI & WIRTH, 1984a: 174 (male, female; Argentina). Distr.- Bolivia, Paraguay,

Argentina, Uruguay.haitiensis DELÉCOLLE, RACCURT & REBHOLTZ, 1986: 108 (male, female; Haiti). Distr.- Haiti.hitchcocki SPINELLI & WIRTH, 1984a: 176 (female; Peru). Distr.- Peru.jamaicensis EDWARDS, 1922: 165 (as variety of loughnani; female; Jamaica). Distr.- USA (Texas,

Florida), Mexico (Yucatán), Central America and Caribbean to Colombia and Venezuela.loughnani EDWARDS, 1922: 165 (female; Jamaica). Distr.- USA (Texas to Georgia), Mexico

(Yucatán), Bahamas, Jamaica, Cuba, Australia (as introduction with Opuntia).panamensis BARBOSA, 1947: 22 (male, female; Panama). Distr.- Mexico (Nayarit, Veracruz,

Chiapas) to Costa Rica, Jamaica.alambiculorum MACFIE, 1948: 81. Mexico.

pilosus WIRTH & BLANTON, 1959: 332 (male, female; Panama). Distr.- Costa Rica, Panama,Brazil (Pará).

poikilonotus MACFIE, 1948: 82 (female; Mexico). Distr.- Mexico (Chiapas), Central America,Venezuela and Trinidad to Brazil (Bahia).

cacozelus MACFIE, 1948: 85. Mexico.hertigi WIRTH & BLANTON, 1953b: 229. Panama.

saltaensis SPINELLI & WIRTH, 1984a: 183 (female; Argentina). Distr.- Western Argentina.uruguayensis RONDEROS, 1990a: 117 (female; Uruguay). Distr.- Uruguay.wirthomyia VARGAS, 1953b: 227 (male; Mexico). Distr.- Mexico (Guerrero).

Subgenus GLAPHIROMYIA VARGAS

GLAPHIROMYIA VARGAS, 1960: 41 (as subgenus of Culicoides). Type species: Culicoidesscopus ROOT & HOFFMAN, by original designation.

parascopus WIRTH & BLANTON, 1978: 238 (male, female; Mexico). Distr.- Mexico (Michoacan).scopus ROOT & HOFFMAN, 1937: 170 (male, female; Mexico). Distr.- Mexico (DF), Costa

Rica, Panama.


Subgenus HAEMATOMYIDIUM GOELDI

HAEMATOMYIDIUM GOELDI, 1905: 137. Type species: Haematomyidium paraense GOELDI,by original designation.

REFERENCES: VITALE, WIRTH & AITKEN, 1981 (key to females of debilipalpis group);WIRTH & FELIPPE-BAUER, 1989 (key to paraensis group).

annuliductus WIRTH, in VITALE et al, 1981: 150 (pupa, male, female; Panama). Distr.- CostaRica, Panama.

austroparaensis SPINELLI, in SPINELLI et al., 2005b: 141 (male, female; Argentina). Distr.-northeastern Argentina.

aragaoi TAVARES & LUNA DIAS, 1980: 393 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).bachmanni SPINELLI, in SPINELLI et al, 2005b: 146 (female; Argentina). Distr.- Paraguay,

northeastern Argentina.bayano WIRTH, in VITALE et al, 1981: 152 (pupa, male, female; Panama). Distr.- Costa Rica,

Panama.crucifer CLASTRIER, 1968: 85 (male, female; French Guiana). Distr.- Trinidad, Guyana, French

Guiana.darlingtonae WIRTH & BLANTON, 1971: 39 (male, female; Trinidad). Distr.- Costa Rica, Trinidad.debilipalpis LUTZ, 1913: 60 (female; Brazil). Distr.- Widespread from USA (Maryland,

Kentucky, Nebraska south to Lousiana and Florida), Guatemala and Belize to Argentina.ichesi RONDEROS & SPINELLI, 1995b: 77. Argentina.

denisae CLASTRIER, 1971: 290 (male, female; French Guiana). Distr.- French Guiana, Brazil(Amazonas).

diversus FELIPPE-BAUER, in FELIPPE-BAUER et al. 2003: 1052 (female; Peru). Distr.- Peru.dureti RONDEROS & SPINELLI, 1995a: 59 (female; Paraguay). Distr.- Northeastern Argentina,

Paraguay.eadsi WIRTH & BLANTON, 1971: 37 (male, female; USA, Texas). Distr.- USA (Florida, Texas),

Cuba, Mexico (Nayarit, Sonora, San Luis Potosi, Yucatán), Guatemala.eldridgei WIRTH & BARRETO, 1978: 561 (male, female; Colombia). Distr.- Colombia.equatoriensis BARBOSA, 1952: 13 (as variety of debilipalpis; female; Ecuador). Distr.- Ecuador.espinolai FELIPPE-BAUER & LOURENCO-DE-OLIVEIRA, 1987: 149 (female; Brazil). Distr.- Brazil

(Minas Gerais).filiductus WIRTH, in VITALE et al., 1981: 155 (pupa, male, female; Panama). Distr.- Belize to

Panama.flinti WIRTH, 1982b: 251 (female; Argentina). Distr.- Northeastern Argentina, southeastern

Brazil, Uruguay.germanus MACFIE, 1940d: 27 (female; Guyana). Distr.- Costa Rica, Guyana.ginesi ORTÍZ, 1951c: 586 (female; Venezuela). Distr.- El Salvador to Panama, Colombia,

Venezuela, Trinidad, Brazil (Pará), northeastern Argentina.glabrior MACFIE, 1940d: 27 (as variety of debilipalpis; female; Guyana). Distr.- Honduras to

Ecuador, Guyana, Surinam, Trinidad, Brazil (Pará).grahambelli FORATTINI, 1956b: 35. Panama.


hoffmani FOX, 1946: 251 (female; Trinidad). Distr.- Belize, Costa Rica, Cayman Islands,Jamaica to Trinidad.

horticola LUTZ, 1913: 61 (female; Brazil). Distr.- Paraguay, eastern Argentina, southeasternBrazil.

imitator ORTÍZ, 1953b: 807 (male, female; Venezuela). Distr.- Guatemala to Panama, Venezuela.insinuatus ORTÍZ & LEÓN, 1955: 577 (female; Ecuador). Distr.- Colombia, Ecuador, Peru,

Trinidad, Guyana, Brazil (Amazonas, Pará, (?) São Paulo).kampa FELIPPE-BAUER, VERAS, CASTELLON & MOREIRA, 2000: 35 (female; Brazil). Distr.-

Brazil (Acre).lahillei (ICHES), 1906: 264 (Ceratopogon, female; Argentina). Distr.- Northeastern Argentina,

southeastern Brazil, Paraguay.limonensis ORTÍZ & LEÓN, 1955: 576 (female; Ecuador). Distr.- Ecuador, Venezuela, Brazil

(Pará, São Paulo, Santa Catarina).neoparaensis TAVARES & SOUZA, 1978: 621 (male, female; Brazil). Distr.- Ecuador, Brazil

(Amazonas, Rio de Janeiro, Santa Catarina).pampaensis SPINELLI & WIRTH, 1984a: 182 (female; Argentina). Distr.- Western Argentina.paraensis (GOELDI), 1905: 137 (Haematomyidium; female; Brazil). Distr.- USA (Colorado,

Nebraska, Pennsylvania, Wisconsin to Louisiana and Florida) to Argentina.undecimpunctatus KIEFFER, 1917b: 307. Argentina.

peruvianus FELIPPE-BAUER, in FELIPPE-BAUER et al., 2003: 1054 (female; Peru). Distr.- Peru.quasiparaensis CLASTRIER, 1971: 286 (male, female; French Guiana). Distr.- Honduras

and El Salvador to Colombia, Peru, French Guiana, Brazil (Amazonas, Rondônia).rachoui TAVARES & SOUZA, 1978: 622 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).spurius WIRTH & BLANTON, 1959: 433 (male, female; Panama). Distr.- Costa Rica, Panama.todatangae WIRTH & BLANTON, 1973: 447 (male, female; Brazil). Distr.- Trinidad, Brazil

(Pará, Santa Catarina).youngi WIRTH & BARRETO, 1978: 562 (male, female; Colombia). Distr.- Colombia.

Subgenus HOFFMANIA FOX

HOFFMANIA FOX, 1948: 21 (as subgenus of Culicoides). Type species: Culicoides inamollaeFOX & HOFFMAN (= Culicoides insignis LUTZ), by original designation.

REFERENCES: SPINELLI, GREINER & WIRTH, 1993 (key to species groups; key to malesand females of guttatus group); WIRTH & BLANTON, 1968a (key to males and females inhylas group).

aitkeni WIRTH & BLANTON, 1968a: 214 (female; Trinidad). Distr.- Trinidad, Brazil (Amazonas).annettae SPINELLI & BORKENT, 2004a: 365 (male, female; Costa Rica). Distr.- Costa Rica.batesi WIRTH & BLANTON, 1973: 426 (male, female; Brazil). Distr.- Guatemala, Colombia,

Ecuador, Bolivia, Brazil (Pará).sanmartini WIRTH & BARRETO, 1978: 553. Colombia.

biestroi SPINELLI & RONDEROS, 1991: 86 (male, female; Argentina). Distr.- Eastern Argentina,Uruguay.


bimaculatus FLOCH & ABONNENC, 1942b: 3 (female; French Guiana). Distr.- French Guiana,Brazil (Pará).

brasilianum FORATTINI, 1956a: 81 (female; Brazil). Distr.- Northeastern Argentina,southeastern Brazil.

brownei SPINELLI, in SPINELLI et al., 1993: 24 (male, female; Colombia). Distr.- Colombia.charruus SPINELLI & Martinez, 1992: 176 (as charrua; pupa, male, female; Uruguay). Distr.-

Paraguay, northeastern Argentina, Uruguay.coutinhoi BARRETTO, 1944: 96 (male; Brazil). Distr.- French Guiana, Brazil (Amazonas,

Pará, São Paulo).davidi SPINELLI, in SPINELLI et al., 1993: 30 (female; Colombia). Distr.- Costa Rica, Colombia,

Trinidad.diabolicus HOFFMAN, 1925: 294 (female; Panama). Distr.- Mexico to Venezuela and Ecuador.diffusus SPINELLI, in SPINELLI et al., 1993: 34 (female; Brazil). Distr.- Brazil (Espirito Santo).fernandoi TAVARES & SOUZA, 1979: 611 (male, female; Brazil). Distr.- Colombia, Brazil

(Espírito Santo, Rio de Janeiro, Santa Catarina), northeastern Argentina, Uruguay.ferreyrai RONDEROS & SPINELLI, 1995a: 61 (female; Argentina). Distr.- Northeastern Argentina.filarifer HOFFMAN, 1939: 172 (female; Mexico). Distr.- Mexico (Veracruz, Chiapas) to

northern Brazil.flavivenulus COSTA LIMA, 1937a: 418 (as flavivenula; female; Brazil). Distr.- Trinidad, French

Guiana, Brazil (Pará, Pernambuco, Mato Grosso, Espírito Santo, São Paulo, Rio deJaneiro, Santa Catarina).

foxi ORTÍZ, 1950c: 461 (male; Puerto Rico). Distr.- Mexico (Chiapas) to Bolivia, PuertoRico to northeastern Argentina.

franklini SPINELLI, in SPINELLI et al., 1993: 45 (male, female; Panama). Distr.- Mexico(Guerrero) to Bolivia, Brazil (Pará).

fusipalpis WIRTH & BLANTON, 1973: 435 (male, female; Brazil). Distr.- El Salvador to Ecuador,Bolivia, French Guiana, Guyana, Brazil (Amazonas, Pará, Bahia, Rio de Janeiro).

guttatus (COQUILLETT), 1904b: 35 (Ceratopogon; female; Brazil). Distr.- Paraguay, northeasternArgentina, southeastern Brazil.

heliconiae FOX & HOFFMAN, 1944: 108 (male, female; Venezuela). Distr.- Belize to Ecuador,Venezuela, Grenada, Trinidad and Tobago.

rozeboomi BARBOSA, 1947: 26. Trinidad.hylas MACFIE, 1940d: 26 (female; Guyana). Distr.- Mexico (Veracruz) to Peru, Brazil

(Amazonas).ignacioi FORATTINI, 1957: 215 (male, female; Brazil). Distr.- Brazil (Minas Gerais, São Paulo,

Rio de Janeiro), Paraguay.saintjusti TAVARES & RUIZ, 1980: 27. Brazil.

insignis LUTZ, 1913: 51 (pupa, male, female; Brazil). Distr.- USA (Alabama, Georgia, Florida),Mexico (Yucatán, Chiapas), Central America and Caribbean to central Argentina.

inamollae FOX & HOFFMAN, 1944: 110. Puerto Rico.painteri FOX, 1946: 257. Honduras.

lutzi COSTA LIMA, 1937a: 419 (female; Brazil). Distr.- Colombia to French Guiana,northeastern Argentina, Brazil (Roraima, Amazonas, Pará, Mato Grosso, Goiás, SãoPaulo, Rio de Janeiro, Santa Catarina).


maruim LUTZ, 1913: 48 (female; Brazil). Distr.- Venezuela to French Guiana, Trinidad,coastal Brazil.

recifei BARBOSA, 1947: 25. Brazil.ocumarensis ORTÍZ, 1950b: 455 (male, female; Venezuela). Distr.- Mexico (Oaxaca, Tabasco)

to northern Brazil (Pará, Rondônia).palpalis MACFIE, 1948: 78 (female; Mexico). Distr.- Mexico (Chiapas) to Peru, Brazil (Amazonas).paraignacioi SPINELLI, in SPINELLI et al., 1993: 66 (male, female; Colombia). Distr.- Belize

to Colombia, French Guiana, Brazil (Amazonas, Pará).paramaruim WIRTH & BLANTON, 1973: 443 (male, female; Brazil). Distr.- Brazil (Pará).plaumanni SPINELLI, in SPINELLI et al., 1993: 69 (female; Argentina). Distr.- Bolivia, Brazil

(Amazonas) northeastern Argentina.polypori WIRTH & BLANTON, 1968a: 212 (male, female; Panama). Distr.- Costa Rica to

Colombia, Brazil (Amazonas).pseudodiabolicus FOX, 1946: 256 (female; Trinidad). Distr.- Mexico (Veracruz) to Peru

and northern Brazil.ruizi FORATTINI, 1954a: 189 (male, female; Brazil). Distr.- Colombia, Brazil (Amazonas, Goiás).tidwelli SPINELLI, in SPINELLI et al., 1993: 74 (male, female; Colombia). Distr.- Honduras to

Colombia, Ecuador.travassosi FORATTINI, 1957: 198 (male, female; Brazil). Distr.- Surinam, Brazil (Amazonas,

Pará, Mato Grosso).trinidadensis HOFFMAN, 1925: 286 (female; Trinidad and Tobago). Distr.- Coastal; Honduras

and El Salvador to Colombia, Ecuador to Surinam, Cuba and Cayman Islands to Trinidad.oliveri FOX & HOFFMAN, 1944: 108. Haiti.wokei BARBOSA, 1947: 28 (preoccupied by Culicoides wokei FOX, 1947). Panama.diminutus BARBOSA, 1951: 163. New name for wokei.

verecundus MACFIE, 1948: 76 (male, female; Mexico). Distr.- Mexico (Chiapas) to Ecuador.contubernalis ORTÍZ & LEÓN, 1955: 574 (as variety of rozeboomi). Ecuador.

xanifer WIRTH & BLANTON, 1968a: 210 (male, female; Panama). Distr.- Honduras to Panama.

Subgenus MACFIELLA FOX

MACFIELLA FOX, 1955: 217 (as subgenus of Culicoides). Type species: Ceratopogon phlebotomusWILLISTON, by original designation.

phlebotomus (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- Coastal;Mexico (Sinaloa) to Ecuador, Jamaica to Brazil (Maranhão, Ceará, Pernambuco, Goiás).

amazonius MACFIE, 1935a: 52. Brazil.willistoni WIRTH & BLANTON, 1953c: 116 (male, female; Panama). Distr.- Mexico (Sonora),

Honduras, Panama.

Subgenus MATAEMYIA VARGAS

MATAEMYIA VARGAS, 1960: 43 (as subgenus of Culicoides). Type species: Culicoides mojingaensisWIRTH & BLANTON, by original designation.


REFERENCE: WIRTH & SORIA, 1981 (key males and females to some species, as discrepansgroup).

albuquerquei WIRTH & BLANTON, 1973: 424 (male, female; Brazil). Distr.- Brazil (Pará).avilaensis ORTÍZ & MIRSA, 1951: 593 (female; Venezuela). Distr.- Venezuela.azureus WIRTH & BLANTON, 1959: 377 (male, female; Panama). Distr.- Panama.barthi TAVARES & SOUZA, 1978: 619 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).bricenoi ORTÍZ, 1951a: 445 (male; Venezuela). Distr.- Ecuador, Venezuela, Bolivia, Brazil

(Amazonas, Pará).cuiabai WIRTH, 1982b: 250 (female; Brazil). Distr.- Brazil (Mato Grosso), northeastern

Argentina.dalessandroi WIRTH & BARRETO, 1978: 556 (male, female; Colombia). Distr.- Costa Rica,

Panama, Colombia.daviesi WIRTH & BLANTON, 1968b: 251 (female; Guyana). Distr.- Peru, Guyana.dicrourus WIRTH & BLANTON, 1955b: 123 (male, female; Panama). Distr.- Costa Rica to Ecuador.discrepans ORTÍZ & MIRSA, 1951: 595 (female; Venezuela). Distr.- Venezuela.lenti TAVARES & LUNA DIAS, 1980: 396 (male; Brazil). Distr.- Bolivia, Brazil (Rio de Janeiro).macieli TAVARES & RUIZ, 1980: 29 (male, female; Brazil). Distr.- Brazil (Bahia, Rio de Janeiro).mojingaensis WIRTH & BLANTON, 1953b: 232 (male, female; Panama). Distr.- Panama.volcanensis WIRTH & BLANTON, 1959: 389 (female; Panama). Distr.- Panama, Colombia.wallacei WIRTH & BLANTON, 1973: 449 (male, female; Brazil). Distr.- Brazil (Pará).

Subgenus OECACTA POEY

OECACTA POEY, 1853: 238. Type species: Oecacta furens POEY, by monotypy.

alahialinus BARBOSA, 1952: 11 (1953: 12) (female; Ecuador). Distr.- Costa Rica, Panama,Colombia, Ecuador.

barbosai WIRTH & BLANTON, 1956a: 161 (male, female; Panama). Distr.- USA (Florida) toEcuador.

cancer HOGUE & WIRTH, 1968: 2 (larva, pupa, male, female; Costa Rica). Distr.- Mexico(Sinaloa), El Salvador, Costa Rica.

furens (POEY), 1853: 238 (Oecacta; female; Cuba). Distr.- USA (Massachusetts to Floridaand Texas), Mexico (Campeche, Sinaloa, Yucatán, Veracruz) and Caribbean to Ecuadorand coastal Brazil.

maculithorax (WILLISTON), 1896: 277 (Ceratopogon). St. Vincent.birabeni CAVALIERI, 1966: 59. Venezuela.

gorgasi WIRTH & BLANTON, 1953b: 232 (male, female; Panama). Distr.- Costa Rica toColombia.

Subgenus PSYCHOPHAENA PHILIPPI

PSYCHOPHAENA PHILIPPI, 1865: 628. Type species: Psychophaena pictipennis PHILIPPI (=Culicoides venezuelensis ORTÍZ & MIRSA), by monotypy.


lacustris RONDEROS, 1990a: 116 (male, female; Argentina). Distr.- Southern Argentina.venezuelensis ORTÍZ & MIRSA, 1950: 137 (Venezuela). Distr.- Costa Rica to Chile and

Central Argentina.pictipennis (PHILIPPI), 1865: 628 (Psychophaena) (preoccupied by Culicoidespictipennis (STAEGER), 1839). Chile.ortizi FOX, 1952: 366. Venezuela.

Subgenus unplaced, acotylus Species Group

acotylus LUTZ, 1913: 69 (female; Brazil). Distr.- Mexico (DF), Honduras, Panama, Venezuela,Trinidad, Surinam, Brazil (Mato Grosso, Pará).

panamericanus FOX, 1947: 90. Mexico.atripalpis WIRTH & BLANTON, 1973: 425 (female; Brazil). Distr.- Brazil (Pará).carsiomelas WIRTH & BLANTON, 1955a: 100 (male, female; Panama). Distr.- Panama,

Colombia, Brazil (Pará).teretipalpis WIRTH & BARRETO, 1978: 557 (male, female; Colombia). Distr.- Colombia.

Subgenus unplaced, carpenteri Species Group

belemensis WIRTH & BLANTON, 1973: 427 (male, female; Brazil). Distr.- Colombia, Brazil(Amazonas, Pará).

camposi ORTÍZ & LEÓN, 1955: 580 (female; Ecuador). Distr.- Costa Rica, Panama, Colombia,Ecuador.

fairchildi WIRTH & BLANTON, 1955a: 102. Panama.carpenteri WIRTH & BLANTON, 1953a: 72 (male, female; Panama). Distr.- Costa Rica, Panama,

Ecuador, Bolivia, Brazil (Amazonas).

Subgenus unplaced, daedalus Species Group

antefurcatus WIRTH & BLANTON, 1959: 315 (male, female; Panama). Distr.- Panama.beaveri WIRTH & BARRETO, 1978: 557 (female; Colombia). Distr.- Colombia.commatis WIRTH & BLANTON, 1959: 321 (male, female; Panama). Distr.- Panama.crescentis WIRTH & BLANTON, 1959: 317 (male, female; Panama). Distr.- Mexico (Chiapas)

to Colombia, northeastern Argentina.cummingi SPINELLI & BORKENT, 2004a: 379 (female; Costa Rica). Distr.- Costa Rica.daedaloides WIRTH & BLANTON, 1959: 330 (male, female; Panama). Distr.- Panama,

Colombia.daedalus MACFIE, 1948: 83 (male; Mexico). Distr.- USA (Arizona, New Mexico), Mexico

(Chiapas) to Colombia.dunni WIRTH & BLANTON, 1959: 328 (female; Panama). Distr.- Costa Rica, Panama.pampoikilus MACFIE, 1948: 79 (female; Mexico). Distr.- USA (Arizona, New Mexico),

Mexico (Chiapas, Oaxaca) to Venezuela.dominicii ORTÍZ, 1951b: 7. Venezuela.

phaeonotus WIRTH & BLANTON, 1959: 326 (male, female; Panama). Distr.- Panama.


picadoae SPINELLI & BORKENT, 2004a: 381 (male, female; Costa Rica). Distr.- Costa Rica.pseudocrescentis TAVARES & LUNA DIAS, 1980: 397 (male; Brazil). Distr.- Brazil (Rio de

Janeiro), Paraguay, northeastern Argentina.

Subgenus unplaced, dasyophrus Species Group

dasyophrus MACFIE, 1940d: 27 (male, female; Guyana). Distr.- Colombia, Ecuador,Venezuela, Guyana, Brazil (Amazonas, Mato Grosso, Pará).

estevezae RONDEROS & SPINELLI, 1994: 47 (female; Argentina). Distr.- Northern Argentina.guerrai WIRTH & BLANTON, 1971: 41 (male, female; Trinidad. Distr.- Trinidad, Brazil (Pará, Bahia).rodriguezi ORTÍZ, 1968: 67 (female; Venezuela). Distr.- Panama, Venezuela.

Subgenus unplaced, eublepharus Species Group

archboldi WIRTH & BLANTON, 1970b: 39 (male, female; Dominica). Distr.- Dominica, Trinidad.caldasi BROWNE, 1980: 535 (female; Colombia). Distr.- Colombia.caucaensis WIRTH & LEE, 1967: 19 (female; Colombia). Distr.- Colombia.eublepharus MACFIE, 1948: 86 (female; Guyana). Distr.- Mexico (Chiapas), Costa Rica to

Ecuador, Venezuela, northern Brazil.transferrans ORTÍZ, 1953a: 801. Venezuela.

florenciae MESSERSMITH, 1972: 167 (female; Colombia). Distr.- Colombia.guadeloupensis FLOCH & ABONNENC, 1950b: 2 (female; Guadeloupe). Distr.- Guadeloupe.guarani RONDEROS & SPINELLI, 1994: 48 (female; Argentina). Distr.- Northeastern Argentina.pabloi BROWNE, 1980: 541 (male, female; Colombia). Distr.- Colombia.propriipennis MACFIE, 1948: 84 (female; Mexico). Distr.- Mexico (Chiapas) to Panama,

Ecuador, Venezuela and northern Brazil.rangeli ORTÍZ & MIRSA, 1952b: 126 (female; Venezuela). Distr.- Mexico (Oaxaca) to Ecuador,

Bolivia, Venezuela, Trinidad, Brazil (Amazonas).donajii VARGAS, 1954: 28. Mexico.patulipalpis WIRTH & BLANTON, 1959: 421. Panama.

tamboensis WIRTH & LEE, 1967: 20 (male, female; Colombia). Distr.- Colombia.zumbadoi SPINELLI & BORKENT, 2004a: 383 (male, female; Costa Rica). Distr.- Costa Rica.

Subgenus unplaced, fluvialis Species Group

balsapambensis ORTÍZ & LEÓN, 1955: 569 (as variety of pifanoi; female; Ecuador). Distr.-Costa Rica to Ecuador, Brazil.

castillae FOX, 1946: 251 (female; Honduras). Distr.- Guatemala to Ecuador, Venezuela, Trinidad.gibsoni WIRTH, 1952c: 246. Guatemala.flochabonnenci ORTÍZ & MIRSA, 1952a: 267. Venezuela.

fernandezi ORTÍZ, 1954: 223 (female; Venezuela). Distr.- Venezuela.fluvialis MACFIE, 1940d: 25 (female; Guyana). Distr.- Honduras to Colombia, Venezuela,

Trinidad, Guyana, Brazil (Amazonas, Pará).leopoldoi ORTÍZ, 1951c: 579 (female; Venezuela). Distr.- Guatemala and Belize to Bolivia

and northeastern Argentina, Trinidad.


lichyi FLOCH & ABONNENC, 1949: 1 (1950a: 69) (female; Venezuela). Distr.- Venezuela.pulchripennis MACFIE, 1939b: 200 (female; Brazil). Distr.- Brazil (Santa Catarina).tetrathyris WIRTH & BLANTON, 1959: 409 (male, female; Panama). Distr.- Honduras, Costa

Rica, Panama, Ecuador, Trinidad, Surinam, northern Brazil.williamsi SPINELLI, in SPINELLI et al., 2005b: 147 (male, female; Paraguay). Distr.- Paraguay,

northeastern Argentina.yaracuyensis ORTÍZ, 1959: 364 (as yaracuyanus; female; Venezuela). Distr.- Venezuela.

Subgenus unplaced, leoni Species Group

benarrochi ORTÍZ & MIRSA, 1952b: 126 (as benarrochei; female; Venezuela). Distr.- Brazil(Rio de Janeiro), Venezuela, Trinidad.

fieldi WIRTH & BLANTON, 1956c: 50 (male, female; Honduras). Distr.- Honduras, CostaRica, Panama.

gabaldoni ORTÍZ, 1954: 221 (female; Venezuela). Distr.- Mexico (Tabasco) to Ecuador,Venezuela, Trinidad, Brazil (Bahia), Paraguay, northeastern Argentina.

glabellus WIRTH & BLANTON, 1956c: 47 (male, female; Panama). Distr.- Honduras toEcuador, Trinidad, Brazil (Bahia, Pará).

leoni BARBOSA, 1952: 17 (1953: 19) (female; Ecuador). Distr.- Ecuador.trifidus SPINELLI & BORKENT, 2004a: 385 (male, female; Costa Rica). Distr.- Costa Rica.

Subgenus unplaced, limai Species Group

antunesi FORATTINI, 1954b: 315 (male, female; Brazil). Distr.- Brazil (Goiás, São Paulo).boliviensis SPINELLI & WIRTH, 1984a: 172 (female; Bolivia). Distr.- Bolivia, Brazil (Minas

Gerais), northeastern Argentina.carvalhoi WIRTH & BLANTON, 1973: 429 (male, female; Brazil). Distr.- Brazil (Pará).duartei TAVARES & LUNA DIAS, 1980: 395 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).galindoi WIRTH & BLANTON, 1953a: 73 (male, female; Panama). Distr.- Costa Rica, Panama.limai BARRETTO, 1944: 99 (male; Brazil). Distr.- El Salvador to Ecuador, Brazil (Pará, Mato

Grosso, São Paulo, Rio de Janeiro, Santa Catarina) to northeastern Argentina.lobatoi FELIPPE-BAUER, in FELIPPE-BAUER & QUINTELAS, 1994: 25 (female; Brazil). Distr.-

Brazil (Rio de Janeiro).lopesi BARRETTO, 1944: 102 (male; Brazil). Distr.- Panama, Surinam, Brazil (São Paulo).santanderi BROWNE, 1980: 536 (female; Colombia). Distr.- Colombia.tenuilobus WIRTH & BLANTON, 1959: 354 (male, female; Panama). Distr.- Guatemala to Panama.vernoni WIRTH & BLANTON, 1973: 448 (male, female; Brazil). Distr.- Costa Rica, Colombia,

Bolivia, Brazil (Pará).

Subgenus unplaced, monticola Species Group

andinus WIRTH & LEE, 1967: 17 (female; Colombia). Distr.- Colombia.magnipalpis WIRTH & BLANTON, 1953a: 76 (male, female; Panama). Distr.- Panama.monticola WIRTH & LEE, 1967: 15 (female; Colombia). Distr.- Costa Rica to Ecuador.

pichindensis BROWNE, 1980: 538. Colombia.


Subgenus unplaced, pachymerus Species Group

almirantei WIRTH & BLANTON, 1959: 454 (male, female; Panama). Distr.- Costa Rica, Panama.atelis WIRTH, 1982b: 249 (female; Panama). Distr.- Panama.caprilesi FOX, 1952: 364 (female; Venezuela). Distr.- Panama, Colombia, Venezuela, Brazil

(Pará, Mato Grosso).kintzi WIRTH & BLANTON, 1953a: 72. Panama.

cylindricornis WIRTH & BLANTON, 1973: 430 (female; Brazil). Distr.- Brazil (Pará).obnoxius FOX, 1952: 365 (female; Venezuela). Distr.- Colombia, Venezuela.pachymerus LUTZ, 1914: 83 (female; Brazil). Distr.- Guatemala to Colombia, Brazil (Amazonas).uniradialis WIRTH & BLANTON, 1953a: 70 (male, female; Panama). Distr.- Panama, Colombia.

Subgenus unplaced, reticulatus Species Group

aureus ORTÍZ, 1951c: 585 (male; Venezuela). Distr.- Panama, Bolivia, Venezuela, Paraguay,Brazil (Amazonas), northeastern Argentina.

miyamotoi WIRTH & BLANTON, 1953b: 231. Panama.fittkaui WIRTH & BLANTON, 1973: 432 (male, female; Brazil). Distr.- Brazil (Pará).forattinii ORTÍZ, 1961: 211 (female; Venezuela). Distr.- Venezuela.goeldii WIRTH & BLANTON, 1973: 437 (male, female; Brazil). Distr.- Brazil (Pará).guamai WIRTH & BLANTON, 1973: 438 (male, female; Brazil). Distr.- Brazil (Pará).guyanensis FLOCH & ABONNENC, 1942a: 4 (male, female; French Guiana). Distr.- Panama,

Venezuela to French Guiana, Trinidad and Tobago, Brazil (Pará, Pernambuco, São Paulo).recifensis BARBOSA, 1943: 263. Brazil.stubalensis FOX, 1946: 254. Trinidad and Tobago.

kuscheli WIRTH & BLANTON, 1978: 236 (female; Chile). Distr.- Northern Chile.lanei ORTÍZ, 1950a: 431 (male; Panama). Distr.- Mexico (Veracruz), Honduras, Costa Rica,

Panama, Venezuela, Trinidad, Brazil (Pará).lyrinotatus WIRTH & BLANTON, 1955b: 126 (male, female; Panama). Distr.- Nicaragua,

Panama, Brazil.macrostigma WIRTH & BLANTON, 1953b: 230 (male, female; Panama). Distr.- Costa Rica

to Colombia.martinezi WIRTH & BLANTON, 1970b: 43 (male, female; Trinidad). Distr.- Trinidad.paucienfuscatus BARBOSA, 1947: 23 (male; Brazil). Distr.- Costa Rica to Peru and

Bolivia,Venezuela, Trinidad, Brazil (Amazonas, Pará).pifanoi ORTÍZ, 1951c: 588 (male, female; Venezuela). Distr.- Belize to Colombia, Venezuela,

Trinidad, Brazil (Pará, Bahia), Paraguay.tricoloratus WIRTH & BLANTON, 1953b: 233. Panama.

reticulatus LUTZ, 1913: 49 (male, female; Brazil). Distr.- Honduras to Colombia, Brazil(Pernambuco, Bahia, São Paulo, Rio de Janeiro).

tavaresi FELIPPE-BAUER & WIRTH, 1988: 261 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).


Subgenus unplaced, stigmalis Species Group

REFERENCE: FELIPPE-BAUER & WIRTH, 1987 (key to males and females).

alvarezi ORTÍZ, 1957: 161 (female; Venezuela). Distr.- Ecuador, Venezuela.deanei FELIPPE-BAUER & WIRTH, 1987: 416 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro)fluviatilis (LUTZ), 1914: 82 (Johannseniella; female; Brazil). Distr.- Colombia, Ecuador, Bolivia,

Venezuela, Trinidad, Brazil (Amazonas).scorzai ORTÍZ, 1956: 93. Venezuela.

stigmalis WIRTH, 1952c: 245 (female; Guatemala). Distr.- Mexico (Oaxaca), Guatemala,Costa Rica, Panama.

Subgenus unplaced, stonei Species Group

melleus (COQUILLETT), 1901a: 604 (Ceratopogon; female; Florida). Distr.- Eastern Canada (NewBrunswick) and USA (to Louisiana and Florida), Bahamas, Mexico (Baja California).

Miscellaneous Unplaced Species

arubae FOX & HOFFMAN, 1944: 109 (male, female; Aruba). Distr.- USA (Texas), Mexico(Yucatán), Aruba and Grenada, to Colombia and Venezuela.

floridensis BECK, 1951: 135 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas, Bermuda.malariologiensis PERRUOLO, 1990: 28 (female; Venezuela). Distr.- Venezuela.pancensis BROWNE, 1980: 537 (male, female; Colombia). Distr.- Colombia.polystictus KIEFFER, 1921b: 181 (female; Paraguay). Distr.- Paraguay.propinquus MACFIE, 1948: 81 (male; Mexico). Distr.- Mexico (Chiapas).trilineatus FOX, 1946: 250 (female; Virgin Islands). Distr.- Guatemala to Panama, Puerto

Rico, Virgin Islands, Dominica, Barbados, Paraguay.unetensis PERRUOLO, 2001: 35 (female; Venezuela). Distr.- Venezuela.wokei FOX, 1947: 90 (female; Panama). Distr.- Costa Rica, Panama.

Genus PARADASYHELEA MACFIE

PARADASYHELEA MACFIE, 1940f: 17. Type species: Dasyhelea brevipalpis INGRAM &MACFIE, by original designation.

REFERENCES: SPINELLI, 1987 (description of female); SPINELLI & GROGAN, 2003 (key tomales and females).

brevipalpis (INGRAM & MACFIE), 1931: 178 (Dasyhelea; male; Argentina). Distr.- SouthwesternArgentina south to Tierra del Fuego.

ingrami SPINELLI & GROGAN, 2003: 572 (male, female; Argentina). Distr.- Southtern Argentina.macfiei SPINELLI & GROGAN, 2003: 574 (male, female; Argentina). Distr.- Southwestern

Argentina, southern Chile.


TRIBE CERATOPOGONINI

Genus ALLOHELEA KIEFFER

ALLOHELEA KIEFFER, 1917b: 364. Type species: Sphaeromias pulchripennis KIEFFER, by originaldesignation.

neotropica WIRTH, 1991a: 503 (male, female; Jamaica). Distr.- Belize, Panama, Colombia, Jamaica.

Genus ALLUAUDOMYIA KIEFFER

ALLUAUDOMYIA KIEFFER, 1913: 12. Type species: Alluaudomyia imparunguis KIEFFER, bymonotypy.

NEOCERATOPOGON MALLOCH, 1915b: 310. Type species: Ceratopogon bellus COQUILLETT,by original designation.

ISOECACTA GARRETT, 1925: 9. Type species: Isoecacta poeyi GARRETT (= Ceratopogon bellusCOQUILLETT), by original designation.

REFERENCES: HUERTA, 1996 (key to males and females from Mexico); SPINELLI & WIRTH,1984c (key to males and females).

amazonica SPINELLI & WIRTH, 1984c: 676 (female, male; Brazil). Distr.- Brazil (Amazonas),northeastern Argentina.

bella (COQUILLETT), 1902: 87 (Ceratopogon; USA, District of Columbia). Distr.- North America(Alaska to California and Nova Scotia and Florida), Mexico (Veracruz), Bahamas,Cayman Islands.

biestroi SPINELLI, 1988: 130 (pupa, male; Argentina). Distr.- Northeastern Argentina,Uruguay.

caribbeana SPINELLI & WIRTH, 1984c: 678 (pupa, male, female; Belize). Distr.- USA(California to Texas, Florida), Central America, circum-Caribbean from Mexico(Yucatán, Morelos, Sinaloa) to Colombia and Venezuela.

catarinensis SPINELLI & WIRTH, 1984c: 681 (female; Brazil). Distr.- Brazil (Santa Catarina).distispinulosa SPINELLI & WIRTH, 1984c: 682 (male, female; Brazil). Distr.- Colombia,

Bolivia, Venezuela, Brazil (Amazonas).estevezae SPINELLI & WIRTH, 1984c: 684 (male, female; Mexico). Distr.- Mexico (Morelos),

Honduras, El Salvador.fittkaui SPINELLI & WIRTH, 1984c: 686 (male, female; Brazil). Distr.- Ecuador, Brazil

(Amazonas, Pará).guarani SPINELLI, 1988: 131 (male, female; Uruguay). Distr.- Paraguay, northeastern

Argentina, Uruguay.leei SPINELLI & WIRTH, 1984c: 687 (male, female; Colombia). Distr.- Colombia, Bolivia.nubeculosa SPINELLI & WIRTH, 1984c: 689 (male, female; Brazil). Distr.- Colombia, Brazil

(Amazonas, Pará).plaumanni SPINELLI & WIRTH, 1984c: 691 (female; Brazil). Distr.- Brazil (Santa Catarina).prima CLASTRIER, 1976: 205 (female; French Guiana). Distr.- Panama, French Guiana.


punctivenosa WIRTH & GROGAN, 1988: 20. New name for punctiradialis. Distr.- Brazil (Pará),Paraguay, northeastern Argentina.

punctiradialis SPINELLI & WIRTH, 1984c: 692 (preoccupied by Alluaudomyia punctiradialisCHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Brazil.

schnacki SPINELLI, 1983b: 403 (larva, pupa, male, female; Argentina). Distr.- EasternArgentina, Uruguay.

sexpunctata SPINELLI & WIRTH, 1984c: 695 (female; Colombia). Distr.- Colombia.spinellii WIRTH & GROGAN, 1988: 21. New name for tripunctata. Distr.- Colombia.

tripunctata SPINELLI & WIRTH, 1984c: 697 (preoccupied by Alluaudomyia tripunctataCHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Colombia.

tenuiannulata SPINELLI & WIRTH, 1984c: 696 (male, female; Guatemala). Distr.- Guatemala,Costa Rica.

youngi SPINELLI & WIRTH, 1984c: 699 (male, female; Colombia). Distr.- Panama, Colombia.

Genus AUSTROHELEA WIRTH & GROGAN

AUSTROHELEA WIRTH & GROGAN, 1988: 22. Type species: Monohelea shannoni WIRTH &BLANTON, by original designation.

REFERENCE: WIRTH & GROGAN, 1988 (description of male).

shannoni (WIRTH & BLANTON), 1972b: 175 (Monohelea; female; Argentina). Distr.- SouthernChile, southwestern Argentina south to Tierra del Fuego.

Genus BAEODASYMYIA CLASTRIER & RACCURT

BAEODASYMYIA CLASTRIER & RACCURT, 1979b: 100. Type species: Baeodasymyia modestaCLASTRIER & RACCURT, by monotypy.

REFERENCE: BORKENT & CRAIG, 1999 (keys to pupae, males, females).

christopheri BORKENT, in BORKENT & CRAIG, 1999: 15 (larva, pupa, male, female; CostaRica). Distr.- Costa Rica.

gustavoi BORKENT, in BORKENT & CRAIG, 1999: 20 (female; Paraguay). Distr.- Paraguay,Argentina (Misiones).

lydiae BORKENT, in BORKENT & CRAIG, 1999: 19 (male, female; Costa Rica). Distr.- Costa Rica.michaeli BORKENT, in BORKENT & CRAIG, 1999: 13 (larva, pupa, male, female; Costa Rica).

Distr.- Costa Rica, Panama.modesta CLASTRIER & RACCURT, 1979b: 100 (male, female; Haiti). Distr.- Haiti.

Genus BAEOHELEA WIRTH & BLANTON

BAEOHELEA WIRTH & BLANTON, 1970c: 95. Type species: Baeohelea nana WIRTH &BLANTON, by original designation.


nana WIRTH & BLANTON, 1970c: 95 (male, female; Dominica). Distr.- Costa Rica to Ecuador,St. Kitts, Dominica.

Genus BAHIAHELEA WIRTH

BAHIAHELEA WIRTH, 1992: 276. Type species: Bahiahelea brasiliensis WIRTH, by originaldesignation.

brasiliensis WIRTH, 1992: 280 (male, female; Brazil). Distr.- Brazil (Bahia).

Genus BORKENTHELEA SPINELLI & GROGAN

BORKENTHELEA SPINELLI & GROGAN, 1993: 321. Type species: Borkenthelea nothofagusSPINELLI & GROGAN, by original designation.

REFERENCE: SPINELLI & GROGAN, 2001 (key to males and females species).

harii SPINELLI & GROGAN, 2001: 148 (male, female; Argentina). Distr. - SouthwesternArgentina, southern Chile.

nerudai SPINELLI & GROGAN, 2001: 151 (male, female; Chile). Distr. - Southern Chile.nothofagus SPINELLI & GROGAN, 1993: 323 (male, female; Argentina). Distr.- Southwestern

Argentina, southern Chile.quatei SPINELLI & GROGAN, 2001: 153 (male, female; Argentina). Distr.- Argentina (Chubut).

Genus BRACHYPOGON KIEFFER

BRACHYPOGON KIEFFER, 1899: 69. Type species: Ceratopogon vitiosus WINNERTZ, byoriginal designation.

REFERENCE: SPINELLI & WIRTH, 1993 (key to subgenera as part of larger key to genera).

Subgenus BRACHYPOGON KIEFFER

REFERENCE: SPINELLI & GROGAN, 1998 (key to males and females).

apunctipennis SPINELLI & GROGAN, 1998: 64 (male; Brazil). Distr.- Brazil (Rondônia).bifidus SPINELLI & GROGAN, 1998: 66 (male, female; Dominica). Distr.- El Salvador, Dominica.bimaculatus SPINELLI & GROGAN, 1998: 67 (male, female; Colombia). Distr.- Mexico

(Yucatán), Colombia, Brazil (Mato Grosso).bonaerensis SPINELLI, 1990: 744 (female; Argentina). Distr.- Argentina (Buenos Aires).calchaqui SPINELLI, 1990: 746 (male, female; Argentina). Distr.- Western Argentina.ecuadorensis SPINELLI & GROGAN, 1998: 68 (male, female; Ecuador). Distr.- Belize to

Ecuador, Jamaica.ethelae SPINELLI & GROGAN, 1998: 62 (male, female; Panama). Distr.- Panama.


fuscivenosus (LUTZ), 1914: 94 (Palpomyia; male, female; Brazil). Distr.- USA (Florida), Mexico(Yucatán, Quintana Roo), Panama, Jamaica, Virgin Islands, Guyana, Brazil (Rio de Janeiro).

impar (JOHANNSEN), 1938: 223 (Ceratopogon; female; Puerto Rico). Distr.- Colombia,Venezuela, Puerto Rico, Brazil (Santa Catarina).

insularis SPINELLI & GROGAN, 1998: 70 (male; Cuba). Distr.- Cuba.monicae SPINELLI & GROGAN, 1998: 70 (male, female; Colombia). Distr.- Colombia.paraensis WIRTH & BLANTON, 1970c: 99 (as parasensis; male, female; Brazil). Distr.- Brazil (Pará).pseudoparaensis SPINELLI & GROGAN, 1998: 64 (male; Brazil). Distr.- Brazil (Pará).ringueleti SPINELLI, 1990: 748 (male, female; Argentina). Distr.- Southern Argentina.schmitzi SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondônia).spatuliformis SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondônia).telesfordi SPINELLI & GROGAN, 1998: 71 (male, female; St. Vincent). Distr.- Costa

Rica, St. Vincent, Grenada.woodruffi SPINELLI & GROGAN, 1998: 72 (male, female; Dominica Republic). Distr.- Mexico

(Yucatán), Dominica Republic.

Subgenus ISOHELEA KIEFFER

ISOHELEA KIEFFER, 1917b: 295. Type species: Ceratopogon lacteipennis ZETTERSTEDT

(misidentified, = Psilohelea sociabilis GOETGHEBUER), by original designation.


borkenti SPINELLI & CAZORLA, 2004: 3 (male, female; Argentina). Distr.- Western, southernArgentina.

cuacuahuitlus HUERTA & BORKENT, 2005: 115 (male; Mexico). Distr.- Mexico (Jalisco).hugoi SPINELLI & GROGAN, 1994: 2 (male, female; Colombia). Distr.- Colombia.mapuche SPINELLI, 1990: 750 (male, female; Argentina). Distr.- Southwestern Argentina.misionensis SPINELLI, 1990: 752 (male; Argentina). Distr.- Northeastern Argentina,

southeastern Brazil.pallidipennis SPINELLI & GROGAN, 1994: 6 (male, female; Honduras). Distr.- Honduras.wirthi SPINELLI, 1990: 753 (female; Argentina). Distr.- Northwestern Argentina.

Genus CACAOHELEA WIRTH & GROGAN

CACAOHELEA WIRTH & GROGAN, 1988: 33. Type species: Cacaohelea youngi WIRTH &GROGAN, by original designation.

youngi WIRTH & GROGAN, 1988: 34 (female; Costa Rica). Distr.- Costa Rica.

Genus CERATOCULICOIDES WIRTH & RATANAWORABHAN

CERATOCULICOIDES WIRTH & RATANAWORABHAN, 1971a: 170. Type species: Helealongipennis WIRTH, by original designation.


aliciae HUERTA & BORKENT, 2005: 112 (male, female; Mexico). Distr.- Mexico (Jalisco).

Genus DIAPHANOBEZZIA INGRAM & MACFIE

DIAPHANOBEZZIA INGRAM & MACFIE, 1931: 223. Type species: Diaphanobezzia pellucidaINGRAM & MACFIE, by original designation.

araucaria SPINELLI, 1996: 77 (female; Argentina). Distr.- Southwestern Argentina.patagonica SPINELLI & GROGAN, 1990: 127 (female; Argentina). Distr.- Southwestern

Argentina.pellucida INGRAM & MACFIE, 1931: 223 (male; Argentina). Distr.- Southwestern Argentina.spinellii WIRTH & GROGAN, 1988: 48 (male, female; Argentina). Distr.- Southwestern

Argentina.

Genus DOWNESHELEA WIRTH & GROGAN

DOWNESHELEA WIRTH & GROGAN, 1988: 50. Type species: Monohelea stonei WIRTH, byoriginal designation.

REFERENCE: LANE & WIRTH, 1964 (key to males and females).

balboa (LANE & WIRTH), 1964: 225 (Monohelea; male, female; Panama). Distr.- Panama.bicornis FELIPPE-BAUER & QUINTELAS, 1993b: 185 (male; Brazil). Distr.- Brazil (Rio de Janeiro).blantoni (LANE & WIRTH), 1964: 217 (Monohelea; male, female; Panama). Distr.- Panama.carioca (TAVARES & SILVA PEREIRA), 1978: 157 (Monohelea; male, female; Brazil). Distr.- Brazil

(Rio de Janeiro).castroi (TAVARES & SILVA PEREIRA), 1978: 159 (Monohelea; male, female; Brazil). Distr.- Brazil

(Rio de Janeiro).cebacoi (LANE & WIRTH), 1964: 218 (Monohelea; male, female; Panama). Distr.- Panama,

Brazil (Pará).charrua FELIPPE-BAUER & SPINELLI, 1994: 161 (male, female; Uruguay). Distr.- Uruguay.chiapasi (LANE & WIRTH), 1964: 219 (Monohelea; male, female; Nicaragua). Distr.- Nicaragua

to Colombia.chirusi (LANE & WIRTH), 1964: 218 (Monohelea; male, female; Panama). Distr.- Nicaragua, Panama.colombiae (LANE & WIRTH), 1964: 220 (Monohelea; male; Colombia). Distr.- Colombia.deanei FELIPPE-BAUER & QUINTELAS, in FELIPPE-BAUER et al., 1995: 395 (male; Trinidad).

Distr.- Trinidad.fluminensis FELIPPE-BAUER & QUINTELAS, 1993a: 33 (male; Brazil). Distr.- Brazil (Rio de Janeiro).fuscipennis (LANE & WIRTH), 1964: 221 (Monohelea). Colombia.guianae (WIRTH), 1953b: 150 (Monohelea; male; Guyana). Distr.- Brazil (Pará), Guyana.multilineata (LUTZ), 1914: 93 (Palpomyia; male, female; Brazil). Distr.- USA (Florida, Puerto

Rico), Mexico (Yucatán), Panama, Guyana, Brazil (Mato Grosso, Rio de Janeiro).panamensis (LANE & WIRTH), 1964: 221 (Monohelea; male, female; Panama). Distr.- Mexico

(Guerrero), Panama, Virgin Islands.


quasidentica FELIPPE-BAUER & QUINTELAS, 1993a: 37 (male; Brazil). Distr.- Brazil (Rio de Janeiro).stonei (WIRTH), 1953b: 148 (Monohelea; male, female; USA, Louisiana). Distr.- Eastern North

America (Ontario, Massachusetts to Florida), Caribbean to Colombia and Trinidad.

Genus ECHINOHELEA MACFIE

ECHINOHELEA MACFIE, 1940c: 187. Type species: Echinohelea ornatipennis MACFIE, byoriginal designation.

REFERENCE: WIRTH, 1994b (key to males and females).

Subgenus ECHINOHELEA MACFIE

blantoni WIRTH, 1994b: 232 (male, female; Panama). Distr.- Mexico (Yucatán, Oaxaca),Panama to Ecuador.

jamaicensis WIRTH, 1994b: 234 (male, female; Jamaica). Distr.- Jamaica.lanei WIRTH, 1951b: 319 (male, female; USA, Virgina). Distr.- USA (Virginia, Michigan and

Massachetts to Florida), Panama, Colombia, Trinidad, Brazil (Pará, Rondônia).leei WIRTH, 1994b: 237 (male; Colombia). Distr.- Colombia.macfiei LANE, 1948: 228 (female; Brazil). Distr.- Brazil (São Paulo).neotropica WIRTH, 1994b: 237 (male, female; Colombia). Distr.- Panama to Ecuador, Brazil

(Amazonas, Pará, Rio de Janeiro).ornatipennis MACFIE, 1940c: 188 (male, female; Guyana). Distr.- Panama, Trinidad, Guyana,

Brazil (Pará).panamensis WIRTH, 1994b: 239 (male, female; Panama). Distr.- Panama to Ecuador.richardsi MACFIE, 1940c: 189 (female; Guyana). Distr.- Guyana, Brazil (Pará, Rondônia,

São Paulo).smarti MACFIE, 1940c: 190 (female; Guyana). Distr.- Colombia, Guyana.

Subgenus ECHINOIDESHELEA WIRTH

ECHINOIDESHELEA WIRTH, 1994b: 231 (as subgenus of Echinohelea). Type species:Echinohelea aitkeni WIRTH, by original designation.

aitkeni WIRTH, 1994b: 231 (male, female; Brazil). Distr.- Brazil (Pará).

Genus FITTKAUHELEA WIRTH & BLANTON

FITTKAUHELEA WIRTH & BLANTON, 1970a: 7. Type species: Fittkauhelea amazonica WIRTH

& BLANTON, by original designation.

amazonica WIRTH & BLANTON, 1970a: 9 (male, female; Brazil). Distr.- Brazil (Amazonas).


Genus ISTHMOHELEA INGRAM & MACFIE

ISTHMOHELEA INGRAM & MACFIE, 1931: 208. Type species: Isthmohelea disjuncta INGRAM

& MACFIE, by original designation.

disjuncta INGRAM & MACFIE, 1931: 209 (male, female; Chile). Distr.- Southern Chile.

Genus LEPTOHELEA WIRTH & BLANTON

LEPTOHELEA WIRTH & BLANTON, 1970a: 12. Type species: Leptohelea micronyx WIRTH &BLANTON, by original designation.

micronyx WIRTH & BLANTON, 1970a: 12 (female; Colombia). Distr.- Costa Rica, Colombia.

Genus MACRUROHELEA INGRAM & MACFIE

MACRUROHELEA INGRAM & MACFIE, 1931: 203. Type species: Macrurohelea caudataINGRAM & MACFIE, by original designation.


caudata INGRAM & MACFIE, 1931: 205 (male, female; Chile). Distr.- Southern Chile.fuscipennis SPINELLI & GROGAN, 1990: 128 (female; Argentina). Distr.- Southwestern Argentina.gentilii SPINELLI & GROGAN, 1984: 963 (male, female; Argentina). Distr.- Southwestern Argentina.irwini GROGAN & WIRTH, 1980b: 140 (male, female; Chile). Distr.- Central and southern Chile.kuscheli WIRTH, 1965c: 49 (female; Chile). Distr.- Central Chile.monotheca SPINELLI & GROGAN, 1984: 965 (female; Argentina). Distr.- Southwestern Argentina.paracaudata GROGAN & WIRTH, 1980b: 141 (male; Chile). Distr.- Central and southern

Chile, southwestern Argentina.setosa WIRTH, 1965c: 49 (male, female; Chile). Distr.- Northern Chile.similis SPINELLI & GROGAN, 1990: 131 (male; Argentina). Distr.- Southwestern Argentina.thoracica INGRAM & MACFIE, 1931: 206 (male, female; Argentina). Distr.- Southwestern

Argentina, southern Chile.wirthi SPINELLI & GROGAN, 1984: 965 (female; Argentina). Distr.- Southwestern Argentina.yamana SPINELLI & GROGAN, 1999: 710 (male, female; Argentina). Distr.- Argentina (Tierra

del Fuego).

Genus MONOHELEA KIEFFER

MONOHELEA KIEFFER, 1917b: 294. Type species: Monohelea hieroglyphica KIEFFER, byoriginal designation.

REFERENCE: LANE & WIRTH, 1964 (key to males and females).


affinis FELIPPE-BAUER & SPINELLI, 1991: 201 (male, female; Brazil). Distr.- Brazil (Amazonas).aguirrei TAVARES & SOUZA, 1980: 97 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro,

Santa Catarina), Argentina (Corrientes, Buenos Aires province).archibaldoi TAVARES & SOUZA, 1980: 98 (male, female; Brazil). Distr.- Brazil (Rio de Janeiro).bidentata FELIPPE-BAUER & SPINELLI, 1994: 163 (male; Argentina). Distr.- Argentina (Buenos Aires).brasiliensis LANE, 1948: 226 (female; Brazil). Distr.- Brazil (Rio de Janeiro).cunasi LANE & WIRTH, 1964: 232 (male, female; Panama). Distr.- Panama.fairchildi LANE & WIRTH, 1964: 214 (male, female; Panama). Distr.- Panama.guaimiesi LANE & WIRTH, 1964: 227 (male, female; Panama). Distr.- Panama.hieroglyphica KIEFFER, 1917b: 312 (male, female; Paraguay). Distr.- Dominica, Brazil (Goiás,

Rio de Janeiro), Paraguay.lanei WIRTH, 1953b: 142 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas,

Costa Rica.maculipennis (COQUILLETT), 1905: 64 (Ceratopogon; female; USA, Florida). Distr.- USA

(Florida, Louisiana), Bahamas, Mexico (Yucatán, Tamaulipas), Guatemala, Panama.maya FELIPPE-BAUER, HUERTA & IBAÑEZ-BERNAL, 2000a: 815 (male, female; Mexico). Distr.-

Mexico (Yucatán).mayeri ORTÍZ, 1950d: 202 (female; Venezuela). Distr.- Costa Rica to Colombia, Venezuela, Trinidad.ornata WIRTH, 1953b: 144 (female; USA, Florida). Distr.- USA (Florida), Panama.poncai LANE & WIRTH, 1964: 228 (male, female; Panama). Distr.- Costa Rica, Panama,

Brazil (Rio de Janeiro).roraimensis FELIPPE-BAUER & SPINELLI, 1991: 202 (male; Brazil). Distr.- Brazil (Roraima).urracaisi LANE & WIRTH, 1964: 231 (male, female; Panama). Distr.- Panama, Guyana.uruguayensis FELIPPE-BAUER & SPINELLI, 1998: 63 (male; Uruguay). Distr.- Uruguay.

Genus NANNOHELEA GROGAN & WIRTH

NANNOHELEA GROGAN & WIRTH, 1980a: 374. Type species: Ceratopogon bourioniCLASTRIER, by original designation.

clastrieri GROGAN & WIRTH, 1980a: 381 (male; Colombia). Distr.- Colombia.

Genus NOTIOHELEA GROGAN & WIRTH

NOTIOHELEA GROGAN & WIRTH, 1979b: 283. Type species: Notiohelea chilensis GROGAN

& WIRTH, by original designation.

chilensis GROGAN & WIRTH, 1979b: 284 (female; Chile). Distr.- Southern Chile.pilosa SPINELLI & GROGAN, 1990: 133 (female; Argentina). Distr.- Southwestern Argentina.

Genus PARABEZZIA MALLOCH

PARABEZZIA MALLOCH, 1915a: 358. Type species: Parabezzia petiolata MALLOCH, by originaldesignation.


REFERENCE: SPINELLI & GROGAN, 1987 (key to females and some males).

alexanderi WIRTH, 1965b: 219 (female; USA, Massachesetts). Distr.- Eastern North America(Ontario and Massachesetts to Florida), Mexico (Veracruz), Belize, El Salvador,Argentina (Misiones).

arenosa CLASTRIER & RACCURT, 1979a: 172 (female; Haiti). Distr.- Haiti.balseiroi SPINELLI & GROGAN, 1987: 13 (pupa, male, female; Argentina). Distr.- Argentina

(Mendoza, Corrientes, Entre Ríos).blantoni WIRTH, 1965b: 220 (female; Panama). Distr.- Panama.brasiliensis SPINELLI & GROGAN, 1987: 34 (female; Brazil). Distr.- Brazil (Mato Grosso).brunnea WIRTH, 1965b: 220 (female; Panama). Distr.- Belize, Panama.caribbeana CLASTRIER & RACCURT, 1979a: 173 (female; Haiti). Distr.- Haiti.cayoensis SPINELLI & GROGAN, 1987: 36 (female; Belize). Distr.- Belize.clastrieri SPINELLI & GROGAN, 1987: 23 (female; El Salvador). Distr.- Belize, El Salvador.costalis WIRTH, 1965b: 221 (male, female; Panama). Distr.- Panama.fuscipennis WIRTH, 1965b: 223 (female; Panama). Distr.- Panama.haitiensis CLASTRIER & RACCURT, 1979a: 170 (female; Haiti). Distr.- Haiti.hondurensis SPINELLI & GROGAN, 1987: 25 (male, female; Honduras). Distr.- Honduras.inaequalis SPINELLI & GROGAN, 1987: 38 (male, female; Brazil). Distr.- Brazil (Amazonas).jamaicensis WIRTH, 1965b: 224 (female; Jamaica). Distr.- Jamaica.neunguis GROGAN & WIRTH, 1977: 65 (female; USA, Virgina). Distr.- USA (Virginia,Maryland), El Salvador.pallida SPINELLI & GROGAN, 1987: 40 (female; Mexico). Distr.- Mexico (Oaxaca).panamensis WIRTH, 1965b: 225 (male, female; Panama). Distr.- Panama.pseudunguis SPINELLI & GROGAN, 1987: 30 (male, female; Panama). Distr.- Panama.raccurti SPINELLI & GROGAN, 1987: 42 (male, female; El Salvador). Distr.- El Salvador,

northern Argentina.spangleri WIRTH, 1965b: 227 (female; Puerto Rico). Distr.- Puerto Rico.uncinata (JOHANNSEN), 1943: 761 (Stilobezzia; female; USA, Alabama). Distr.- USA (Alabama,

Maryland, Virginia, Arkansas), Jamaica.unguis WIRTH, 1965b: 228 (male, female; USA, Arizona). Distr.- USA (Arizona, Texas) to Colombia.wirthi CLASTRIER & RACCURT, 1979a: 169 (female; Haiti). Distr.- Haiti.

Genus PARASTILOBEZZIA WIRTH & BLANTON

PARASTILOBEZZIA WIRTH & BLANTON, 1970a: 10. Type species: Parastilobezzia leei WIRTH

& BLANTON, by original designation.

leei WIRTH & BLANTON, 1970a: 10 (male, female; Colombia). Distr.- Costa Rica, Colombia.

Genus RHYNCHOHELEA WIRTH & BLANTON

RHYNCHOHELEA WIRTH & BLANTON, 1970c: 96. Type species: Rhynchohelea monilicornisWIRTH & BLANTON, by original designation.


monilicornis WIRTH & BLANTON, 1970c: 98 (female; USA, Florida). Distr.- USA (Florida),Costa Rica.

Genus SCHIZONYXHELEA CLASTRIER

SCHIZONYXHELEA CLASTRIER, 1984: 1. Type species: Schizonyxhelea guyana CLASTRIER,by original designation.

forattinii WIRTH & GROGAN, 1988: 81 (male, female; Brazil). Distr.- Costa Rica, Brazil(Santa Catarina).

guyana CLASTRIER, 1984: 2 (female; French Guiana). Distr.- Mexico (Yucatán), FrenchGuiana, Brazil (Mato Grosso).

Genus STILOBEZZIA KIEFFER

STILOBEZZIA KIEFFER, 1911a: 118. Type species: Stilobezzia festiva KIEFFER, by originaldesignation. HARTOMYIA MALLOCH, 1915a: 339. Type species: Ceratopogon pictusCOQUILLETT (= Stilobezzia coquilletti KIEFFER), by original designation.

REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM &MACFIE,1931 (key males and females from Patagonia and southern Chile); LANE & FORATTINI,1961 (key to males and females from Neotropical Region north of Patagonia andsouthern Chile); WIRTH & GROGAN, 1988 (key to subgenera, as part of generic key).

Subgenus ACANTHOHELEA KIEFFER

ACANTHOHELEA KIEFFER, 1917a: 198. Type species: Acanthohelea pruinosa KIEFFER, bymonotypy.

amazonica CLASTRIER, 1991: 306 (male, female; French Guiana). Distr.- French Guiana, Peru.atrichopogon LANE & FORATTINI, 1956: 208 (male, female; Panama). Distr.- Panama.bicinctipes INGRAM & MACFIE, 1931: 195 (female; Chile). Distr.- Southern Chile.edwardsi INGRAM & MACFIE, 1931: 198 (male, female; Argentina). Distr.- Southwestern Argentina.furva INGRAM & MACFIE, 1931: 200 (female; Argentina). Distr.- Southern Chile, southwestern

Argentina.guianae (MACFIE), 1940d: 28 (Acanthohelea; male, female; Guyana). Distr.- Guyana.hirsuta INGRAM & MACFIE, 1931: 201 (male, female; Argentina). Distr.- Southwestern Argentina.maia LANE & FORATTINI, 1958: 204 (male, female; Panama). Distr.- Panama, Colombia.manaosensis LANE & FORATTINI, 1958: 205 (male, female; Brazil). Distr.- Brazil (Amazonas).nigerrima INGRAM & MACFIE, 1931: 196 (female; Argentina). Distr.- Southwestern Argentina.ornata LANE & FORATTINI, 1958: 206 (female; Panama). Distr.- Panama.ornaticrus INGRAM & MACFIE, 1931: 194 (female; Chile). Distr.- Southern Chile.patagonica INGRAM & MACFIE, 1931: 196 (male, female; Argentina). Distr.- Southwestern

Argentina.


rava INGRAM & MACFIE, 1931: 203 (male, female; Chile). Distr.- Southern Chile.sanctibernardini KIEFFER, 1917b: 308 (female; Paraguay). Distr.- Paraguay.succinea INGRAM & MACFIE, 1931: 200 (male, female; Argentina). Distr.- Chile, southwestern

Argentina south to Tierra del Fuego.tibialis LANE & FORATTINI, 1956: 209 (male, female; Panama). Distr.- Panama.varia INGRAM & MACFIE, 1931: 191 (male, female; Chile). Distr.- Southern Chile,

southwestern Argentina south to Tierra del Fuego.

Subgenus EUKRAIOHELEA INGRAM & MACFIE

EUKRAIOHELEA INGRAM & MACFIE, 1921: 347. Type species: Eukraiohelea africana INGRAM

& MACFIE, designation by MACFIE, 1940f: 22.

REFERENCE: WIRTH & SPINELLI, 1992b (key males and females).

amnigena (MACFIE), 1935a: 56 (Eukraiohelea; female; Brazil). Distr.- USA (Florida, SouthCarolina), (?) Jamaica, (?) Puerto Rico, Brazil (Maranhão), northeastern Argentina.

dorsofasciata (LUTZ), 1914: 96 (Palpomyia; female; Brazil). Distr.- Brazil (Rio de Janeiro).elegantula (JOHANNSEN), 1907: 109 (Bezzia; female; USA, Kansas). Distr.- USA (Kansas to

Maryland to Louisiana and Florida), Mexico (Yucatán), Panama, (?) Puerto Rico,Paraguay, Argentina.

maculitibia LANE & FORATTINI, 1956: 207. Panama.subsessilis KIEFFER, 1917b: 311. Paraguay.

Subgenus STILOBEZZIA KIEFFER

albicoxa LANE & FORATTINI, 1956: 210 (male, female; Panama). Distr.- Panama.albocincta, KIEFFER 1917b: 309 (female; Paraguay). Distr.- Paraguay.americana KIEFFER, 1917b: 310 (female; Paraguay). Distr.- Paraguay.antennalis (COQUILLETT), 1901a: 606 (Ceratopogon; female; USA, District of Columbia).

Distr.- North America (British Columbia to Ontario to Texas and Florida), Panama.beckae WIRTH, 1953a: 69 (male, female; USA, Florida). Distr.- USA (Maryland to Mississippi,

Florida), Mexico (Yucatán), Panama, Peru.bicolor LANE, 1947a: 208 (male; Brazil). Distr.- Panama, Brazil (São Paulo), Argentina

(Buenos Aires).bimaculata LANE & FORATTINI, 1956: 211 (male, female; Panama). Distr.-Panama, Trinidad,

Brazil (Rio de Janeiro).bispinosa KIEFFER, 1917b: 310 (female; Paraguay). Distr.- Brazil (São Paulo), Paraguay.blantoni LANE & FORATTINI, 1956: 211 (male, female; Panama). Distr.- Panama.caribe LANE & FORATTINI, 1958: 208 (male; Panama). Distr.- Panama.chaconi MACFIE, 1938: 166 (male; Trinidad). Distr.- Costa Rica, Trinidad, Brazil (Goiás, São Paulo).coquilletti KIEFFER, 1917b: 308. (New name for pictus COQUILLETT, female; USA, Virginia).

Distr.- USA (Virgina, Illinois, Maryland), Mexico (Yucatán, Chiapas) to Panama,Trinidad, Brazil (São Paulo).


picta (COQUILLETT), 1905: 60 (Ceratopogon) (preoccupied by Sphaeromias pictus (MEIGEN),1818). USA (Virginia).

coracina KIEFFER, 1917b: 311 (male; Paraguay). Distr.- Brazil (São Paulo), Paraguay,Argentina (Buenos Aires).

diminuta LANE & FORATTINI, 1958: 209 (male, female; Panama). Distr.- Panama, Dominica.dryadum MACFIE, 1940b: 186 (female; Guyana). Distr.- Panama, Guyana.dubitans LANE, FORATTINI & RABELLO, 1955: 85 (pupa, male; Brazil). Distr.- Brazil (São Paulo).dureti LANE & FORATTINI, 1958: 210 (male; Brazil). Distr.- Brazil (São Paulo).esmeralda LANE & FORATTINI, 1958: 211 (male, female; Panama). Distr.- Panama.femoralis LANE & FORATTINI, 1956: 212 (male; Panama). Distr.- Mexico (Yucatán), Panama.fiebrigi KIEFFER, 1917b: 309 (female; Paraguay). Distr.- Brazil (São Paulo, Rio de Janeiro),

Paraguay, Argentina (Buenos Aires province).glauca MACFIE, 1939b: 204 (male; Brazil). Distr.- USA (Maryland to Mississippi and Florida),

Mexico (Chiapas) to southeastern Brazil.fluminensis LANE, 1947a: 210. Brazil.

grandis LANE & FORATTINI, 1958: 213 (female; Panama). Distr.- Panama.kiefferi LANE, 1947a: 205 (female; Brazil). Distr.- USA (Florida), Mexico (Yucatán,

Tamaulipas), Panama, Brazil (Minas Gerais).punctipes WIRTH, 1953a: 79. USA (Florida).

macfiei LANE, 1947a: 213 (male, female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro).maculata LANE, 1947a: 207 (male; Brazil). Distr.- Mexico (Yucatán), Panama, Venezuela,

Brazil (Rio de Janeiro).merceri CAZORLA & SPINELLI, in CAZORLA et al., 2005: 290 (male, female, pupa; Peru). Distr.- Peru.modesta LANE, 1947a: 206 (female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro).nigroflava LANE & FORATTINI, 1958: 215 (male, female; Costa Rica). Distr.- Costa Rica,

Panama, Puerto Rico.obscura LANE & FORATTINI, 1958: 216 (male, female; Panama). Distr.- Panama.pallescens LANE & FORATTINI, 1958: 218 (male, female; Panama). Distr.- Panama, Brazil

(São Paulo).panamensis LANE & FORATTINI, 1958: 218 (male, female; Panama). Distr.- Panama, Puerto

Rico, Brazil (São Paulo, Rio de Janeiro).paulistensis LANE, 1947a: 200 (male, female; Brazil). Distr.- Panama, USA (Puerto Rico),

Brazil (São Paulo, Rio de Janeiro).punctulata LANE, 1947a: 204 (female; Brazil). Distr.- Mexico (Tabasco), Panama, Perú,

Brazil (Mato Grosso, Rio de Janeiro).rabelloi LANE, 1947a: 203 (male, female; Brazil). Distr.- USA (Maryland to Louisiana and

Florida) to Argentina.silvicola MACFIE, 1940b: 185 (male; Guyana). Distr.- Guyana.similans LANE & FORATTINI, 1956: 214 (male, female; Panama). Distr.- Panama.simplex LANE & FORATTINI, 1958: 222 (male, female; Panama). Distr.- Panama.thomsenae WIRTH, 1953a: 83 (male, female; USA, Florida). Distr.- USA (Florida), Mexico

(Yucatán), Panama.scutata LANE & FORATTINI, 1961: 92. Panama.

transversa LANE & FORATTINI, 1958: 222 (male, female; Panama). Distr.- Panama.


travassosi LANE, 1947a: 210 (female; Brazil). Distr.- Brazil (Espírito Santo).venezuelensis ORTÍZ, 1950d: 199 (as variety of glauca; female; Venezuela). Distr.- Venezuela.williamsi CAZORLA & SPINELLI, in CAZORLA et al., 2005: 292 (male, female; Peru). Distr.- Peru.wirthi LANE & FORATTINI, 1956: 214 (male, female; Panama). Distr.- Panama.wygodzinskyi LANE, 1947a: 212 (female; Brazil). Distr.- Panama, Brazil (São Paulo).

TRIBE HETEROMYIINI WIRTH, 1962: 275

Genus CLINOHELEA KIEFFER

CLINOHELEA KIEFFER, 1917b: 295. Type species: Ceratopogon variegatus WINNERTZ

(= Ceratopogon unimaculata MACQUART), by original designation.

Subgenus CERATOBEZZIA KIEFFER

CERATOBEZZIA KIEFFER, 1917b: 326. Type species: Ceratobezzia fallax KIEFFER, byoriginal designation.

fallax KIEFFER, 1917b: 326 (female; Paraguay). Distr.- El Salvador to Colombia, Venezuela,Brazil (Amazonas, Pará, Mato Grosso), Paraguay, northeastern Argentina.

barrettoi LANE & DURET, 1954: 249. Brazil.

Subgenus CLINOHELEA KIEFFER

REFERENCE: SPINELLI & DURET, 1993 (key to males and females).

albopennis LANE, 1944: 259 (female; Brazil). Distr.- Brazil (Pará, São Paulo), Paraguay,northeastern Argentina.

argentina LANE & DURET, 1954: 248 (female; Argentina). Distr.- Northeastern Argentina.damascenoi LANE & DURET, 1954: 250 (female; Brazil). Distr.- Peru, Bolivia, French Guiana,

Brazil (Pará), Paraguay.horacioi LANE, 1944: 257 (male, female; Brazil). Distr.- Mexico (Yucatán, Tabasco, Morelos)

to northeastern Argentina.muzoni SPINELLI & DURET, 1993: 46 (male, female; Colombia). Distr.- Colombia.neivai LANE, 1944: 252 (male, female; Brazil). Distr.- Panama, Brazil (Goiás, Minas Gerais,

São Paulo, Rio de Janeiro), Paraguay.nigripes MACFIE, 1939b: 205 (female; Brazil). Distr.- Costa Rica to Colombia, Brazil (São

Paulo, Santa Catarina), northeastern Argentina.rubriceps KIEFFER, 1917b: 318 (female; Paraguay). Distr.- Paraguay.saltanensis LANE & DURET, 1954: 252 (female; Argentina). Distr.- Brazil (Rondônia, Santa

Catarina), Paraguay, northern Argentina.townsendi LANE, 1944: 256 (female; Brazil). Distr.- Panama to Argentina (Corrientes, Buenos

Aires province).townesi LANE, 1944: 254. Brazil.


wygodzinskyi LANE, 1948: 231 (female; Brazil). Distr.- Brazil (Rio de Janeiro).

Genus HETEROMYIA SAY

HETEROMYIA SAY, 1825: plate 35. Type species: Heteromyia fasciata SAY, by monotypy.PACHYLEPTUS WALKER, 1856: 426. Type species: Pachyleptus fasciatus WALKER (= Heteromyia

nigra KIEFFER), by monotypy.

REFERENCE: DURET & LANE, 1955 (key to males and females).

antequerae (LYNCH ARRIBÁLZAGA), 1893: 227 (Pachyleptus; female; Argentina). Distr.-Northeastern Argentina.

bejaranoi DURET & LANE, 1955: 36 (male, female; Argentina). Distr.- NortheasternArgentina.

castanea LANE, 1946a: 214 (female; Brazil). Distr.- Brazil (São Paulo).chaquensis DURET & LANE, 1955: 37 (female; Argentina). Distr.- Northeastern Argentina.clavata WILLISTON, 1900: 225 (female; Mexico). Distr.- Mexico (Veracruz) to Colombia,

Brazil (Amazonas), northeastern Argentina.rufa KIEFFER, 1917b: 325. Colombia.caloptera KIEFFER, 1919b: 192. Unnecessary new name for H. rufa KIEFFER, not LOEW, 1861.

correntina DURET & LANE, 1955: 39 (male, female; Argentina). Distr.- Panama, Brazil(Rondônia), northeastern Argentina.

kiefferi LANE, 1946a: 213 (female; Brazil). Distr.- Brazil (Rio de Janeiro).lamprogaster EDWARDS, 1933: 87 (female; Argentina). Distr.- Northeastern Argentina.nigra KIEFFER, 1917b: 326 (female; Paraguay). Distr.- Brazil (São Paulo, Rio de Janeiro,

Paraná), Paraguay.fasciatus (WALKER), 1856: 426 (Pachyleptus) (preoccupied by Heteromyia fasciata SAY, 1825).

South America.orellana (ROBACK), 1957: 1 (Palpomyia; female; Peru). Distr.- Peru.wokei WIRTH & GROGAN, 1977: 181 (pupa, male, female; Nicaragua). Distr.- Nicaragua.

Genus PELLUCIDOMYIA MACFIE

PELLUCIDOMYIA MACFIE, 1939a: 99. Type species: Pellucidomyia ugandae MACFIE, byoriginal designation.

MACFIEHELEA LANE, 1946a: 208. Type species: Macfiehelea oliveirai LANE, by originaldesignation.

REFERENCE: WIRTH, 1960 (World key to males and females).

blantoni (LANE), 1956b: 435 (Macfiehelea; female; Panama). Distr.- Panama.lanei WIRTH & RATANAWORABHAN, 1971b: 446 (female; Colombia). Distr.- Colombia, Jamaica.oliveirai (LANE), 1946a: 209 (Macfiehelea; female; Brazil). Distr.- Brazil (Mato Grosso, Minas Gerais).wirthi (LANE), 1956b: 437 (Macfiehelea; female; Panama). Distr.- USA (Texas) to Colombia.


Genus PHYSOHELEA GROGAN & WIRTH

REFERENCE: SPINELLI, 1994 (description of male P. turgidipes).

PHYSOHELEA GROGAN & WIRTH, 1979a: 53. Type species: Neurohelea oedidactyla INGRAM

& MACFIE, by original designation.

oedidactyla (INGRAM & MACFIE), 1931: 212 (Neurohelea; female; Argentina). Distr.-Southwestern Argentina.

turgidipes (INGRAM & MACFIE), 1931: 214 (Neurohelea; female; Chile). Distr.- Southern Chile.

TRIBE SPHAEROMIINI NEWMAN, 1834: 388

Genus AUSTROSPHAEROMIAS SPINELLI

AUSTROSPHAEROMIAS SPINELLI, 1997a: 224. Type species: Austrosphaeromias apricans(INGRAM & MACFIE), by original designation.

apricans (INGRAM & MACFIE), 1931: 219 (Palpomyia; male, female; Argentina). Distr.-Southwestern Argentina, southern Chile.

sentior (INGRAM & MACFIE), 1931: 222 (Palpomyia, as variety of apricans). Chile.wirthi SPINELLI, 1997a: 228 (female; Argentina). Distr.- Southwestern Argentina.

Genus GROGANHELEA SPINELLI & DIPPOLITO

GROGANHELEA SPINELLI & DIPPOLITO, in SPINELLI et al., 1995: 166. Type species:Groganhelea rondoniensis SPINELLI & DIPPOLITO, by original designation.

rondoniensis SPINELLI & DIPPOLITO, in SPINELLI et al., 1995: 167 (female; Brazil). Distr.-Brazil (Rondônia), Uruguay.

Genus JOHANNSENOMYIA MALLOCH

JOHANNSENOMYIA MALLOCH, 1915a: 332. Type species: Johannsenomyia halteralisMALLOCH (= Ceratopogon argentata LOEW), designation by WIRTH, 1952a: 211.

DICROHELEA KIEFFER, 1917b: 363. Type species: Palpomyia filicornis KIEFFER, designationby MACFIE, 1940f: 26.

blantoni (LANE & WIRTH ), 1961: 81 (Dicrohelea; female; Panama). Distr.- Panama.lanei WIRTH, 1965a: 4 (female; Brazil). Distr.- Brazil (Santa Catarina).

Genus LANEHELEA WIRTH & BLANTON

LANEHELEA WIRTH & BLANTON, 1972a: 433. Type species: Lanehelea leei WIRTH andBLANTON, by original designation.


leei WIRTH & BLANTON, 1972a: 434 (female; Colombia). Distr.- Colombia.spinifemur WIRTH & BLANTON, 1972a: 436 (female; Colombia). Distr.- Colombia.

Genus MALLOCHOHELEA WIRTH

MALLOCHOHELEA WIRTH, 1962: 278. Type species: Johannsenomyia albibasis MALLOCH,by original designation.

aenipes (MACFIE), 1940c: 193 (Bezzia; female; Guyana). Distr.- Guyanalimitrofe SPINELLI & FELIPPE-BAUER, 1990a: 87 (female; Argentina). Distr.- Northeastern

Argentina, Uruguay.nemoralis (MACFIE), 1940c: 190 (Johannsenomyia; male; Guyana). Distr.- Guyana.nigripes (MACFIE), 1939b: 216 (Johannsenomyia; female; Brazil). Distr.- Brazil (Santa Catarina).pullata (WIRTH), 1952a: 213 (Johannsenomyia; female; USA, California). Distr.- USA (California,

Arizona, Texas) to Brazil (Amapá).bicellii (LANE), 1961b: 449 (Nilobezzia). Brazil.

termophila (SPINELLI), 1984: 197 (Neobezzia; male, female; Argentina). Distr.- Brazil(Rondônia), northern Argentina.

Genus NEOBEZZIA WIRTH & RATANAWORABHAN

NEOBEZZIA WIRTH & RATANAWORABHAN, 1972a: 477. Type species: Neobezzia clavipesWIRTH & RATANAWORABHAN, by original designation.

REFERENCE: WIRTH & RATANAWORABHAN, 1972a (key to females).

albitarsis WIRTH & RATANAWORABHAN, 1972a: 478 (female; Brazil). Distr.- Brazil (Pará).amnicola (MACFIE), 1940d: 30 (Bezzia; female; Guyana). Distr.- Belize to Brazil (Amazonas,

Rondônia, Mato Grosso, São Paulo).blantoni WIRTH & RATANAWORABHAN, 1972a: 482 (female; Panama). Distr.- Nicaragua to

Colombia, Brazil (Amazonas, Pará, Rondônia), northeastern Argentina.brasiliae (LANE), 1961a: 37 (Macropeza; female; Brazil). Distr.- Brazil (Brasilia, D.F.).clavipes WIRTH & RATANAWORABHAN, 1972a: 485 (female; Brazil). Distr.- Panama, Brazil (Amazonas).costaricae WIRTH & RATANAWORABHAN, 1972a: 486 (female; Costa Rica). Distr.- Costa Rica,

Panama.fittkaui WIRTH & RATANAWORABHAN, 1972a: 489 (female; Brazil). Distr.- Brazil (Amazonas).wirthi SPINELLI & FELIPPE-BAUER, 1990a: 88 (male, female; Argentina). Distr.- Northeastern

Argentina.

Genus NILOBEZZIA KIEFFER

NILOBEZZIA KIEFFER, 1921a: 24. Type species: Nilobezzia armata KIEFFER, by monotypy.

REFERENCE: LANE, 1961b (key to males and females).


neotropica (MACFIE), 1940a: 78 (Bezzia; female; Brazil). Distr.- Brazil (Rondônia, Bahia).paraensis (LANE), 1958: 33 (Bezzia; female; Brazil). Distr.- Brazil (Pará).schwarzii (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Texas). Distr.- USA (South

Carolina to Texas and Florida), Mexico (Yucatán), Bahamas, Cuba, Panama, Brazil(Rio de Janeiro), northeastern Argentina.

banksi (GERRY), 1933: 94 (Bezzia). Cuba.brasiliensis (LANE), 1958: 28 (Bezzia). Brazil.

Genus SPHAEROHELEA SPINELLI & FELIPPE-BAUER

SPHAEROHELEA SPINELLI & FELIPPE-BAUER, 1990b: 195. Type species: Sphaerohelea biestroiSPINELLI & FELIPPE-BAUER, by original designation.

biestroi SPINELLI & FELIPPE-BAUER, 1990b: 197 (female; Argentina). Distr.- NortheasternArgentina.

TRIBE PALPOMYIINI ENDERLEIN, 1936: 49

Genus AMEROHELEA GROGAN & WIRTH

AMEROHELEA GROGAN & WIRTH, 1981: 1280. Type species: Amerohelea galindoi GROGAN

& WIRTH, by original designation.

REFERENCES: GROGAN & WIRTH, 1981 (key to females); HUERTA, 1996 (key to femalesfrom Mexico).

dalcyi GROGAN & WIRTH, 1981: 1289 (female; Brazil). Distr.- Brazil (Amazonas).fasciata GROGAN & WIRTH, 1981: 1283 (male, female; Belize). Distr.- Mexico (Sonora,

Morelos, Tabasco, Chiapas) to Colombia.frontispina (DOW & TURNER), 1976: 138 (Bezzia; female; USA, Texas). Distr.- USA

(California to Texas) south to Colombia and Venezuela.galindoi GROGAN & WIRTH, 1981: 1294 (male, female; Colombia). Distr.- Panama,

Colombia,Venezuela, northern Argentina.nelsoni GROGAN & WIRTH, 1981: 1290 (female; Brazil). Distr.- Brazil (Amazonas).pseudofasciata GROGAN & WIRTH, 1981: 1286 (female; Brazil). Distr.- Southeastern Brazil,

Argentina (Misiones, Río Negro).ronderosi GROGAN & WIRTH, 1981: 1298 (male, female; Colombia). Distr.- Colombia.similis SPINELLI, 1989: 27 (female; Uruguay). Distr.- Northern Argentina, Uruguay.sordidipes (MACFIE), 1939b: 209 (Palpomyia; female; Brazil). Distr.- Bolivia, Brazil (Pará,

Minas Gerais, São Paulo, Santa Catarina).spinellii GROGAN & WIRTH, 1981: 1292 (female; Colombia). Distr.- Colombia.vargasi GROGAN & WIRTH, 1981: 1296 (male, female; Belize) Distr.- Mexico (Veracruz,

Oaxaca), Belize to Costa Rica.


Genus BEZZIA KIEFFER

BEZZIA KIEFFER, 1899: 69. Type species: Ceratopogon ornatus MEIGEN, by original designation.

REFERENCES: HUERTA, 1996 (key to males and females from Mexico); SPINELLI & WIRTH,1989a (key to subgenera and species groups for males and females, key to males andfemales of nobilis and punctipennis groups); SPINELLI & WIRTH, 1989b (key to femalesand some males of glabra and brevicornis groups); SPINELLI & WIRTH, 1990 (key to malesand females of gibbera group); SPINELLI & WIRTH, 1991 (key to males and females ofdentifemur and venustula groups).

aitkeni SPINELLI & WIRTH, 1991: 5 (female; Brazil). Distr.- Brazil (Pará).albuquerquei LANE, 1961a: 43 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).araucana SPINELLI & WIRTH, 1990: 14 (male, female; Argentina). Distr.- Southwestern

Argentina.bivittata (COQUILLETT), 1905: 60 (Ceratopogon; female; USA, California). Distr.- North America

(Alaska to Nova Scotia to California and Florida), Panama.blantoni SPINELLI & WIRTH, 1989b: 771 (pupa, male, female; Honduras). Distr.- Belize to

Costa Rica, Puerto Rico, Argentina (Corrientes, Buenos Aires).brevicornis (KIEFFER), 1917b: 328 (Allobezzia; male, female; Paraguay). Distr.- Paraguay,

Argentina (Corrientes, Entre Ríos, Buenos Aires, Río Negro), Uruguay.bromeliae SPINELLI & WIRTH, 1991: 9 (pupa, male, female; Panama). Distr.- Panama.capitata WIRTH & GROGAN, 1983: 503 (male; Honduras). Distr.- USA (Arizona) to Costa Rica.carioca LANE, 1958: 30 (female; Brazil). Distr.- Brazil (Rio de Janeiro).catarinensis SPINELLI & WIRTH, 1990: 17 (male, female; Brazil). Distr.- Brazil (Santa Catarina)cayoensis SPINELLI & WIRTH, 1991: 6 (male, female; Belize). Distr.- Belize.chilensis SPINELLI & RONDEROS, 2001: 752 (Male, female; Chile). Distri.- Chile, Argentina

(Salta, Rio Negro).clavipennis SPINELLI & WIRTH, 1989b: 775 (female; Brazil). Distr.- Colombia, Ecuador,

Venezuela, Brazil (Amazonas, Rondônia).dentifemur SPINELLI & WIRTH, 1991: 3 (male, female; Colombia). Distr.- Belize, Colombia,

Trinidad, Brazil (Pará).filiducta SPINELLI & WIRTH, 1991: 3 (as filiductus; female; Colombia). Distr.- Honduras,

Colombia.flinti SPINELLI & WIRTH, 1989a: 113 (male, female; Dominica). Distr.- Dominica.fluminensis LANE, 1948: 236 (female; Brazil). Distr.- Brazil (Brasilia, D.F., Rio de Janeiro),

Argentina (Corrientes).fusca SPINELLI & WIRTH, 1991: 9 (female; Colombia). Distr.- Colombia.gibbera (COQUILLETT), 1905: 60 (Ceratopogon; female; Cuba). Distr.- USA (Arizona to Florida)

to Panama, Cuba, Jamaica, Tobago.glabra (COQUILLETT), 1902: 85 (Ceratopogon; female; USA, Florida). Distr.- North America

(Alaska to New Brunswick to California and Florida), Belize, El Salvador.globulosa SPINELLI & WIRTH, 1990: 18 (male, female; Puerto Rico). Distr.- Puerto Rico.goianensis LANE, 1961a: 44 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).


grogani SPINELLI & WIRTH, 1990: 19 (male, female; Panama). Distr.- Mexico (Morelos),Panama, Colombia.

hondurensis SPINELLI & WIRTH, 1990: 20 (male, female; Honduras). Distr.- Mexico(Veracruz, Chiapas), Honduras, El Salvador, Panama.

jamaicensis SPINELLI & WIRTH, 1989a: 115 (male, female; Jamaica). Distr.- Jamaica, Haiti.jubata SPINELLI & WIRTH, 1990: 20 (male, female; Colombia). Distr.- Colombia.leei SPINELLI & WIRTH, 1990: 21 (male, female; Colombia). Distr.- Colombia.lenkoi LANE, 1958: 31 (male, female; Brazil). Distr.- Panama, Brazil (Amazonas, Pará, Brasilia,

D.F., São Paulo, Rio de Janeiro).mathisi SPINELLI & WIRTH, 1989a: 117 (female; Ecuador). Distr.- Ecuador.mazaruni MACFIE, 1940c: 193 (female; Guyana). Distr.- USA (California, Florida), El

Salvador, Venezuela, Guyana, Caribbean to Brazil (Amazonas, Rondônia).megatheca SPINELLI & WIRTH, 1990: 22 (female; Colombia). Distr.- Colombia, Brazil

(Rondônia).mesotibialis SPINELLI & WIRTH, 1990: 23 (female; Belize). Distr.- Belize, Trinidad.nigritibialis SPINELLI & WIRTH, 1991: 5 (female; Belize). Distr.- Belize, Costa Rica.nobilis (WINNERTZ), 1852: 79 (Ceratopogon; Germany). Distr.- Palaearctic, North America

(Alaska to Nova Scotia) to southern Argentina.acanthodes MACFIE, 1940c: 192. Guyana.atlantica WIRTH & WILLIAMS, 1957: 13. Bermuda.

pseudogibbera SPINELLI & WIRTH, 1990: 23 (male, female; Panama). Distr.- Belize,Honduras, Panama.

pseudovenustula SPINELLI & WIRTH, 1991: 8 (female; Brazil). Distr.- Brazil (Pará).pulchripes KIEFFER, 1917b: 330 (female; Paraguay). Distr.- Widespread, from Mexico

(Tabasco) to northeastern Argentina.punctipennis (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- USA

(California, Nevada, Arizona, Texas, Florida), Mexico (Yucatán, Tabasco, QuintanaRoo), El Salvador, Honduras, Jamaica, Dominica, Brazil (Amazonas), Argentina(Buenos Aires province).

raposoensis SPINELLI & WIRTH, 1991: 4 (male, female; Colombia). Distr.- Mexico (Yucatán),Colombia.

roldani SPINELLI & WIRTH, 1981: 187 (larva, pupa, male, female; Argentina). Distr.- EasternArgentina (Entre Rios, Buenos Aires).

schmitzorum DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 54 (female; Brazil). Distr.-Brazil (Rondônia).

setigera SPINELLI & WIRTH, 1990: 24 (male, female; Colombia). Distr.- El Salvador, Colombia.snowi LANE, 1958: 34 (male, female; Guatemala). Distr.- Guatemala, Panama, Colombia,

Trinidad, Brazil (Pará).subfusca MACFIE, 1939b: 218 (female; Brazil). Distr.- Brazil (Santa Catarina).venustula (WILLISTON), 1896: 278 (Ceratopogon; female; St. Vincent). Distr.- Widespread,

from Mexico (Veracruz, Morelos, Chiapas) to Argentina.concoloripes MACFIE, 1940d: 31. Guyana.

woodruffi SPINELLI & WIRTH, 1989a: 120 (female; Jamaica). Distr.- Jamaica.


Genus CLASTRIEROMYIA SPINELLI & GROGAN

CLASTRIEROMYIA SPINELLI & GROGAN, 1985: 330. Type species: Clastrieromyia schnackiSPINELLI & GROGAN, by original designation.

REFERENCE: SPINELLI & GROGAN, 1986 (key to females).

dycei SPINELLI & GROGAN, 1986: 456 (pupa, male, female; Uruguay). Distr.- Uruguay, easternand central Argentina.

kremeri SPINELLI & GROGAN, 1985: 332 (female; Brazil). Distr.- Brazil (Amazonas).schnacki SPINELLI & GROGAN, 1985: 331 (female; Ecuador). Distr.- Ecuador, northern

Argentina.uruguayensis SPINELLI & GROGAN, 1986: 458 (female; Uruguay). Distr.- Northeastern and

central Argentina, Uruguay.

Genus PACHYHELEA WIRTH

PACHYHELEA WIRTH, 1959a: 50. Type species: Ceratopogon magnus COQUILLETT

(= Ceratopogon pachymerus WILLISTON), by original designation.

albidiventris (KIEFFER), 1917b: 316 (Sphaeromias; female; Colombia). Distr.- Colombia.pachymera (WILLISTON), 1900: 224 (Ceratopogon; female; Mexico). Distr.- USA (Texas) to

central Argentina.latifemoris (INGRAM & MACFIE), 1931: 231 (Johannsenomyia). Argentina.

Genus PALPOMYIA MEIGEN

PALPOMYIA MEIGEN, 1818: 82. Type species: Ceratopogon flavipes MEIGEN, by monotypy.Generic name first published in synonymy with Ceratopogon but available under ICZNCode Article 11(e).

REFERENCES: INGRAM & MACFIE, 1931 (key males and females from Patagonia andsouthern Chile); LANE, 1960 (key to females).

aculeata INGRAM & MACFIE, 1931: 215 (female; Chile). Distr.- Southern Chile.almeidai (LANE), 1946b: 219 (Sphaeromias; female; Brazil). Distr.- Brazil (São Paulo).barrettoi LANE, 1947b: 442 (female; Brazil). Distr.- Brazil (Rio de Janeiro).boliviensis KIEFFER, 1917b: 324 (female; Bolivia). Distr.- Bolivia.brasiliensis MACFIE, 1939b: 213 (female; Brazil). Distr.- Brazil (Minas Gerais, Santa Catarina).callangana KIEFFER, 1917b: 323 (female; Peru). Distr.- Peru.carioca LANE, 1960: 384 (Brazil; female). Distr.- Brazil (Rio de Janeiro).carrerai (LANE), 1948: 235 (Dicrobezzia, as carreirai; female; Brazil). Distr.- Brazil (São Paulo).castanea MACFIE, 1939b: 211 (female; Brazil). Distr.- Brazil (São Paulo, Santa Catarina).catarinensis LANE, 1960: 385 (female; Brazil). Distr.- Brazil (Santa Catarina).


chilensis INGRAM & MACFIE, 1931: 218 (female; Chile). Distr.- Southern Chile, ? Brazil (SãoPaulo).

columbiana KIEFFER, 1917b: 323 (female; Colombia). Distr.- Colombia.conifera MACFIE, 1939b: 214 (male; Brazil). Distr.- Brazil (Brasilia, D.F., Goiás, Santa Catarina).coroicoensis WIRTH, 1974: 54. New name for boliviensis. Distr.- Bolivia.

boliviensis (KIEFFER), 1917b: 331 (Jenkinsia; female) (preoccupied by Palpomyia boliviensisKIEFFER, 1917b). Bolivia.

crassicrus KIEFFER, 1917b: 321 (female; Paraguay). Distr.- Brazil (São Paulo), Paraguay.guarani LANE, 1946b: 221 (pupa, male, female; Brazil). Distr.- Brazil (São Paulo, Espírito

Santo, Rio de Janeiro), Paraguay.guyana CLASTRIER, 1992: 117 (male, female; French Guiana). Distr.- French Guiana.hispida BORKENT, in BORKENT & WIRTH, 1997: 132. New name for multispinosa. Distr.- El

Salvador to Colombia.multispinosa SPINELLI & GROGAN, 1989: 6 (male, female) (preoccupied by Palpomyia

multispinosa (PIERCE), 1966). El Salvador.iberaensis SPINELLI & CAZORLA, 2006: 1068 (female, male genitalia; northeastern Argentina).

Distr.- Argentina.inermicollis KIEFFER, 1917b: 322 (female; Paraguay). Distr.- Paraguay.insularis SPINELLI & GROGAN, 1989: 3 (male, female; Puerto Rico). Distr.- Jamaica, Dominica,

Puerto Rico.johannseni (LANE), 1948: 233 (Dicrobezzia; female; Brazil). Distr.- Brazil (São Paulo).lacustris LANE, FORATTINI & RABELLO, 1955: 83 (pupa, male; Brazil). Distr.- Brazil (São Paulo).limnochares (MACFIE), 1940a: 76 (Sphaeromias; male; Brazil). Distr.- Northeastern Brazil.lutzi LANE, 1947b: 440 (female; Brazil). Distr.- Brazil (Rio de Janeiro).mellichroa MACFIE, 1939b: 212 (female; Brazil). Distr.- Brazil (Santa Catarina).microchela KIEFFER, 1917b: 321 (female; Colombia). Distr.- Colombia.nigroflava LANE, 1947b: 446 (female; Brazil). Distr.- Brazil (São Paulo).nigroscutellata LANE, 1947b: 444 (female; Brazil). Distr.- Brazil (São Paulo, Rio de Janeiro).oliveirai LANE, 1947b: 443 (female; Brazil). Distr.- Brazil (Rondônia, Minas Gerais).pampana LANE, 1960: 386 (female; Brazil). Distr.- Brazil (Rio Grande do Sul).paraensis LANE, 1960: 387 (female; Brazil). Distr.- Brazil (Pará).patagonica INGRAM & MACFIE, 1931: 222 (female; Argentina). Distr.- Southwestern

Argentina.paulistensis LANE, 1947b: 445 (female; Brazil). Distr.- Panama, Brazil (São Paulo).pseudolacustris DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 55 (male, female; Brazil).

Distr.- Brazil (Rondônia).scabra (COQUILLETT), 1905: 62 (Ceratopogon; female; Mexico). Distr.- Mexico (Tabasco),

Argentina (Buenos Aires province).spinifemur (LANE), 1948: 234 (Dicrobezzia; male, female genitalia; Brazil). Distr.- Brazil

(São Paulo, Rio de Janeiro).spinosa LUTZ, 1914: 93 (female; Brazil). Distr.- Brazil (Rio de Janeiro).subaspera (COQUILLETT), 1901a: 606 (Ceratopogon; female; USA, New Mexico). Distr.- North

America (Alberta to Ontario to California, Florida), Mexico (Chihuahua), Argentina.maculicrus INGRAM & MACFIE, 1931: 230. Argentina.


subfuscula INGRAM & MACFIE, 1931: 216 (male, female; Chile). Distr.- Southern Chile,southwestern Argentina.

tamioi LANE, 1960: 388 (female; Brazil). Distr.- Brazil (Rio de Janeiro).tanycornis BORKENT, in BORKENT & WIRTH, 1997: 134. New name for longicornis. Distr.- St.

Vincent; Brazil (Brasilia, D.F.).longicornis (WILLISTON), 1896: 280 (Ceratopogon; female) (preoccupied by Ceratopogon longicornis

WALTL, 1837). St. Vincent.tenuicrus KIEFFER, 1917b: 320 (female; Paraguay). Distr.- Paraguay.umbella MACFIE, 1939b: 208 (female; Brazil). Distr.- Brazil (Mato Grosso, Rio de Janeiro,

Santa Catarina, Rio Grande do Sul).versicolor MACFIE, 1939b: 215 (female; Brazil). Distr.- Brazil (Rondônia, Santa Catarina).wirthi LANE, FORATTINI & RABELLO, 1955: 82 (pupa, female; Brazil). Distr.- Brazil (São Paulo).

Genus PHAENOBEZZIA HAESELBARTH

PHAENOBEZZIA HAESELBARTH, 1965: 297. Type species: Probezzia pistiae INGRAM &MACFIE, by original designation.

REFERENCE: SPINELLI & WIRTH, 1986a (key to males and females).

ateles (MACFIE), 1940a: 78 (Bezzia; male; Brazil). Distr.- Colombia, Ecuador, Brazil (Bahia).astyla SPINELLI & WIRTH, 1986a: 232. Colombia.

maya SPINELLI & WIRTH, 1986a: 234 (male, female; Belize). Distr.- USA (Texas) to Brazil(Rondônia).

TRIBE STENOXENINI COQUILLETT, 1899: 61

Genus PARYPHOCONUS ENDERLEIN

PARYPHOCONUS ENDERLEIN, 1912: 57. Type species: Paryphoconus angustipennisENDERLEIN, by original designation.

REFERENCES: HUERTA, 1996 (key to females from Mexico); SPINELLI & WIRTH, 1984b(key to females).

aemulus MACFIE, 1940b: 180 (female; Guyana). Distr.- Peru, Guyana, Brazil (Amazonas,Mato Grosso).

amapaensis LANE, 1961b: 450 (female; Brazil). Distr.- Brazil (Amapá).angustipennis ENDERLEIN, 1912: 57 (female; Brazil). Distr.- Colombia, Peru, Venezuela,

Brazil (Goiás, São Paulo, Santa Catarina), northeastern Argentina.anomalicornis KIEFFER, 1917b: 333 (female; Colombia and Paraguay). Distr.- Mexico San Luis

Potosi) to Colombia, Venezuela, Brazil (Amazonas), Paraguay, northeastern Argentina.apicalis SPINELLI & WIRTH, 1984b: 889 (female; Ecuador). Distr.- Panama, Ecuador.barrettoi LANE, 1946a: 203 (female; Brazil). Distr.- Brazil (Goiás, São Paulo).


batesi LANE, 1961c: 455 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).brunneipennis SPINELLI & WIRTH, 1984b: 890 (male, female; Brazil). Distr.- Colombia,

Brazil (Amazonas).ecuadorensis SPINELLI & WIRTH, 1984b: 892 (female; Ecuador). Distr.- Ecuador.enderleini LANE, 1956a: 301 (female; Brazil). Distr.- Brazil (Mato Grosso).fittkaui SPINELLI & WIRTH, 1984b: 892 (female; Brazil). Distr.- Brazil (Amazonas).flavidus (JOHANNSEN), 1943: 761 (Ceratobezzia; female; Guyana). Distr.- Guyana, Brazil

(Amazonas, Mato Grosso).lanei WIRTH, 1959b: 236. Brazil.

flinti SPINELLI & WIRTH, 1984b: 894 (female; Mexico). Distr.- Mexico (Tabasco, Chiapas), Belize.fusciradialis SPINELLI & WIRTH, 1984b: 895 (female; Brazil). Distr.- Brazil (Pará).fuscus LANE, 1946a: 206 (female; Brazil). Distr.- Brazil (Goiás).goianensis LANE, 1961c: 456 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).grandis MACFIE, 1939c: 6 (female; Brazil). Distr.- Mexico (on train quarantined in Arizona),

Belize, Venezuela, Brazil (Pará, Santa Catarina).guianae MACFIE, 1940b: 180 (female; Guyana). Distr.- Ecuador, Guyana, Brazil (Pará, Mato

Grosso, São Paulo).kiefferi LANE, 1956a: 302 (female; Brazil). Distr.- Brazil (Goiás).latipennis SPINELLI & WIRTH, 1984b: 898 (male, female; Colombia). Distr.- Colombia, Venezuela.leei SPINELLI & WIRTH, 1984b: 898 (female; Colombia.). Distr.- Colombia.macfiei LANE, 1946a: 203 (female; Brazil). Distr.- Ecuador, Venezuela, Brazil (Pará, Mato

Grosso, São Paulo).maya SPINELLI & WIRTH, 1984b: 899 (female; Belize). Distr.- Mexico (San Luis Potosi), Belize.mayeri WIRTH, 1959b: 236 (female; Brazil). Distr.- Brazil (Amazonas, Rondônia).misionensis SPINELLI, 1998: 52 (female; Argentina). Distr.- Northeastern Argentina.neotropicalis (LANE), 1948: 229 (Macropeza; female; Brazil). Distr.- Brazil (São Paulo).nigripes MACFIE, 1939c: 8 (female; Argentina). Distr.- Guyana, Brazil (Pará, Rondônia),

northeastern Argentina.nubifer MACFIE, 1939c: 5 (female; Brazil). Distr.- Bolivia, Brazil (Amazonas, Rio de Janeiro,

Santo Catarina), Paraguay, northeastern Argentina, Uruguay.oliveirai LANE, 1956a: 303 (female; Brazil). Distr.- Colombia, Brazil (Amazonas, Pará).paranaensis SPINELLI & WIRTH, 1984b: 902 (female; Argentina). Distr.- Northeastern Argentina.paulistensis LANE, 1961c: 457 (female; Brazil). Distr.- Brazil (São Paulo).sonorensis WIRTH & RATANAWORABHAN, 1972b: 1374 (female; Mexico). Distr.- USA

(Oklahoma), Mexico (Sonora), Belize, Panama.steineri SPINELLI & WIRTH, 1984b: 905 (female; Peru). Distr.- Panama, Peru.subflavus MACFIE, 1940d: 23 (female; Guyana). Distr.- Colombia, Guyana, Brazil (Amazonas,

Pará, Rondônia).travassosi LANE, 1956a: 304. Brazil.

taragui SPINELLI, 1998: 53 (female; Argentina). Distr.- Northeastern Argentina.telmatophilus (MACFIE), 1940a: 77 (Bezzia; male; Brazil). Distr.- Northeastern Brazil.terminalis (COQUILLETT), 1904a: 90 (Ceratopogon; female; Nicaragua). Distr.- Belize to Peru

and Ecuador, Brazil (Amazonas, Santa Catarina).unimaculatus MACFIE, 1940b: 179 (female; Guyana). Distr.- Guyana, Brazil (Pará).


wirthi LANE, 1961c: 458 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).wygodzinskyi LANE, 1946a: 208 (female; Brazil). Distr.- Brazil (Rio de Janeiro).

Genus STENOXENUS COQUILLETT

STENOXENUS COQUILLETT, 1899: 61. Type species: Stenoxenus johnsoni COQUILLETT, bymonotypy.

REFERENCE: WIRTH & RATANAWORABHAN, 1972b (key to females).

aductus DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 56 (female; Brazil). Distr.- Brazil(Rondônia).

arcuatus WIRTH & RATANAWORABHAN, 1972b: 1379 (female; Panama). Distr.- Panama.blantoni WIRTH & RATANAWORABHAN, 1972b: 1379 (female; Panama). Distr.- Panama.brasiliensis MACFIE, 1939b: 140 (female; Brazil). Distr.- Brazil (Amazonas, Santa Catarina),

northeastern Argentina.carrerai LANE, 1956a: 305 (as carreirai; female; Brazil). Distr.- Brazil (Goiás).dimorphus KIEFFER, 1909: 47 (male, female; Peru). Distr.- Peru.excentricus LANE, 1961c: 453 (female; Brazil). Distr.- Brazil (Pará).fulvus JOHANNSEN, 1927: 70 (female; Peru). Distr.- Peru.johnsoni COQUILLETT, 1899: 61 (female; USA, New Jersey). Distr.- USA (Missouri, New

Jersey, Texas) to Panama.lanei WIRTH & RATANAWORABHAN, 1972b: 1381 (female; Panama). Distr.- Panama.limpidus WIRTH & RATANAWORABHAN, 1972b: 1383 (female; Costa Rica). Distr.- Costa Rica.marginalis WIRTH & RATANAWORABHAN, 1972b: 1385 (female; Panama). Distr.- Panama.niger LANE, 1948: 230 (as nigrus; male, female; Brazil). Distr.- Brazil (Goiás).paraensis LANE, 1956a: 306 (female; Brazil). Distr.- Brazil (Pará).pseudofulvus SPINELLI, 1998: 54 (female; Argentina). Distr.- Northeastern Argentina,

Uruguay.setiger MACFIE, 1939b: 138 (female; Brazil). Distr.- Jamaica, Dominican Republic, Brazil

(Santa Catarina), northeastern Argentina.

Unplaced genus of Forcipomyiinae

DIDYMOPHLEPS WEYENBERGH, 1883: 108. Type species: Didymophleps hortorumWEYENBERGH, by monotypy.

Unplaced species of Forcipomyiinae

Didymophleps hortorum WEYENBERGH, 1883: 110. Argentina.

Unplaced species of Ceratopogonidae

Ceratopogon claripennis LYNCH ARRIBÁLZAGA, 1893: 229. Argentina.


7IDENTIFICATION

BORKENT & WIRTH (1997) listed the living and extinct species of biting midges for the Worldand an updated version is available on the web (http://www.inhs.uiuc.edu/cee/FLYTREE/)

CeratopogonidaeCatalog.pdf). BORKENT & SPINELLI (2000) provide a catalog of theceratopogonids south of the USA which included those species from the Nearctic portionof Mexico. Additional information was given concerning which life stages were originallydescribed and the distribution of each of the 1066 mentioned species. References to allpertinent taxonomic literature was provided for each genus, including available keys. Here,we provide an updated catalog of those species restricted to the Neotropical Region, asdefined by GRIFFITHS (1980) (Table 4). The number of species in each genus is given inTable 5. WIRTH & GROGAN (1988) provided a revision of the tribe Ceratopogonini, themost diverse tribe at the generic level. This revision included a key to the genera andsubgenera, and a diagnosis, distribution and list of included species for each genus.

The best work for identifying Neotropical Culicoides is a wing atlas published by WIRTH

et al. (1988), which contains wing photographs and a summary of the numerical charactersof the majority of the species presently known for the region. This work may be usedtogether with the revision of the genus for Panama by WIRTH & BLANTON (1959), and thekeys for the Amazon Basin by WIRTH & BLANTON (1973), for the species occurring southof the Amazon Basin by SPINELLI & WIRTH (1986b), and for Argentina (SPINELLI et al.,2005b). Description of further species may be found in SPINELLI & BORKENT (2004a) forCosta Rica, BRICKLE & HAGAN (1999) for Belize, FELIPPE-BAUER & QUINTELAS (1994) andFELIPPE-BAUER et al. (2000a, b) for Brazil, FELIPPE-BAUER et al. (2003, 2005) for Peru andPERRUOLO (1990, 2001) for Venezuela.

Adult ceratopogonids are easily collected with a variety of methods and are bestpreserved in 70% alcohol. These specimens can be identified to genus while in fluid butgenerally need to be placed on microscope slides to be identified to species. Specimens thatare not slide mounted should be pinned, as old alcohol material is impossible to slidemount and study properly. For more information, see chapter ‘Capture and study ofCeratopogonidae’.


Table 5. Numbers of named valid species in each genus of Neotropical Ceratopogonidae.

Taxon Number of species

Subfamily Leptoconopinae NOÈ

Leptoconops SKUSE 12Subfamily Forcipomyiinae LENZ

Atrichopogon KIEFFER 100Forcipomyia MEIGEN 211

Subfamily Dasyheleinae LENZ

Dasyhelea KIEFFER 58Subfamily Ceratopogoninae NEWMAN

Tribe Culicoidini KIEFFER

Culicoides LATREILLE 266Paradasyhelea MACFIE 3

Tribe CeratopogoniniAllohelea KIEFFER 1Alluaudomyia KIEFFER 19Austrohelea WIRTH & GROGAN 1Baeodasymyia CLASTRIER & RACCURT 5Baeohelea WIRTH & BLANTON 1Bahiahelea WIRTH 1Borkenthelea SPINELLI & GROGAN 4Brachypogon KIEFFER 25Cacaohelea WIRTH & GROGAN 1Ceratoculicoides WIRTH & RATANAWORABHAN 1Diaphanobezzia INGRAM & MACFIE 4Downeshelea WIRTH & GROGAN 18Echinohelea MACFIE 11Fittkauhelea WIRTH & BLANTON 1Isthmohelea INGRAM & MACFIE 1Leptohelea WIRTH & BLANTON 1Macrurohelea INGRAM & MACFIE 12Monohelea KIEFFER 18Nannohelea GROGAN & WIRTH 1Notiohelea GROGAN & WIRTH 2Parabezzia MALLOCH 24Parastilobezzia WIRTH & BLANTON 1Rhynchohelea WIRTH & BLANTON 1Schizonyxhelea CLASTRIER 2Stilobezzia KIEFFER 65

Tribe Heteromyiini WIRTH

Clinohelea KIEFFER 12Heteromyia SAY 11Pellucidomyia MACFIE 4


Physohelea GROGAN & WIRTH 2Tribe Sphaeromiini NEWMAN

Austrosphaeromias SPINELLI 2Groganhelea SPINELLI & DIPPOLITO 1Johannsenomyia MALLOCH 2Lanehelea WIRTH & BLANTON 2Mallochohelea WIRTH 6Neobezzia WIRTH & RATANAWORABHAN 8Nilobezzia KIEFFER 3Sphaerohelea SPINELLI & FELIPPE-BAUER 1

Tribe Palpomyiini ENDERLEIN

Amerohelea GROGAN & WIRTH 11Bezzia KIEFFER 46Clastrieromyia SPINELLI & GROGAN 4Pachyhelea WIRTH 2Palpomyia MEIGEN 46Phaenobezzia HAESELBARTH 2

Tribe Stenoxenini COQUILLETT

Paryphoconus ENDERLEIN 40Stenoxenus COQUILLETT 16

Unplaced genus Didymophleps WEYENBERGH 1

Unplaced species 2

Total number of species (not including unplaced taxa) 1095

Table 5. Continued.

Taxon Number of species

DIAGNOSIS OF THE CERATOPOGONIDAE

The following distinguishes Ceratopogonidae from all other Diptera: small-medium sized bit-ing midges with wing length = 0.4-7.0 mm. Ocelli absent. Antenna generally with 13 flagellom-eres (some have fewer but always 6 or more present). Most males with plumose antennae. Mostfemales with serrate mandibles. Anepisternum divided into anterior and posterior portions byanepisternal cleft. Postnotum lacking medial longitudinal groove. Wing with 1-2 radial veinsreaching margin, with two median vein branches (posterior one may be weak) reaching wingmargin. Wings overlapping each other over the abdomen in living individuals at rest (with ex-ceptions in a few species of Stilobezzia). Foreleg shorter than hind leg. Tarsomere 1 of each legequal to or longer than tarsomere 2 (with a few exceptions in Forcipomyia).

Adults are similar to some Chironomidae but may be distinguished by the combina-tion of a lack of a medial groove on the postnotum (otherwise only in Podonominae) anda lack of a m-cu crossvein (present in Podonominae).


Pupae with well-developed, undivided respiratory organs with series of small spira-cles. Third leg curled under wing sheath. Apex of the abdomen not curled under thorax.Two pointed anal processes.

Larvae with well-developed head capsule, well-developed pharyngeal complex, andno open spiracles.

Eggs are not diagnosable at this time (but see key below).

KEY TO THE GENERA OF ADULT CERATOPOGONIDAE IN THE NEOTROPICAL REGION

There are currently 1095 species of Ceratopogonidae recorded from the Neotropical Region(Table 5) but there are many more awaiting description, especially in the genera Forcipomyia,Atrichopogon, Dasyhelea, Culicoides and Stilobezzia. Species of the family may be found at all altitudesand in virtually every habitat where there is even a little moisture. The following key includesgenera known from the Neotropics and includes males, for the first time, of all genera (malesnot known for Groganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, and Sphaerohelea).

This key is intended to work for both alcohol and slide mounted material. We haveindicated in bold type those character states that are best seen with a compound microscope(i.e. with slide mounted material), although with skill and patience many of these characterstates may also be observed in alcohol specimens. Some wing veins may be difficult to seein some taxa; for alcohol specimens, tilting the wing may help to see faint wing veins andwith slide material, phase contrast microscopy provides the best resolution.

When the two claws at the end of a leg are the same size, they are referred to as beingequal; conversely when they differ in size they are called unequal. The costal ratio is thelength of the costa from the wing base (measured from the arculus), divided by the totalwing length. Terms for wing veins and cells follow the system of the Manual of NearcticDiptera (MCALPINE et al., 1981), with modifications proposed by SZADZIEWSKI (1996) andare fully labeled here in Figs. 10C, D and 11I. As is widely used by other Dipterists, veinsare in capital letters and those of cells in small letters.

1 Eyes widely separated dorsally, lacking frontal suture (Fig. 6A); palpus with 4segments (only one beyond elongate and/or swollen third) and with membranousgap between segment 3 and 4+5 (Fig. 9A); wing without crossvein r-m (Fig. 10A);female with radial cells fused into swollen unit (Fig. 10A); without palisade setaeon first tarsomere of hind leg (Fig. 14B); male gonostylus with an apical,socketed peg (Fig. 19A); female cercus very elongate (Fig. 22E) .............................................................................................................................................. Leptoconops SKUSE

– Eyes narrowly or widely separated dorsally, with frontal suture (Figs. 6B-G);palpus with 2-5 segments but never with a membranous gap between the last twosegments (Figs. 9B-E); wing with crossvein r-m (Figs. 10B-J); radial cells distinctor, if fused or absent (Figs. 11A, F), r-m clearly present; with (Fig. 14A) or without(Fig. 14B) palisade setae on first tarsomere of hind leg; male gonostylus withoutapical, socketed peg (Figs. 19B-F, 20A-F) although apex may be pointed (Fig.19E); female cercus short (Figs. 22F, 23A-E) .............................................................. 2


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

B. Culicoides ‡

Palpus

A. Leptoconops

C. Culicoides †

E. Echinohelea

D. Rhynchohelea

G. FittkauheleaF. Parabezzia

LaciniaMandibleLabrumLabium

Frontal suture

Frontal suture

Labium

Clypeus


2(1) Wing with (Figs. 10B, E, G) or without (Figs. 10D, J, 11A-C) numerousmacrotrichia on membrane; hind first tarsomere with scattered setae or, at most, arow of strong bristles (but these not forming a tight row, especially basally on thetarsomere) (Fig. 14B); female with two small, equal claws on each leg (Figs. 14C-E); with (Figs. 14C, D) or without (Figs. 14E, F) well-developed empodia .......... 3

– Wing bare or with scattered macrotrichia on apical half of wing (a few specieshave some macrotrichia more basally on the wing but these are scattered) (Figs.11E, I); hind first tarsomere with row of palisade setae with at least more basalsetae with abutting bases, some groups with an additional pronounced basal stoutspine (Figs. 14A, G-I); females with two small, equal claws on each leg or with 1-2 claws on one or more of the legs which, in some, are large (Figs. 14F-I, 15A-C,G, H, 16A-E); without well-developed empodia ......................................................... 7

3(2) Costa reaching well beyond middle of wing (Figs. 10C, D); both radial cells well-developed, with second radial cell elongate and moderately open (Figs. 10C, D);paratergite well-developed, with 1 strong and 0-5 more slender, shorter setae (Fig.9F) ...................................................................................................... Atrichopogon KIEFFER

– Costa short or long; if long, then second radial cell either wide (Figs. 10F, G) ornarrow (Figs. 10B, E); paratergite narrow (as in Fig. 9G) or well-developed,without setae or, if setae present, these number at least 10 and all are equal in sizeand length ............................................................................................................................ 4

4(3) Apical flagellomere with terminal nipple which is constricted basally (Figs. 7A, B);claws strongly curved (Figs. 14C, D); empodia vestigial or well-developed(Figs. 14C, D) ..................................................................................... Forcipomyia MEIGEN

– Apical flagellomere without terminal nipple (although it may be strongly tapered)(Fig. 7C); claws only slightly curved (Fig. 14E); empodia poorly developed ....... 5

5(4) Eyes broadly abutting medially; antennal flagellomeres without sensillacoeloconica; male with short setae on first flagellomere, less than 1/2 thelength of those on following flagellomeres (Fig. 7C); male flagellomeressculptured (Fig. 7C); without tibial spur on foreleg; wing membrane withoutdistinct pattern of dark pigmentation (Fig. 10E) .......................... Dasyhelea KIEFFER

– Eyes contiguous to widely separated medially (Figs. 6B, C); some antennalflagellomeres with sensilla coeloconica; male with setae on firstflagellomere about as long as those on following flagellomeres; maleflagellomeres not sculptured; with tibial spur on foreleg; wing membranewith or without distinct patterns of dark and/or light pigmentation (Figs. 10F-H);CULICOIDINI .................................................................................................................. 6

6(5) Wing with 1-2 radial cells; costa extending beyond 0.50 of wing length (10F, G);females with or without toothed mandibles; wing membrane with or withoutdistinct patterns of dark and/or light pigmentation (10F, G) .................................................................................................................................................. Culicoides LATREILLE

– Wing with both radial cells obliterated (Fig. 10H); costa not reaching beyond 0.43of wing length; females with reduced, vestigial mandibles; wing membranewithout distinct patterns of dark and/or light pigmentation (Fig. 10H) ...............................................................................................................................Paradasyhelea MACFIE


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

C. Dasyhelea †

D. Rhynchohelea ‡

B. Forcipomyia ‡

A. Forcipomyia ‡


7(2) M forking beyond r-m (i.e. medial fork petiolate, in some, with base quite short(Fig. 11I), in some, base of M2 not present (Figs. 10I, J, 11A) ................................ 8

– M forking at or before r-m (i.e. medial fork sessile, not petiolate) (Figs. 11G, 12F-J, 13A-I); base of M2 always present ............................................................................ 52

8(7) Wing with reduced radial cells (Figs. 11A, B, D, F); costal ratio less than 0.65 (Figs.11A, B, D, F); female claws more or less equal (the male of Rhynchohelea isunknown but likely with broadly separated eyes, antennal flagellomeres 10 and 11fused and a katepisternal seta) ........................................................................................ 9

– Wing with at least one well-developed radial cell (Figs. 10J, 11E, I); costal ratiomore than 0.50; female claws equal or unequal, some with only a single claw onone or more of the legs .................................................................................................. 15

9(8) Male antennal flagellomeres 10 and 11 fused or with less than 13flagellomeres; wing with or without radial cells; katepisternum with 1 or morelateral setae (can be seen in alcohol specimens by holding specimen upright andlooking along plane of thoracic pleura in outline) (Fig. 9G) .................................. 10

– Male antenna with flagellomeres 10 and 11 separate and with 13flagellomeres present; wing without radial cell (cells completely fused) (Figs.11A, B, D); katepisternum without setae (Fig. 9H) ............................................. 13

10(9) Female antenna with 12 flagellomeres (Fig. 7D); female labium stout, truncate,with apical hooklike structures (Fig. 6D) ...................................................................................................................................Rhynchohelea WIRTH & BLANTON (male unknown)

– Female antenna with 13 flagellomeres; female proboscis normal (Figs. 6B, C, E-G).............................................................................................................................................. 11

11(10) Male flagellum with 7-8 flagellomeres (Fig. 8A); palpus with 3 segments (Fig. 9C)............................................................................................. Nannohelea GROGAN & WIRTH

– Male flagellum with at least 9 flagellomeres (almost always with 13 but with 10and 11 fused); palpus with 5 segments ........................................................................ 12

12(11) Anepisternum lacking setae; male antenna with flagellomeres 2-6 fused (6-11also fused) .........................................................................Brachypogon KIEFFER (in part)

– Anepisternum with at least 1 seta on posterior margin (Fig. 9G); male antennawith flagellomeres 2-5 articulated, not fused (flagellomeres 7-11 fused, sometimesincluding flagellomere 6) ........................ Ceratoculicoides WIRTH & RATANAWORABHAN

13(9) Palpus with 5 segments; wing length 0.7-0.8 mm; male with well-developedparameres .................................................................................... Schizonyxhelea CLASTRIER

– Palpus with 2 segments (Fig. 9B); wing length < 0.6 mm; male without parameres.............................................................................................................................................. 14

14(13) Fore tibia with apical spur; macrotrichia confined to margin and apex of wing(in some, also a few on apices of M1 and M2); costal ratio > 0.40 (Fig. 11B); maleantenna with 6 separate flagellomeres (Fig. 8C); male with abdominal tergites 1-3pale and 4-8 brown ........................................................... Baeohelea WIRTH & BLANTON

– Fore tibia without apical spur; macrotrichia long and abundant over entirewing; costal ratio < 0.36; male antenna with 12 flagellomeres but with 2-7, 8-9,and 10-11 fused (Fig. 8B); male with abdomen uniformly brown ............................................................................................................... Baeodasymyia CLASTRIER & RACCURT


Figure 8. Antennae. A: Male Nannohelea bourioni (CLASTRIER) (Palaearctic Region). B: MaleBaeodasymyia modesta CLASTRIER and RACCURT (Neotropical Region). C: Male Baeohelea nana WIRTH &BLANTON (Neotropical Region). D: Male Borkenthelea harii (Neotropical Region). (A-B from WIRTH

& GROGAN, 1988; C courtesy of INBio, Costa Rica).

A. Nannohelea †

C. Baeohelea †

D. Borkenthelea †

B. Baeodasymyia †

Sensilla coeloconica


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

F. Atrichopogon G. Ceratoculicoides

H. Parabezzia

A. Leptoconops

C. Nannohelea D. LeptoheleaB. Baeohelea

E. Parabezzia

Fused 4+5

Paratergite

Anepisternum

Katepisternum

ForecoxaMidcoxa


15(8) Wing with characteristic pattern of darkly mottled pigmentation, more or lessscattered on membrane (Fig. 12C); thorax and at least hind femur and tibia withpatterned pigmentation; male aedeagus with distinctive posteromedial loop (Fig.21A) ....................................................................................................... Monohelea KIEFFER1

– Wing without pigmentation or, if present, restricted to radial sector or withdiscrete dots, stripes or bands (Figs. 11E, H), or if with substantial pattern ofpigmentation (Fig. 11H) then thorax and all femora and tibiae uniformly darkbrown; other taxa with variably pigmented thorax and legs; aedeagus, if present,without posteromedial loop ........................................................................................... 16

16(15) Costa extending nearly to tip of wing; female wing with costa extending beyondapex of second radial cell (Figs. 12B, D, E) (extension also in male wing ofBahiahelea and Leptohelea) ................................................................................................. 17

– Costa not extending beyond 0.85 of wing length (Figs. 11E, I); female wing withcosta ending at or extending slightly past apex of second radial cell ................... 25

17(16) Halter saclike, without constriction below knob (Fig. 13J) ...............................................................................................................................Diaphanobezzia INGRAM & MACFIE

– Halter with knob and distinct stem .............................................................................. 1818(17) Male ..................................................................................................................................... 19– Female ................................................................................................................................. 2119(18) Wing with costa not extending beyond apex of second radial cell ..........................

...................................................................................... Parastilobezzia WIRTH & BLANTON

– Wing with costa extending well beyond apex of second radial cell ..................... 2020(19) With flagellomeres 10-13 more elongate than preceding flagellomeres; antennal

plume well developed; parameres fused into large, single sphere (Fig. 19F) .............................................................................................................................. Bahiahelea WIRTH

– With flagellomeres 2-13 more or less equal in length; antennal plume short;parameres elongate, slender ........................................... Leptohelea WIRTH & BLANTON

21(18) Palpus with three segments, with last segment small and short (Fig. 9D); claws small,equal, without basal inner tooth (Fig. 15D) ................. Leptohelea WIRTH & BLANTON

– Palpus with 4-5 segments, with last segment elongate (Fig. 9E); claws variable ....... 2222(21) Clypeus not fused to head (as in Fig. 6B); wing with first radial cell small, second

elongate (Fig. 12E); with single claw (with thick basal tooth) on each leg (Fig. 15E)...................................................................................... Parastilobezzia WIRTH & BLANTON

– Clypeus laterally fused to head (Figs. 6F, G); wing with 1 radial cell (Fig. 12B); with2 claws on each leg, although hind leg claws may be unequal ............................... 23

23(22) Eyes moderately separated, bare (Fig. 6F); claws without basal inner tooth ........................................................................................................................ Parabezzia MALLOCH

– Eyes broadly separated, bare or pubescent (Fig. 6G); claws with basal inner tooth.............................................................................................................................................. 24

24(23) Palpus with 4 segments (Fig. 6G); eyes pubescent; spermathecae withoutdistinct necks .............................................................. Fittkauhelea WIRTH & BLANTON

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


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

I. Cacaohelea J. Brachypogon

A. Leptoconops

C. Atrichopogon D. Atrichopogon

F. Culicoides

G. Culicoides H. Paradasyhelea

B. Forcipomyia

E. Dasyhelea

R1R3

r-m c

A

CuP

M1

M2CuA1CuA2

anal

First radial cell Second radial cell

r3

m1

m2

cua1


Figure 11. Female wings. A: Rhynchohelea monilicornis WIRTH & BLANTON (Nearctic and NeotropicalRegions). B: Baeohelea nana WIRTH & BLANTON (Neotropical Region). C: Leptohelea micronyx WIRTH &BLANTON (Neotropical Region). D: Schizonyxhelea forattinii WIRTH & GROGAN (Neotropical Region).E: Alluaudomyia bella (COQUILLETT) (Nearctic Region). F: Brachypogon paraensis WIRTH & BLANTON

(Neotropical Region). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Alloheleajohannseni (WIRTH) (Nearctic Region). I: Stilobezzia lutea (MALLOCH) (Nearctic Region). J: Borkentheleanothofagus SPINELLI & WIRTH (Neotropical Region). (A, E, G, I from MCALPINE et al., 1981; C, D, F-H from WIRTH & GROGAN, 1988).

A. Rhynchohelea

C. Leptohelea D. Schizonyxhelea

F. Brachypogon

G. Echinohelea H. Allohelea

B. Baeohelea

E. Alluaudomyia

I. Stilobezzia J. Borkenthelea

R1 R2 R3


Figure 12. Wings. A: Female Austrohelea shannoni WIRTH & GROGAN (Neotropical Region). B:Female Parabezzia petiolata MALLOCH (Nearctic Region). C: Female Monohelea hieroglyphica KIEFFER

(Neotropical Region). D: Female Bahiahelea brasiliensis WIRTH (Neotropical Region). E: FemaleParastilobezzia leei WIRTH & BLANTON (Neotropical Region). F: Female Downeshelea sp. G: FemalePhysohelea oedidactyla (INGRAM & MACFIE) (Neotropical Region). H: Female Clinohelea bimaculata(LOEW) (Nearctic Region). I: Female Paryphoconus angustipennis ENDERLEIN (Neotropical Region). J:Male Stenoxenus coomani SÉGUY (Oriental Region) (A, E from WIRTH & GROGAN, 1988; B-C, H-Jfrom MCALPINE et al., 1981).

A. Austrohelea

C. Monohelea D. Bahiahelea

F. Downeshelea

G. Physohelea H. Clinohelea

B. Parabezzia

E. Parastilobezzia

I. Paryphoconus J. Stenoxenus †


Figure 13. Female wings. A: Stenoxenus coomani SÉGUY (Oriental Region). B: Palpomyia plebeja (LOEW)(Nearctic Region). C: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). D:Phaenobezzia pistiae (INGRAM & MACFIE) (Afrotropical Region). E: Neobezzia blantoni WIRTH &RATANAWORABHAN (Neotropical Region). F: Groganhelea rondoniensis SPINELLI & DIPPOLITO

(Neotropical Region). G: Mallochohelea smithi (Lewis) (Nearctic Region). H: Amerohelea sp.(Neotropical Region) I: Clastrieromyia schnacki SPINELLI & GROGAN (Neotropical Region). J: HalterDiaphanobezzia araucaria SPINELLI (Neotropical Region) (A-D, G from MCALPINE et al., 1981; H fromGROGAN & WIRTH, 1981; F from SPINELLI & DIPPOLITO, 1995; I from SPINELLI & GROGAN, 1986; Efrom WIRTH & RATANAWORABHAN, 1972a; J from SPINELLI, 1996).

A. Stenoxenus

C. Bezzia D. Phaenobezzia

F. Groganhelea

G. Mallochohelea H. Amerohelea

B. Palpomyia

E. Neobezzia

I. Clastrieromyia

J. Diaphanobezzia


– Palpus with 5 segments; eyes bare; spermathecae with distinct slender necks.................................................................................................................... Bahiahelea WIRTH

25(16) Male ..................................................................................................................................... 26– Female ................................................................................................................................. 3926(25) Hind leg with single long claw and small basal tooth (Fig. 14H) ..............................

................................................................................................................... Allohelea KIEFFER

– Hind leg with two equal or subequal claws ................................................................ 2727(26) Katepisternum with 1 or more lateral setae (can be seen in alcohol specimens

by holding specimen upright and looking along plane of thoracic pleura inoutline) (Fig. 9G) ...............................................................Brachypogon KIEFFER (in part)

– Katepisternum without setae .................................................................................... 2828(27) Wing with 1 radial cell (Fig. 11E) ................................................................................. 29– Wing with 2 radial cells ................................................................................................... 3329(28) Fore coxa with long spine-like or slender, tapering setae (Fig. 9H) ..........................

.............................................................................................................. Parabezzia MALLOCH

– Fore coxa with only short, simple setae (sometimes rubbed off in alcoholspecimens) ......................................................................................................................... 30

30(29) Wing with pigmentation spots at least immediately basal to or over r-m and overapex of radial cell (Fig. 11E), some with further pigmentation ............................................................................................................................................Alluaudomyia KIEFFER

– Wing without pigmentation spots ................................................................................ 3131(30) Palpus with 5 segments; hind tarsomere 1 with at least 1 1/2 rows of palisade

setae; aedeagus composed of 2 lateral sclerites (so that aedeagus appears tobe split medially) (Fig. 20E) or with aedeagus absent but with a transversebar ventral to the aedeagus (Fig. 20D) ..................... Stilobezzia KIEFFER (in part)

– Palpus with 4 segments (as in Fig. 9E); hind tarsomere 1 with only 1 row ofpalisade setae; aedeagus continuous medially (as in Fig. 21C) ......................... 32

32(31) Eyes without pubescence; genitalia with fused, very short, parameres .............................................................................. undescribed genus near Parabezzia MALLOCH

– Eyes pubescent; genitalia without parameres ................................................................................................................................................. Fittkauhelea WIRTH & BLANTON

33(28) Wing with patches of pigmentation, including discrete lighter patches on M2 andCuA2 (Fig. 12F); aedeagus continuous medially (Fig. 20F) ............................................................................................................................... Downeshelea WIRTH & GROGAN

– Wing with or without pattern of pigmentation but if pattern present, thenwithout discrete patches on M2 and CuA2 (posterior portion of wing may begenerally more darkly pigmented) (Figs. 11J, 12A) and with aedeagus dividedmedially (Figs. 20B, C) .................................................................................................. 34

34(33) Antenna with apical 5 flagellomeres more elongate than basal 8 flagellomeres(female-like antenna), without plume .......................... Cacaohelea WIRTH & GROGAN

– Antenna with apical 3 flagellomeres more elongate than basal 10 flagellomeres(male-like antenna), with or without plume ............................................................... 35

35(34) Radial cells separated by a short longitudinal vein (R2); wing without macrotrichia;sternite 9 elongate (Fig. 20C) ......................................... Isthmohelea INGRAM & MACFIE


– Both radial cells close to one another, separated by a short transverse vein (R2)(Figs. 11J, 12A); wing with or without macrotrichia; sternite 9 short or moderatelyelongate (Figs. 20D, E) .................................................................................................... 36

36(35) Flagellomeres 3-10 fused (Fig. 8D) ........................ Borkenthelea SPINELLI & GROGAN

– Antenna with flagellomeres separate, not fused ........................................................ 3737(36) Length of tarsomeres 2-5 of hind leg about 1.4-1.7as long as length of tarsomere

1 .......................................................................................... Austrohelea WIRTH & GROGAN

– Length of tarsomeres 2-5 of hind leg about as long as tarsomere 1 ................... 3838(37) Hind tarsomere 1 with at least 1 1/2 rows of palisade setae; wing with or without

macrotrichia; aedeagus composed of 2 lateral sclerites (so that aedeagusappears to be split medially) (Fig. 20E) or with aedeagus absent but with atransverse bar ventral to the aedeagus (Fig. 20D); antennal flagellomereswithout sensilla coeloconica ........................................................Stilobezzia KIEFFER

– Hind tarsomere 1 with only 1 row of palisade setae (as in Fig. 14A); wing withoutmacrotrichia; aedeagus undivided, somewhat triangular; some antennalflagellomeres with sensilla coeloconica ............. Macrurohelea INGRAM & MACFIE

39(25) Katepisternum with 1 or more lateral setae (can be seen in alcohol specimensby holding specimen upright and looking along plane of thoracic pleura inoutline) (Fig. 9G) ...............................................................Brachypogon KIEFFER (in part)

– Katepisternum without seta ...................................................................................... 4040(39) Wing with 1 radial cell (Figs. 11C, E) .......................................................................... 41– Wing with 2 radial cells (first may be small) (Figs. 10I, 11H, I) ............................ 4341(40) Palpus with 4 segments (as in Fig. 9E); claws on hind legs equal .............................

.................................................................. undescribed genus near Parabezzia MALLOCH

– Palpus with 5 segments; claws on hind legs a single talon or unequal (Figs. 14F,15A-C) ................................................................................................................................ 42

42(41) Wing with pigmentation spots immediately basal to or over r-m and over apex ofradial cell (Fig. 11E); hind tarsomere 1 with only one row of palisade setae .....................................................................................................................Alluaudomyia KIEFFER

– Wing without pigmentation spots; hind tarsomere 1 with at least 1 1/2 rows ofpalisade setae ....................................................................... Stilobezzia KIEFFER (in part)

43(40) Tenth segment of female elongated and bent anteroventrally (Fig. 22F) ..........................................................................................................Macrurohelea INGRAM & MACFIE

– Tenth segment of female not elongated, directed posteriorly ............................... 4444(43) Wing with patches of pigmentation, either 2-3 dark bands (Fig. 11H) or at least

with patches of pigmentation at r-m and just posterior to the apex of the secondradial cell ............................................................................................................................ 45

– Wing without patches of discrete pigmentation ....................................................... 4745(44) Fore- and midlegs each with one claw (each with a smaller basal tooth) ................

................................................................................................ Stilobezzia KIEFFER (in part)– Fore- and midlegs each with 2 equal claws ................................................................ 4646(45) Hind claw with basal tooth (Fig. 14H) ............................................. Allohelea KIEFFER

– Hind claw without basal tooth (Fig. 14I) .................. Downeshelea WIRTH & GROGAN

47(44) Fore- and midlegs each with 1 claw (some with a smaller basal tooth) ............... 48


– Fore- and midlegs each with paired, equal, or unequal-sized claws (as in Fig. 14F).............................................................................................................................................. 49

48(47) First radial cell small, with R3 thickened (Fig. 10I); second radial cell somewhatbroad; medial vein reduced to faint trace on basal portion; single claw on each legshort, stout and sharp (Fig. 15F).. ................................ Cacaohelea WIRTH & GROGAN

– First radial cell small to well-developed, with R3 at most thickened at apex (Fig.11I); second radial cell narrow; medial veins clearly visible but with base of M2sometimes absent; claw on at least hind leg elongate (Figs. 15A-C) ........................................................................................................................ Stilobezzia KIEFFER (in part)

49(47) Radial cells separated by a short longitudinal vein (R2); wing with a fewmacrotrichia restricted to extreme periphery at tip; antennal flagellomereswithout sensilla coeloconica ..................................... Isthmohelea INGRAM & MACFIE

– Radial cells separated by a short transverse vein (R2); wing with or withoutmacrotrichia on apical 1/4; some antennal flagellomeres with sensillacoeloconica (either only on flagellomere 1 or only on flagellomeres 8 and12) ........................................................................................................................................ 50

50(49) Wing without macrotrichia; mandibles reduced, vestigial; eyes bare orpubescent; claws small, equal in all legs ..................... Notiohelea GROGAN & WIRTH

– Wing with macrotrichia on apical 1/4; mandibles toothed; eyes pubescent;only fore- and midlegs with small, equal claws, or claws unequal on all legs (as inFig. 14F) ............................................................................................................................. 51

51(50) Radial cells subequal, small; R3 thickened; midleg claws unequal and somewhatsimilar to those of hind leg; sensilla coeloconica only on flagellomere 1 ................................................................................................ Borkenthelea SPINELLI & GROGAN

– Second radial cell about twice as long as first radial cell; R3 not thickened; midlegwith small, equal claws, much shorter than elongate single claw (with basal tooth)of hind leg; sensilla coeloconica only on flagellomeres 8 and 12 .................................................................................................................. Austrohelea WIRTH & GROGAN

52(7) Tarsomere 5 of foreleg swollen (Figs. 15G, 17A, C); HETEROMYIINI ........... 53– Tarsomere 5 of foreleg not swollen ............................................................................. 5653(52) M barely sessile, forking at r-m (Fig. 12G) .................. Physohelea GROGAN & WIRTH

– M broadly sessile, forking before r-m (Fig. 12H) ...................................................... 5454(53) Fore femur swollen, with numerous thick ventral spines (Fig. 17C) ........................

........................................................................................................................ Heteromyia SAY

– Fore femur slender, with at most a few slender spines ........................................... 5555(54) Fourth tarsomere of all legs cylindrical or slightly cordate and lacking strong

spines; female hind leg strikingly elongate, with tarsomere 2 at least 3/4 the lengthof tarsomere 1 (Fig. 15H); hind claw very long, either single or with a small basaltooth, much longer than claws of fore and midlegs (Fig. 15H) .............................................................................................................................................. Pellucidomyia MACFIE

– Fourth tarsomere of foreleg cordate of mid and hind leg bifid and spinose (Fig.17B); female hind leg not much longer than other legs, with tarsomere 2 less thantwice the length of tarsomere 1; both hind claws about same length as those onfore- and midleg ................................................................................... Clinohelea KIEFFER


Figure 14. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Ceratopogon MEIGEN

first tarsomere. B: Forcipomyia sp. first tarsomere. C: Atrichopogon levis (COQUILLETT) (NearcticRegion). D: Forcipomyia fusicornis (COQUILLETT) (Nearctic and Neotropical Regions). E: Dasyheleapseudoincisurata WAUGH & WIRTH (Nearctic Region). F: Alluaudomyia bella (COQUILLETT) (NearcticRegion). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Allohelea neotropicaWIRTH (Neotropical Region). I: Downeshelea sp. (C-E, G from MCALPINE et al., 1981; F, H-I fromWIRTH & GROGAN, 1988).

A. Most Ceratopogoninae

C. Atrichopogon D. Forcipomyia

F. Alluaudomyia

G. Echinohelea H. Allohelea

B. Other subfamilies

E. Dasyhelea

I. Downeshelea

Palisade setae

Empodium

H

H

H

H

H

HMF

H H H


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

HMFHMF HM

F

HF

HM

F HMF

H

MF

A. Stilobezzia

C. Stilobezzia D. Leptohelea

F. Cacaohelea G. Clinohelea

H. Pellucidomyia

B. Stilobezzia

E. Parastilobezzia


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

H

MF

HMF

H

H

H

M

F

A. Neobezzia

C. Groganhelea

D. Groganhelea

B. Neobezzia

E. Phaenobezzia


Figure 17. Female fifth tarsomeres (F = foreleg, M = midleg, H = hind leg). A: Clinohelea bimaculata(LOEW) (Nearctic Region). B: Clinohelea bimaculata (LOEW) (Nearctic Region). C: Heteromyia fasciataSAY (Nearctic Region). D: Heteromyia fasciata SAY (Nearctic Region). E: Heteromyia fasciata SAY

(Nearctic Region). F: Stenoxenus coomani SÉGUY (Oriental Region). G: Johannsenomyia argentata (LOEW)(Nearctic Region). H: Johannsenomyia argentata (LOEW) (Nearctic Region). I: Johannsenomyia argentata(LOEW) (Nearctic Region) (all from MCALPINE et al., 1981).

F

F

H

F

M

H

H

HM

A. Clinohelea

C. Heteromyia D. Heteromyia

F. Stenoxenus G. Johannsenomyia

H. Johannsenomyia

B. Clinohelea

E. Heteromyia

I. Johannsenomyia


56(52) Male genitalia large, with gonocoxites fused dorsally and ventrally (Fig. 21B);female genitalia with sternite 8 large and plate like (Fig. 23A) ................................................................................................................................................... Echinohelea MACFIE

– Male genitalia small or large, if large then gonocoxites not fused; female genitaliawith sternite 8 moderately developed .......................................................................... 57

57(56) Male (male of Groganhelea, Sphaerohelea are unknown) .............................................. 58– Female ................................................................................................................................. 7258(57) Gonostylus much reduced in size (Figs. 21D, 22B) or fused with gonocoxite (Fig.

22D) .................................................................................................................................... 59– Gonostylus large, clearly present (Figs. 21F, 22A, C) (male Lanehelea are not

known but likely key to at least this point) ................................................................. 6159(58) Wing with 2 clearly defined radial cells (Fig. 13I); gonostylus well- defined and

with pointed apex (Fig. 21D) .................................Clastrieromyia SPINELLI & GROGAN

– Wing with 1 radial cell or with, at most, a faint indication of a division betweenfirst and second radial cell (Fig. 13D); gonostylus either small and apicallyrounded (Fig. 22B) or not clearly defined and fused with gonocoxite (Fig. 22D).............................................................................................................................................. 60

60(59) Anepisternum with numerous elongate setae; gonocoxite short and gonostyluspresent but small Fig. 22B) ................................................. Phaenobezzia HAESELBARTH

– Anepisternum without setae; gonocoxite and gonostylus fused and elongate (Fig.22D) .......................................................................................................Nilobezzia KIEFFER

61(58) Genitalia with gonocoxite and tergite 9 elongate (Fig. 22C) .................................. 62– Genitalia more stout, with gonocoxite and tergite 9 not unusually elongate (Figs.

21F, 22A) ............................................................................................................................ 6462(61) Wing with 1 radial cell (as in Fig. 13D) .......Neobezzia WIRTH & RATANAWORABHAN

– Wing with 2 radial cells (Fig. 13G) ............................................................................... 6363(62) Forefemur with ventral thick spines .................................. Austrosphaeromias SPINELLI

– Forefemur without ventral thick spines ....................................... Mallochohelea WIRTH

(Note. Some Mallochohelea species have males with non-elongate genitalia)64(61) Tarsomere 5 of at least hind leg with stout, blunt spines (Fig. 17H); apex of tergite

9 with single row of thick, stout setae (Fig. 21C) .............. Johannsenomyia MALLOCH

– Tarsomere 5 without thick spines, or if present, these more slender and sharplypointed (Fig. 16A); tergite 9 with scattered setae, never arranged in a single row ofthick, stout setae ............................................................................................................... 65

65(64) With 1-2 stout setae on apex of the cercus (Fig. 21E); parameres fused, withfusion restricted to most anteromedial portion (Fig. 21E) ............................................................................................................................... Amerohelea GROGAN & WIRTH

– With several to many setae on the cercus; parameres separate or with fusionvarious ............................................................................................................................... 66

66(65) Forefemur with stout spines (Fig. 18A) ...................................................................... 67– Forefemur without stout spines .................................................................................... 6867(66) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part)– Wing with 2 radial cells (Fig. 13B) ................................... Palpomyia MEIGEN (in part)68(66) Palpus with 4 segments ............................................................... Stenoxenus COQUILLETT


– Palpus with 5 segments ................................................................................................... 6969(68) Thorax slender and dorsoventrally flattened; paratergite with numerous small

setae; 1 radial cell (Fig. 12I) .....................................................Paryphoconus ENDERLEIN

– Thorax stouter and not dorsoventrally flattened; paratergite without setae; 1-2radial cells .......................................................................................................................... 70

70(69) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part)– Wing with 2 radial cells (Fig. 13B) ............................................................................... 7171(70) Hind femur swollen (Fig. 18C); abdomen white, contrasting with dark thorax .....

.....................................................................................................................Pachyhelea WIRTH

– Hind femur not swollen (Fig. 18A); abdomen light to dark brown, similar to darkthorax ...................................................................................... Palpomyia MEIGEN (in part)

72(57) Wing with M2 strongly curving near base (Fig. 13A); palpus with 4 segments ................................................................................................................ Stenoxenus COQUILLETT

– Wing with M2 with not or only slightly curved near base (Figs. 13B-I); palpus with5 segments ......................................................................................................................... 73

73(72) Tarsomere 5 armed ventrally with stout black blunt spines (Figs. 16A, B, 17G-I)(these spines somewhat sharp pointed in Austrosphaeromias); abdomen withouttergal apodemes; sternite 8 with or without (Sphaerohelea only) postero-lateral hair tufts, with long, medially overlapping, curved setae (bestexamined at 100X) (Fig. 23B); ....................................... SPHAEROMIINI..........74

– Tarsomere 5 unarmed, or if with some ventral spines, these are slender and withsharp tips (Fig. 16E); abdomen with (Figs. 23C-E) or without tergal apodemes(not visible in some teneral or young specimens); sternite 8 without hair tufts.............................................................................................................................................. 81

74(73) Femora without spines; claws equal on foreleg, unequal on mid- and hind legs(Figs. 17G-I) ............................................................................... Johannsenomyia MALLOCH

– Femora with or without spines; claws equal on all legs or with only forelegsunequal ............................................................................................................................... 75

75(74) Claws gently curved distally, with slender basal tooth on internal side ............... 76– Claws straight or flattened distally, with blunt or slender external basal tooth (Figs.

16A, B) ............................................................................................................................... 7876(75) Forefemur swollen, with numerous thick ventral spines .............................................

....................................................................................................Austrosphaeromias SPINELLI

– Fore femur slender, unarmed ........................................................................................ 7777(76) Costa extending to wing tip; tarsomere 5 with 2-3 pairs of ventral spines .............

............................................................................. Sphaerohelea SPINELLI & FELIPPE-BAUER

– Costa not extending beyond 0.85 of wing length; tarsomere 5 with 4-5 pairs ofventral spines ......................................................................Lanehelea WIRTH & BLANTON

78(75) Wing with 1 radial cell; costa long, extending nearly to wing tip with costal ratioover 0.87; femora without ventral spines .................................................................... 79

– Wing with 1-2 radial cells; costa short, extending to less than 0.84 of wing length;femora with or without ventral spines ......................................................................... 80


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

A. Palpomyia

C. PachyheleaB. Bezzia

HMF

HF


Figure 19. Male genitalia, ventral view. A: Leptoconops torrens (TOWNSEND) (Nearctic Region). B:Forcipomyia acidicola (TOKUNAGA) (Holarctic Region). C: Dasyhelea pseudoincisurata WAUGH & WIRTH

(Nearctic Region). D: Culicoides haematopotus MALLOCH (Nearctic and Neotropical Regions). E:Stenoxenus coomani SÉGUY (Oriental Region). F: Bahiahelea brasiliensis WIRTH (Neotropical Region) (A-E from Manual of Nearctic Diptera; F from BORKENT, unpubl.).

A. Leptoconops

C. Dasyhelea D. Culicoides

F. Bahiahelea

B. Forcipomyia

E. Stenoxenus

Parameres

Parameres

Parameres

Parameres

Parameres

Aedeagus


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

A. Leptohelea

C. Isthmohelea D. Stilobezzia

F. Downeshelea

B. Cacaohelea

E. Stilobezzia

Aedeagus

Parameres

Gonocoxite

Gonostylus

Cercus


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

Aedeagus

Gonostylus

Paramer

A. Monohelea

C. Johannsenomyia D. Clastrieromyia

F. Paryphoconus

B. Echinohelea

E. Amerohelea


79(78) External basal tooth of tarsal claws pointed (Fig. 16A); tarsus of hind leg notelongate (Fig. 16B); wing with anal lobe poorly developed ......................................................................................................................Neobezzia WIRTH & RATANAWORABHAN

– External basal tooth of tarsal claws blunt (Fig. 16C); tarsus of hind leg elongate(Fig. 16D); wing with anal lobe well developed (Fig. 13F) ............................................................................................................................... Groganhelea SPINELLI & DIPPOLITO

80(78) Wing with 2 radial cells (Fig. 13G); wing grayish with veins darker brown, anteriorveins prominent; forefemur without ventral thick spines ........ Mallochohelea WIRTH

– Wing with 1 radial cell; wing whitish, anterior veins pale; forefemur with ventralthick spines ...........................................................................................Nilobezzia KIEFFER

81(73) Thorax slender and dorsoventrally flattened; paratergite with numerous smallsetae; 1 radial cell (Fig. 12I) .....................................................Paryphoconus ENDERLEIN

– Thorax stouter and not dorsoventrally flattened; paratergite without setae; wingwith 1-2 radial cells .................................................................. PALPOMYIINI..........82

82(81) Abdomen with 1 pair of tergal apodemes arising near lateral margins of sternite 7;1 spermatheca present, with no evidence of a reduced second one ..................................................................................................................... Amerohelea GROGAN & WIRTH

– Abdomen with 1 or more pairs of tergal apodemes (Figs. 23C-E), if only 1 pairpresent, these arise from more medially on posterior margin of sternite 7; 1-2spermathecae but if apparently only one, a small reduced second one is present.............................................................................................................................................. 83

83(82) Wing with 2 radial cells (Fig. 13B) ............................................................................... 84– Wing with 1 radial cell (Fig. 13C) ................................................................................. 8684(83) Forefemur without stout spines; hind femur greatly swollen (Fig. 18C); abdomen

white, contrasting with dark thorax ....................................................Pachyhelea WIRTH

– Forefemur with or without stout spines (Fig. 18A); hind femur not greatly swollen(Fig. 18A); abdomen light to dark brown, similar to dark thorax ......................... 85

85(84) Flagellum short, only 1.2-1.4 times longer than the breadth of the head; wingmembrane unpigmented; anal lobe of wing well-developed (Fig. 13I); abdomenwith single pair of apodemes arising from the anterior margin of tergite 7 andextending to about the anterior margin of tergite 6 (Fig. 23C) .......................................................................................................................Clastrieromyia SPINELLI & GROGAN

– Flagellum elongate, at least 2.5 times longer than the breadth of the head; wingmembrane pigmented; anal lobe of wing poorly developed (Fig. 13B); abdomenwith 2 or more pairs of elongate apodemes arising from the anterior margin of atleast tergites 6-7 and extending much further anteriorly into abdominal cavity(Fig. 23E) ............................................................................................... Palpomyia MEIGEN

86(83) Costa moderately short with costal ratio 0.67-0.75 (Fig. 13C); forefemur with (Fig.18B) or without stout spines; tarsomere 5 with or without stout ventral setae ............................................................................................................ Bezzia KIEFFER (in part)

– Costa more elongate with costal ratio more than 0.80 (Fig. 13D); forefemurwithout stout spines; tarsomere 5 with slender ventral setae with sharp, bent tips(Fig. 16E) ................................................................................ Phaenobezzia HAESELBARTH


KEY TO THE GENERA OF EGGS OF CERATOPOGONIDAE IN THE NEOTROPICAL REGION

There are only a few species of Ceratopogonidae for which the eggs have been described(Table 1). The following key is therefore very tentative.

There are no keys to the eggs of Diptera which would allow those of Cerato-pogonidae to be distinguished from those of other families. Nevertheless, someCeratopogonidae have eggs which are either different from those of all other Diptera(Dasyhelea, Stenoxenus and Paryphoconus) or are laid in a distinctive matrix (Alluaudomyia andmost Sphaeromiini).

1 Egg C-shaped (Fig. 24A) .................................................................................... Dasyhelea– Egg with, at most, a slight curvature (Figs. 24B-F) .................................................... 22(1) Egg with distinctive subapical frill and terminal nipple-like constriction at anterior

end (Fig. 24F) ........................................................................ Stenoxenus and Paryphoconus– Egg without any well developed ornamentation at either end; small spicules may

be present (Figs. 24A-E) ................................................................................................... 33(2) Egg oval (Fig. 24B) ............................ Leptoconops (in part), Forcipomyiinae (in part)– Egg elongate, slender (Figs. 24C-E) ............................................................................... 44(3) Eggs arranged in an elongate ribbon, with eggs lying at an angle to the length of

the ribbon (Fig. 24E) ............................................................................................................................. most Sphaeromiini (known for Johannsenomyia, Mallochohelea, Macropeza)

– Eggs laid separately or in group; if in a ribbon, eggs are laid side by side (Figs.24C-D) .................................................................................................................................. 5

5(4) Eggs laid in ribbon (Fig. 24D) ..................................... Alluaudomyia (at least in part)– Eggs laid singly or in a loose group (but not attached to one another

longitudinally) (Fig. 24C) .......................................................... other Ceratopogonidae

KEY TO THE GENERA OF LARVAE OF CERATOPOGONIDAE IN THE NEOTROPICAL REGION

The immatures of Ceratopogonidae are among the most poorly known of all nematocer-ous families (Table 2). Here we provide a tentative key to the larvae of some commongenera. A world key to the genera of larvae and pupae is being prepared by the seniorauthor. It is important to realize that only 11 of 51 known genera of NeotropicalCeratopogonidae have been described as larvae in this region (Table 2), although anadditional 11 genera are known as larvae from elsewhere.

1 Proleg present on prothorax (Figs. 25B, C); head capsule hypognathous toprognathous (Figs. 25B, C); with strong setae on head capsule and body (Figs. 25B,C) (Forcipomyiinae) ........................................................................................................... 2

– Without proleg on thorax (Figs. 25A, D-F); head capsule somewhat hypognathousor prognathous (Figs. 25A, D-F); without strong setae on head capsule and bodyanterior to anal segment (Figs. 25A, D-F) .................................................................... 3

2(1) Body somewhat flattened dorsoventrally, with lateral tubercles (Fig. 25B) .......................................................................................................................................... Atrichopogon


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

Gonostylus

Parameres

Parameres

Parameres

CercusTenth segment

A. Bezzia

C. Mallochohelea D. Nilobezzia

F. Macrurohelea

B. Phaenobezzia

E. Leptoconops


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

Sternite 8

Sternite 8

Sternite 9Sternite 10

Cercus

Tergal apodeme

Tergal apodeme

A. Echinohelea

C. Clastrieromyia

D. Paryphoconus

B. Probezzia

E. Palpomyia


Figure 24. Eggs. A: Dasyhelea flavifrons (GUÉRIN) (Palearctic Region). B: Forcipomyia mcateei WIRTH

(Nearctic Region). C: Culicoides pulicaris (LINNAEUS) (Palearctic Region). D: Alluaudomyia needhamiTHOMSEN (Nearctic Region). E: Probezzia concinna (GUÉRIN) (Palearctic Region). F: Paryphoconusangustipennis ENDERLEIN (Neotropical Region). (A from SZADZIEWSKI et al., 1997; B from CHAN &LEROUX, 1965 (as F. saundersi); C from GUTSEVICH & GLUKHOVA, 1970; D from THOMSEN, 1937; Efrom WESENBERG-LUND, 1914; F from MAYER, 1952).

A. Dasyhelea

C. Culicoides D. Alluaudomyia

F. Paryphoconus

B. Forcipomyia

E. Probezzia


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

A. Leptoconops

C. Forcipomyia

D. Dasyhelea

F. Bezzia

B. Atrichopogon

E. Culicoides

Posterior proleg

Posterior proleg

Prothoracic proleg

Prothoracic proleg

Posterior proleg


– Body cylindrical in cross-section, without lateral tubercles (Fig. 25C) ........ Forcipomyia3(1) Head capsule directed anteroventrally; end of abdomen with series of hooks,

sometimes arranged as two prolegs, (often retracted and difficult to see in alcoholmaterial), never with elongate setae (Fig. 25D) ............................................. Dasyhelea

– Head capsule prognathous; end of abdomen simple, without hooks, with orwithout elongate setae (Figs. 25A, E, F) ....................................................................... 4

4(3) Head capsule somewhat poorly developed pigmented, with well-developed,posteriorly directed apodemes extending into prothorax (Figs. 26A, B); abdomenwith segments secondarily divided (Fig. 25A) ............................................ Leptoconops

– Head capsule well-developed, lacking apodemes posterior to margin of headcapsule (Figs. 26C-F); abdomen with segments normal, not secondarily divided(Figs. 25E, F) (Ceratopogoninae) ................................................................................... 5

5(4) Head capsule with all setae simple (Figs. 26C, D) ................................. Culicoides– Head capsule with one or more of setae s, u and posterior o plumose (Figs.

26E, F; seta u not shown, it is generally lateral on head capsule) ........................... 66(5) Posteroventral margin of head capsule with posteriorly projecting process (Fig.

26F) ................................................................................................. Sphaeromiini (in part)– Posteroventral margin of head capsule a straight or somewhat curved line,

without posteriorly projecting process (Fig. 26E) ............................................................................................................ Sphaeromiini (in part), Palpomyiini, Ceratopogonini

CLAVE PARA LOS GÉNEROS DE ADULTOS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL

Existen hasta la fecha 1095 especies de Ceratopogonidae descriptas de la región Neotropical(Tabla 5), aunque muchas más aguardan para a ser descriptas, particularmente en los génerosForcipomyia, Atrichopogon, Dasyhelea, Culicoides y Stilobezzia. Especies de esta familia pueden serhalladas en todas las altitudes y virtualmente en todo hábitat donde exista por lo menos algode humedad. La siguiente clave incluye a los géneros conocidos de la región Neotropical y,por primera vez, a los machos de todos los géneros (no son conocidos los machos deGroganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, y Sphaerohelea)

Esta clave está destinada a identificar tanto material conservado en alcohol comomaterial montado en lámina. Hemos señalado en negrita los caracteres que se observanmejor mediante el uso de microscopio (por ej., en láminas), aunque con práctica ypaciencia muchos de estos caracteres también pueden ser observados en materialconservado en alcohol. En algunos grupos, algunas nervaduras de las alas puedes serdificiles de observar; con ejemplares conservados en alcohol, es posible reconocer débilestrazos de las nervaduras al examinarlos de perfil a la vez que se inclina levemente el ala,mientras que con material montado en lámina, la mejor resolución es la que brinda elmicroscopio de contraste de fase.

Cuando las dos uñas al final de la pata son del mismo tamaño, se las refiere comoiguales; cuando son de diferente tamaño, como desiguales. La relación costal es el largo dela costa desde la base del ala (medida desde el arculus), dividido por el largo total del ala.La terminología adoptada para las nervaduras y celdas de las alas sigue aquella del“Manual of Nearctic Diptera” (MCALPINE et al., 1981), con las modificaciones propuestas


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

A. Leptoconops

C. Culicoides D. Culicoides

F. Nilobezzia

B. Leptoconops

E. Nilobezzia

Apodemes

Seta s

Projecting process

Seta o


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

1 Ojos ampliamente separados, sin sutura frontal (Fig. 6A); palpo con 4 segmentos(sólo uno más allá del tercero, el cual es alargado o hinchado) y con intervalomembranoso entre los segmentos 3 y 4+5 (Fig. 9A); ala sin nervadura transversalr-m (Fig. 10A); hembra con las celdas radiales fusionadas en un estigma (Fig.10A); sin hileras de setas empalizadas en el primer tarsómero de la pata posterior(Fig. 14B); gonostilo del macho con una púa alveolada apical (Fig. 19A);cerco de la hembra muy alargado (Fig. 22E) .................................. Leptoconops SKUSE

– Ojos estrecha o anchamente separados en el dorso, con sutura frontal (Figs. 6B-G); palpo con 2-5 segmentos, pero nunca con intervalo membranoso entre los dosúltimos segmentos (Figs. 9B-E); ala con nervadura transversal r-m (Figs. 10B-J);celdas radiales distintivas o, si están fusionadas o ausentes (Figs. 11A, F), lanervadura transversal r-m está claramente presente; con (Fig. 14A) o sin (Fig.14B) hileras de setas empalizadas en el primer tarsómero de la pata posterior;gonostilo del macho sin púa alveolada apical (Figs. 19B-F, 20A-F) aunqueel ápice puede ser puntiagudo (Fig. 19E); cerco de la hembra corto (Figs.22F, 23A-E) ......................................................................................................................... 2

2(1) Ala con (Figs. 10B, E, G) o sin (Figs. 10D, J, 11A-C) numerosas macrotriquias enla membrana; primer tarsómero posterior con setas dispersas o, a lo sumo, conuna hilera de pelos robustos (pero éstos no forman un hilera apretada,especialmente en la base del tarsómero) (Fig. 14B); hembra con dos uñaspequeñas, iguales, en cada pata (Figs. 14C-E); con (Figs. 14C, D) o sin (Figs. 14E,F) empodio bien desarrollado ......................................................................................... 3

– Ala desnuda o con macrotriquias dispersas en la mitad distal del ala (unas pocasespecies tienen algunas macrotriquias en la base del ala, pero éstas se hallandispersas) (Figs. 11E, I); primer tarsómero posterior con hilera de setas con almenos más setas basales cuyas bases contactan, algunos grupos con una fuerteespina basal adicional (Figs. 14A, G-I); hembra con dos uñas pequeñas, iguales, encada pata, o con 1-2 uñas en una o más patas, las cuales en algunas son largas(Figs. 14F-I, 15A-C, G, H, 16A-E); sin empodio bien desarrollado ....................... 7

3(2) Costa sobrepasando la mitad del largo del ala (Figs. 10C, D); ambas celdas radialesbien desarrolladas, con la segunda celda radial larga y abierta moderadamente(Figs. 10C, D); paratergito bien desarrollado, con una seta fuerte y 0-5 setas finasmás cortas (Fig. 9F) ........................................................................ Atrichopogon KIEFFER

– Costa corta o larga; si es larga, entonces la seguna celda radial es ancha (Figs. 10F,G) o angosta (Figs. 10B, E); paratergito angosto (como en Fig. 9G) o biendesarrollado, sin setas o, si hay setas presentes, son al menos 10 e iguales entamaño y longitud .............................................................................................................. 4

4(3) Flagelómero apical con pezón terminal, el cual es estrechado basalmente (Figs.7A, B); uñas fuertemente curvas (Figs. 14C, D); empodio vestigial o biendesarrollado (Figs. 14C, D) ........................................................... Forcipomyia MEIGEN


– Flagelómero apical sin pezón terminal (aunque puede ser fuertementeadelgazado) (Fig. 7C); uñas sólo ligeramente curvas (Fig. 14E); empodiopobremente desarrollado .............................................................................................. 5

5(4) Ojos anchamente colindantes en la línea media; flagelómeros antenales sinsensilas celocónicas; macho con setas cortas en el primer flagelómero,menores a la mitad del largo de aquellas de los flagelómeros subsiguientes(7C); flagelómeros del macho esculturados (Fig. 7C); sin espolón en la pataanterior; membrana del ala sin diseño distintivo de pigmentación oscura (Fig.10E) ......................................................................................................... Dasyhelea KIEFFER

– Ojos contiguos a anchamente separados en la línea media (Figs. 6B, C); algunosflagelómeros antenales con sensilas celocónicas; macho con setas delprimer flagelómero más o menos del mismo largo que aquellas de losflagelómeros subsiguientes; flagelómeros del macho no esculturados; conespolón tibial en la pata anterior; membrana del ala con o sin diseñosdistintivos de pigmentación oscura (Figs. 10F-H); CULICOIDINI ....................... 6

6(5) Ala con 1-2 celdas radiales; costa sobrepasando la mitad del largo del ala (Figs.10F, G); hembras con o sin mandíbulas dentadas; membrana alar con o sindiseños distintivos de pigmentación oscura y/o clara (Figs. 10F, G) ...................................................................................................................................... Culicoides LATREILLE

– Ala con ambas celdas radiales obliteradas (Fig. 10H); costa no alcanzándo más alláde 0,43 del largo del ala; hembras con mandíbulas reducidas, vestigiales;membran alar sin diseños distintivos de pigmentación oscura y/o clara (Fig. 10H)..............................................................................................................Paradasyhelea MACFIE

7(2) M bifurcándose más allá de r-m (v.g. bifurcación mediana peciolada, en algunos debase muy corta) (Fig. 11I), en algunos la base de M2 no está presente (Figs. 10I, J,11A) ....................................................................................................................................... 8

– M bifurcándose al nivel o anterior de r-m (v.g. bifurcación mediana sésil, nopeciolada) (Figs. 11G, 12F-J, 13A-I); base de M2 siempre presente ...................... 52

8(7) Ala con celdas radiales reducidas (Figs. 11A, B, D, F); relación costal menor que0,65 (Figs. 11A, B, D, F); uñas de la hembra más o menos iguales (el macho deRhynchohelea es desconocido, pero probablemente con ojos anchamente separados,flagelómeros antenales 10 y 11 fusionados, y con una seta katepisternal) ............ 9

– Ala con al menos una celda radial bien desarrollada (Figs. 10J, 11E, I); relacióncostal mayor que 0,50; uñas de la hembra iguales o desiguales, algunas con sólouna uña en una o más de las patas ............................................................................... 15

9(8) Macho con los flagelómeros antenales 10 y 11 fusionados o con menos de 13flagelómeros; ala con o sin celdas radiales; katepisterno con 1 o más setaslaterales (pueden ser observadas en ejemplares preservados en alcohol,tomándolos verticalmente y observándolos de perfil a lo largo del plano de lapleura torácica) (Fig. 9G) ................................................................................................ 10

– Antena del macho con flagelómeros 10 y 11 separados y con 13flagelómeros presentes; ala sin celda radial (celdas completamente fusionadas)(Figs. 11A, B, D); katepisterno sin setas (Fig. 9H) ................................................ 13


10(9) Antena de la hembra con 12 flagelómeros (Fig. 7D); labio de la hembra robusto,truncado, con estructuras apicales en forma de gancho (Fig. 6D) ............................................................................. Rhynchohelea WIRTH & BLANTON (macho desconocido)

– Antena de la hembra con 13 flagelómeros; proboscis de la hembra normal (Figs.6B, C, E-G) ........................................................................................................................ 11

11(10) Flagelo del macho con 7-8 flagelómeros (Fig. 8A); palpo con 3 segmentos (Fig.9C) ...................................................................................... Nannohelea GROGAN & WIRTH

– Flagelo del macho con al menos 9 flagelómeros (casi siempre con 13, pero con 10y 11 fusionados); palpo con 5 segmentos ................................................................... 12

12(11) Anepisterno sin setas; antena del macho con flagelómeros 2-6 fusionados (6-11 también fusionados) .............................................. Brachypogon KIEFFER (en parte)

– Anepisterno con al menos 1 seta en el margen posterior (Fig. 9G); antena delmacho con flagelómeros 2-5 articulados, no fusionados (flagelómeros 7-11fusionados, a veces incluso el flagelómero 6) ...................................................................................................................................... Ceratoculicoides WIRTH & RATANAWORABHAN

13(9) Palpo con 5 segmentos; largo del ala 0,7-0,8 mm; macho con parámeros biendesarrollados .............................................................................. Schizonyxhelea CLASTRIER

– Palpo con 2 segmentos (Fig. 9B); largo del ala < 0,6 mm; macho sin parámeros.............................................................................................................................................. 14

14(13) Tibia anterior con espolón apical; macrotriquias confinadas a los márgenesdistales del ala (en algunos, también unas pocas en el ápice de M1 y M2); relacióncostal > 0,40 (Fig. 11B); antena del macho con 6 flagelómeros separados (Fig. 8C);macho con tergitos abdominales 1-3 claros y 4-8 pardos .......................................................................................................................................... Baeohelea WIRTH & BLANTON

– Tibia anterior sin espolón apical; macrotriquias largas y abundantes sobre todoel ala; relación costal ratio < 0,36; antena del macho con 12 flagelómeros, perocon 2-7, 8-9, y 10-11 fusionados (Fig. 8B); macho con abdomen uniformementepardo ........................................................................ Baeodasymyia CLASTRIER & RACCURT

15(8) Ala con diseño característico de una pigmentación oscura moteada, más o menosdispersa sobre la membrana (Fig. 12C); tórax y al menos fémur y tibia posteriorescon diseño de pigmentación; edeago del macho con lazo posteromedianocaracterístico (Fig. 21A) .................................................................... Monohelea KIEFFER1

– Ala sin pigmentación o, si es pigmentada, pigmentación restricta al sector radial, ocon puntos, rayas o bandas discretas (Figs. 11E, H), o si el patrón de pigmentaciónes substancial (Fig. 11H) entonces tórax y todos los fémures y tibiasuniformemente pardos oscuros; otros taxa con tórax y patas pigmentados demanera variable; edeago, si está presente, sin rizo posteromediano……16

16(15) Costa extendiéndose hasta cerca del extremo del ala; costa del ala de la hembraextendiéndose más allá del ápice de la segunda celda radial (Figs. 12B, D, E)(extensión también presente en machos de Bahiahelea y Leptohelea) ...................... 17

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


– Costa no extendiéndose más allá de 0,85 del largo del ala (Figs. 11E, I); costa delala de la hembra finalizando en el ápice de la segunda celda radial o extendiéndosemás allá ligeramente ......................................................................................................... 25

17(16) Halterio en forma de saco, sin constricción por debajo del capitelo (Fig. 13J) ...........................................................................................Diaphanobezzia INGRAM & MACFIE

– Halterio con capitelo y pedicelo bien definidos ........................................................ 1818(17) Macho ................................................................................................................................ .19– Hembra ............................................................................................................................... 2119(18) Ala con costa no extendiéndose más allá del ápice de la segunda celda radial ......

...................................................................................... Parastilobezzia WIRTH & BLANTON

– Ala con costa extendiéndose apreciablemente más allá del ápice de la segundacelda radial ......................................................................................................................... 20

20(19) Con flagelómeros 10-13 más alargados que los flagelómeros precedentes;penacho antenal bien desarrollado; parámeros fusionados en una esferavoluminosa (Fig. 19F) ............................................................................ Bahiahelea WIRTH

– Con flagelómeros 2-13 más o menos iguales en longitud; penacho antenal corto;parámeros alargados, delgados ...................................... Leptohelea WIRTH & BLANTON

21(18) Palpo con tres segmentos, último segmento corto y pequeño (Fig. 9D); uñaspequeñas, iguales, sin diente basal interno (Fig. 15D) ................................................................................................................................................ Leptohelea WIRTH & BLANTON

– Palpo con 4-5 segmentos, último segmento alargado (Fig. 9E); uñas variables ................................................................................................................................................... 22

22(21) Clípeo no fusionado a la cabeza (como en la Fig. 6B); ala con la primera celdaradial pequeña, segunda alargada (Fig. 12E); con una uña (con diente basalgrueso) en cada pata (Fig. 15E) ............................ Parastilobezzia WIRTH & BLANTON

– Clípeo fusionado lateralmente a la cabeza (Figs. 6F, G); ala con 1 celda radial (Fig.12B); con 2 uñas en cada pata, a pesar de que las uñas de la pata posterior puedenser desiguales ..................................................................................................................... 23

23(22) Ojos moderadamente separados, sin pubescencia (Fig. 6F); uñas sin diente basalinterno ................................................................................................ Parabezzia MALLOCH

– Ojos anchamente separados, con o sin pubescencia (Fig. 6G); uñas con dientebasal interno ...................................................................................................................... 24

24(23) Palpo con 4 segmentos; ojos pubescentes; espermatecas sin cuellosdiferenciados ............................................................... Fittkauhelea WIRTH & BLANTON

– Palpo con 5 segmentos; ojos sin pubescencia; espermatecas con cuellosdiferenciados delgados ...................................................................... Bahiahelea WIRTH

25(16) Macho ................................................................................................................................. 26– Hembra ............................................................................................................................... 3926(25) Pata posterior con una uña larga y diente basal pequeño (Fig. 14H) .......................

................................................................................................................... Allohelea KIEFFER

– Pata posterior con dos uñas iguales o subiguales ...................................................... 2727(26) Katepisterno con 1 o más setas laterales (pueden ser observados en ejemplares

preservados en alcohol tomándolos verticalmente y observándolos de perfil a lolargo del plano de la pleura torácica) (Fig. 9G) ....... Brachypogon KIEFFER (en parte)


– Katepisterno sin setas .................................................................................................. 2828(27) Ala con 1 celda radial (Fig. 11E) .................................................................................. 29– Ala con 2 celdas radiales ................................................................................................. 3329(28) Coxa anterior con setas espinosas largas, o setas delgadas afinadas distalmente

(Fig. 9H) ............................................................................................. Parabezzia MALLOCH

– Coxa anterior con sólo setas cortas, simples (a veces perdidas en ejemplares enalcohol) ............................................................................................................................... 30

30(29) Ala con manchas pigmentadas, al menos inmediatamente basal a, o sobre r-m ysobre el ápice de la celda radial (Fig. 11E), algunos con pigmentación adicional............................................................................................................Alluaudomyia KIEFFER

– Ala sin manchas pigmentadas ........................................................................................ 3131(30) Palpo con 5 segmentos; tarsómero 1 de la pata posterior con al menos 1 1/2

hileras de setas empalizadas; edeago compuesto por 2 escleritos laterales (porlo que el edeago parece estar fisurado en la línea media) (Fig. 20E) o conedeago ausente pero con una barra transversa ventral al edeago (Fig. 20D)..............................................................................................Stilobezzia KIEFFER (en parte)

– Palpo con 4 segmentos (como en Fig. 9E); tarsómero 1 de la pata posterior consólo 1 hilera de setas empalizadas; edeago continuo en la línea media (como enFig. 21C) ............................................................................................................................. 32

32(31) Ojos sin pubescencia; genitalia con parámeros fusionados, muy cortos ........................................................................... undescribed genus near Parabezzia MALLOCH

– Ojos pubescentes; genitalia sin parámeros ........ Fittkauhelea WIRTH & BLANTON

33(28) Ala con áreas pigmentadas, incluyendo áreas poco notorias, discontínuas en M2and CuA2 (Fig. 12F); edeago continuo en la línea media (Fig. 20F) ............................................................................................................... Downeshelea WIRTH & GROGAN

– Ala con o sin diseño de pigmentación, pero si está presente, entonces sin áreaspoco notorias, discontínuas en M2 and CuA2 (porción posterior del ala puede sermás oscura) (Figs. 11J, 12A) y con edeago dividido en la línea media (Figs. 20B, C).............................................................................................................................................. 34

34(33) Antena con los 5 flagelómeros apicales más alargados que los 8 flagelómerosbasales (antena tipo-hembra), sin penacho ................. Cacaohelea WIRTH & GROGAN

– Antena con los 3 flagelómeros apicales más alargados que los 10 flagelómerosbasales (antena tipo-macho), con o sin penacho ....................................................... 35

35(34). Celdas radiales separadas por una nervadura longitudinal corta (R2); ala sinmacrotriquias; esternito 9 alargado (Fig. 20C) ........... Isthmohelea INGRAM & MACFIE

– Las 2 celdas radiales cercanas una de la otra, separadas por una nervaduratransversal corta (R2) (Figs. 11J, 12A); ala con o sin macrotriquias; esternito 9corto o moderadamente alargado (Figs. 20D, E) ...................................................... 36

36(35) Flagelómeros 3-10 fusionados (Fig. 8D) ............... Borkenthelea SPINELLI & GROGAN

– Antena con flagelómeros separados, no fusionados ................................................ 3737(36) Largo combinado de los tarsómeros 2-5 de la pata posterior aproximadamente

1,4-1,7 veces mayor que el tarsómero 1 ..................... Austrohelea WIRTH & GROGAN

– Largo combinado de los tarsómeros 2-5 de la pata posterior aproximadamente delmismo tamaño que el tarsómero 1 ............................................................................... 38


38(37) Tarsómero 1 de la pata posterior con al menos 1 1/2 hileras de setas empalizadas;ala con o sin macrotriquias; edeago compuesto por 2 escleritos laterales (porlo que el edeago parece estar fisurado en la línea media) (Fig. 20E) o conedeago ausente pero con una barra transversa ventral al edeago (Fig. 20D);flagelómeros antenales sin sensilas celocónicas ..................Stilobezzia KIEFFER

– Tarsómero 1 de la pata posterior con sólo 1 hilera de setas empalizadas (como enFig. 14A); ala sin macrotríquias; edeago no dividido, más o menos triangular;algunos flagelómeros antenales con sensilas celocónicas ............................................................................................................................Macrurohelea INGRAM & MACFIE

39(25) Katepisterno con 1 o más setas laterales (pueden ser observados en ejemplarespreservados en alcohol tomándolos verticalmente y observándolos de perfil a lolargo del plano de la pleura torácica) (Fig. 9G) ......... Brachypogon KIEFFER (en parte)

– Katepisterno sin setas .................................................................................................. 4040(39) Ala con 1 celda radial (Figs. 11C, E) ............................................................................ 41– Ala con 2 celdas radiales (la primera puede ser pequeña) (Figs. 10I, 11H, I) ........ 4341(40) Palpo con 4 segmentos (como en Fig. 9E); uñas de la pata posterior iguales ........

...................................................... género no descripto cercano a Parabezzia MALLOCH

– Palpo con 5 segmentos; uñas de la pata posterior desiguales o representadas porun talón (Figs. 14F, 15A-C) ............................................................................................ 42

42(41) Ala con manchas pigmentadas inmediatamente basal a, o sobre r-m y sobre elápice de la celda radial (Fig. 11E); tarsómero 1 de la pata posterior con sólo 1hilera de setas empalizadas ...........................................................Alluaudomyia KIEFFER

– Ala sin manchas pigmentadas; tarsómero 1 de la pata posterior con al menos 1 1/2hilera de setas empalizadas .............................................Stilobezzia KIEFFER (en parte)

43(40) Décimo segmento de la hembra alargado y doblado anteroventralmente (Fig. 22F)...........................................................................................Macrurohelea INGRAM & MACFIE

– Décimo segmento de la hembra no alargado, dirigido posteriormente ............... 4444(43) Alas con áreas pigmentadas, con 2-3 bandas oscuras (Fig. 11H), o al menos con

áreas pigmentadas en r-m e inmediatamente posterior al ápice de la segunda celdaradial .................................................................................................................................... 45

– Ala sin áreas pigmentadas discretas ............................................................................. 4745(43) Patas anterior y media, cada una con una uña (cada una con un diente basal más

pequeño) ............................................................................Stilobezzia KIEFFER (en parte)– Patas anterior y media, cada una con 2 uñas iguales ................................................ 4646(45) Uña de la pata posterior con diente basal (Fig. 14H) .................... Allohelea KIEFFER

– Uña de la pata posterior sin diente basal (Fig. 14I) ..... Downeshelea WIRTH & GROGAN

47(44) Patas anterior y media, cada una con 1 uña (algunos con diente basal máspequeño) ............................................................................................................................ 48

– Patas anterior y media, cada una con uñas pares, iguales, o desiguales (como enFig. 14F) ............................................................................................................................. 49

48(47) Primera celda radial pequeña, R3 engrosada (Fig. 10I); segunda celda radial más omenos ancha; nervadura mediana reducida a un trazo poco perceptible en la base;una uña corta, robusta, puntiaguda en cada pata (Fig. 15F) ..................................................................................................................................... Cacaohelea WIRTH & GROGAN


– Primera celda radial pequeña a bien desarrollada, R3 a lo sumo engrosada en elápice (Fig. 11I); segunda celda radial angosta; nervadura mediana claramentevisible, pero algunas veces la base de M2 está ausente; uña, de al menos la pataposterior, alargada (Figs. 15A-C) ..................................Stilobezzia KIEFFER (en parte)

49(47) Celdas radiales separadas por una nervadura longitudinal corta (R2); ala conmacrotriquias escasas, restrictas al ápice extremo; flagelómeros antenales sinsensilas celocónicas ...................................................... Isthmohelea INGRAM & MACFIE

– Celdas radiales separadas por una nervadura transversal corta (R2); ala con o sinmacrotriquias en el 1/4 apical; algunos flagelómeros antenales con sensilascelocónicas (sólo en el flagelómero 1 o sólo en flagelómeros 8 y 12) ........ 50

50(49) Ala sin macrotriquias; mandíbulas reducidas, vestigiales; ojos con o sinpubescencia; uñas pequeñas, iguales, en todas las patas .......................................................................................................................................... Notiohelea GROGAN & WIRTH

– Ala con macrotriquias en el 1/4 apical; mandíbulas dentadas; ojospubescentes; sólo patas anterior y media con uñas pequeñas, iguales, o uñasdesiguales en todas las patas (como en Fig. 14F) ...................................................... 51

51(50) Celdas radiales subiguales, pequeñas; R3 engrosada; uñas de la pata mediadesiguales y más o menos similares a las de la pata posterior; sólo flagelómero 1con sensilas celocónicas ....................................... Borkenthelea SPINELLI & GROGAN

– Segunda celda radial aproximadamente el doble de largo que primera celda radial;R3 no engrosada; pata media con uñas pequeñas, iguales, más cortas que la uñaúnica, alargada (con diente basal) de la pata posterior; sólo flagelómeros 8 y 12con sensilas celocónicas ............................................ Austrohelea WIRTH & GROGAN

52(7) Tarsómero 5 de la pata anterior hinchado (Figs. 15G, 17A, C); ................................................................................................................................. HETEROMYIINI..........53

– Tarsómero 5 de la pata anterior no hinchado ............................................................ 5653(52) M sésil, bifurcándose a nivel de r-m (Fig. 12G) ......... Physohelea GROGAN & WIRTH

– M anchamente sésil, bifurcándose anteriormente a r-m (Fig. 12H) ...................... 5454(53) Fémur anterior hinchado, con numerosas espinas gruesas ventrales (Fig. 17C) ....

........................................................................................................................ Heteromyia SAY

– Fémur anterior delgado, a lo sumo con unas pocas espinas delgadas .................. 5555(54) Tarsómero 4 de todas las patas cilíndrico o ligeramente cordiforme, sin espinas

fuertes; pata posterior de la hembra llamativamente alargada, con tarsómero 2 almemos 3/4 del largo del tarsómero 1 (Fig. 15H); uña posterior muy larga, única ocon diente basal pequeño, más larga que las uñas de las patas anterior y media (Fig.15H) .................................................................................................... Pellucidomyia MACFIE

– Tarsómero 4 de la pata anterior cordiforme, tarsómeros 4 de las patas media yposterior bífido y espinoso (Fig. 17B); pata posterior de la hembra no mucho máslarga que las otras patas, con tarsómero 2 menor que el doble del largo deltarsómero 1; ambas uñas posteriores aproximadamente del mismo largo queaquellas de las patas anterior y media .............................................. Clinohelea KIEFFER

56(52) Genitalia masculina grande, con gonocoxitos fusionados dorsalmente yventralmente (Fig. 21B); genitalia femenina con esternito 8 alargado, laminar (Fig.23A) ........................................................................................................ Echinohelea MACFIE


– Genitalia masculina pequeña o grande, si es grande entonces los gonocoxitos noestán fusionados; genitalia femenina con esternito 8 moderadamente desarrollado.............................................................................................................................................. 57

57(56) Macho (macho de Groganhelea, Sphaerohelea desconocidos) ...................................... 58– Hembra ............................................................................................................................... 7258(57) Gonostilo muy reducido en tamaño (Figs. 21D, 22B) o fusionado con el

gonocoxito (Fig. 22D) ..................................................................................................... 59– Gonostilo grande, claramente presente (Figs. 21F, 22A, C) (macho de Lanehelea

desconocido, pero probablemente se ubicaría en este sitio de la clave) .............. 6159(58) Ala con 2 celdas radiales claramente definidas (Fig. 13I); gonostilo bien definido,

con ápice puntiagudo (Fig. 21D) ..........................Clastrieromyia SPINELLI & GROGAN

– Ala con 1 celda radial o con, al menos, un indicio poco perceptible de divisiónentre primera y segunda celda radial (Fig. 13D); gonostilo pequeño y redondeadoapicalmente (Fig. 22B) o no claramente definido y fusionado con el gonocoxito(Fig. 22D) ........................................................................................................................... 60

60(59) Anepisterno con numerosas setas alargadas; gonocoxito corto y gonostilopresente pero pequeño (Fig. 22B) ..................................... Phaenobezzia HAESELBARTH

– Anepisterno sin setas; gonocoxito y gonostilo fusionados y alargados (Fig. 22D).................................................................................................................Nilobezzia KIEFFER

61(58) Genitalia con gonocoxito y tergito 9 alargados (Fig. 22C) ..................................... 62– Genitalia más robusta, con gonocoxito y tergito 9 no inusualmente alargados

(Figs. 21F, 22A) ................................................................................................................. 6462(61) Ala con 1 celda radial (como en Fig. 13D) ......Neobezzia WIRTH & RATANAWORABHAN

– Ala con 2 celdas radiales (Fig. 13G) ............................................................................ 6363(62) Fémur anterior con espinas guesas ventrales ................... Austrosphaeromias SPINELLI

– Fémur anterior sin espinas gruesas ventrales .............................. Mallochohelea WIRTH

(Nota. Los machos de algunas especies de Mallochohelea no tienen genitalia alargada)64(61) Tarsómere 5 de al menos la pata posterior con espinas robustas, romas (Fig. 17H);

ápice del tergito 9 con una hilera de setas gruesas, robustas (Fig. 21C) ......................................................................................................................... Johannsenomyia MALLOCH

– Tarsómero 5 sin espinas gruesas, o si están presentes, son más delgadas yfinamente puntiagudas (Fig. 16A); tergito 9 con setas dispersas, nunca dispuestasen una hilera de setas gruesas, robustas ...................................................................... 65

65(64) Con 1-2 setas robustas en el ápice de los cercos (Fig. 21E); parámerosfusionados, fusión restricta a la porción más anteromedia (Fig. 21E) ......................................................................................................... Amerohelea GROGAN & WIRTH

– Con algunas a muchas setas en los cercos; parámeros separados o condiferente grado de fusión ........................................................................................... 66

66(65) Fémur anterior con espinas robustas (Fig. 18A) ....................................................... 67– Fémur anterior sin espinas robustas ............................................................................ 6867(66) Ala con 1 celda radial (Fig. 13C) ........................................ Bezzia KIEFFER (en parte)– Ala con 2 celdas radiales (Fig. 13B) .............................. Palpomyia MEIGEN (en parte)68(66) Palpo con 4 segmentos ............................................................... Stenoxenus COQUILLETT

– Palpo con 5 segmentos ................................................................................................... 69


69(68) Tórax delgado y aplanado dorsoventralmente; paratergito con numerosas setaspequeñas; 1 celda radial (Fig. 12I) ..........................................Paryphoconus ENDERLEIN

– Tórax más robusto y no aplanado dorsoventralmente; paratergito sin setas; 1-2celdas radiales .................................................................................................................... 70

70(69) Ala con 1 celda radial (Fig. 13C) ........................................ Bezzia KIEFFER (en parte)– Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 7171(70) Fémur posterior hinchado (Fig. 18C); abdomen blanco, contrastando con el tórax

oscuro ........................................................................................................Pachyhelea WIRTH

– Fémur posterior no hinchado (Fig. 18A); abdomen claro a pardo oscuro, similar altórax oscuro ........................................................................ Palpomyia MEIGEN (en parte)

72(57) Ala con M2 fuertemente curvada cerca de la base (Fig. 13A); palpo con 4segmentos ...................................................................................... Stenoxenus COQUILLETT

– Ala con M2 no curvada cerca de la base, o sólo ligeramente (Figs. 13B-I); palpocon 5 segmentos ............................................................................................................... 73

73(72) Tarsómero 5 armado ventralmente con espinas robustas, negras, romas (Figs.16A, B, 17G-I) (estas espinas algo puntiagudas en Austrosphaeromias); abdomen sinapodemas tergales; esternito 8 con o sin (sólo Sphaerohelea) mechones depelos posterolaterales, con setas largas, curvadas, superpuestas en la líneamedia (mejor visión a 100X) (Fig. 23B); ...................... SPHAEROMIINI..........74

– Tarsómero 5 inerme, o si existen algunas espinas ventrales, son delgadas ypuntiagudas (Fig. 16E); abdomen con (Figs. 23C-E) o sin apodemas tergales (novisibles en algunos ejemplares jóvenes o pobremente esclerotizados); esternito 8sin mechones de pelos ................................................................................................ 81

74(73) Fémures sin espinas; uñas iguales en la pata anterior, desiguales en las patas mediay posterior (Figs. 17G-I) .......................................................... Johannsenomyia MALLOCH

– Fémures con o sin espinas; uñas iguales en todas las patas o sólo desiguales en lapata anterior ...................................................................................................................... 75

75(74) Uñas suavemente curvas distalmente, con diente basal delgado en el margeninterno ................................................................................................................................ 76

– Uñas rectas o aplanadas distalmente, con diente basal delgado o romo en elmargen externo (Figs. 16A, B) ...................................................................................... 78

76(75) Fémur anterior hinchado, con numerosas espinas ventrales gruesas ............................................................................................................................Austrosphaeromias SPINELLI

– Fémur anterior delgado, inerme ................................................................................... 7777(76) Costa extendiéndose hasta el extremo distal del ala; tarsómero 5 con 2-3 pares de

espinas ventrales .............................................. Sphaerohelea SPINELLI & FELIPPE-BAUER

– Costa no extendiéndose más allá de 0,85 del largo del ala; tarsómero 5 con 4-5pares de espinas ventrales ...............................................Lanehelea WIRTH & BLANTON

78(75) Ala con 1 celda radial; costa larga, extendiéndose casi hasta el extremo distal delala; relación costal mayor que 0,87; fémures sin espinas ventrales ....................... 79

– Ala con 1-2 celdas radiales; costa corta, no llegando a 0,84 del largo del ala;fémures con o sin espinas ventrales ............................................................................. 80


79(78) Diente basal externo de las uñas tarsales puntiagudo (Fig. 16A); tarso de la pataposterior no alargado (Fig. 16B); ala con lóbulo anal pobremente desarrollado .................................................................................Neobezzia WIRTH & RATANAWORABHAN

– Diente basal externo de las uñas tarsales romo (Fig. 16C); tarso de la pataposterior alargado (Fig. 16D); ala con lóbulo anal bien desarrollado (Fig. 13F) .......................................................................................... Groganhelea SPINELLI & DIPPOLITO

80(78) Ala con 2 celdas radiales (Fig. 13G); ala grisácea, con nervaduras anteriores pardasoscuras; fémur anterior sin espinas gruesas ventrales ............... Mallochohelea WIRTH

– Ala con 1 celda radial; ala blancuzca, nervaduras anteriores claras; fémur anteriorcon espinas gruesas ventrales ...........................................................Nilobezzia KIEFFER

81(73) Tórax delgado y aplanado dorsoventralmente; paratergito con numerosas setaspequeñas; 1 celda radial (Fig. 12I) ..........................................Paryphoconus ENDERLEIN

– Tórax más robusto y no aplanado dorsoventralmente; paratergito sin setas; ala con1-2 celdas radiales; ................................................................... PALPOMYIINI..........82

82(81) Abdomen con 1 par de apodemas tergales que surgen cerca de los márgeneslaterales del esternito 7; 1 espermateca presente, sin trazos ni evidencias de unasegunda reducida ............................................................. Amerohelea GROGAN & WIRTH

– Abdomen con 1 o más pares de apodemas tergales (Figs. 23C-E), si sólo estápresente 1 par, estos apodemas surgen más cerca de la línea media en el margenposterior del esternito 7; 1-2 espermatecas, pero si existe sólo 1, una segundareducida está presente ..................................................................................................... 83

83(82) Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 84– Ala con 1 celda radial (Fig. 13C) .................................................................................. 8684(83) Fémur anterior sin espinas robustas; fémur posterior muy hinchado (Fig. 18C);

abdomen blanco, contrastando con el tórax oscuro ........................Pachyhelea WIRTH

– Fémur anterior con o sin espinas robustas (Fig. 18A); fémur posterior no muyhinchado (Fig. 18A); abdomen claro a pardo oscuro, similar al tórax oscuro ........... 85

85(84) Flagelo corto, sólo 1,2-1,4 veces más largo que el ancho de la cabeza; membranaalar sin pigmentación; ala con lóbulo anal bien desarrollado (Fig. 13I); abdomencon un par de apodemas que surgen del margen anterior del tergito 7 y seextienden hasta el margen anterior del tergito 6 (Fig. 23C) ..............................................................................................................................Clastrieromyia SPINELLI & GROGAN

– Flagelo alargado, a menos 2,5 veces más largo que el ancho de la cabeza;membrana alar pigmentada; ala con lóbulo anal pobremente desarrollado (Fig.13B); abdomen con 2 o más pares de apodemas alargados que surgen del margenanterior de al menos los tergitos 6-7 y se extienden anteriormente dentro decavidad abdominal (Fig. 23E) ............................................................ Palpomyia MEIGEN

86(83) Costa moderadamente corta, relación costal 0,67-0,75 (Fig. 13C); fémur anteriorcon (Fig. 18B) o sin espinas robustas; tarsómero 5 con o sin setas ventralesrobustas .................................................................................... Bezzia KIEFFER (en parte)

– Costa más larga, relación costal ratio mayor que 0,80 (Fig. 13D); fémur anteriorsin espinas robustas; tarsómero 5 con setas ventrales delgadas con extremosafinados y doblados (Fig. 16E) ........................................... Phaenobezzia HAESELBARTH


CLAVE PARA LOS HUEVOS DE GÉNEROS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL

Solamente han sido descriptos los huevos de pocas especies de ceratopogónidos (Table 1).La clave siguiente es, por lo tanto, muy tentativa.

No se hallan disponibles claves para huevos de Diptera en las cuales los huevos deCeratopogonidae pudan ser distinguidos de aquellos pertenecientes a otras familias. Sinembargo, algunos Ceratopogonidae poseen huevos que son diferentes de todos los demás(Dasyhelea, Stenoxenus y Paryphoconus) o las oviposiciones tienen lugar en una matrizcaracterística (Alluaudomyia y la mayoría de los Sphaeromiini).

1 Huevo en forma de C (Fig. 24A) ...................................................................... Dasyhelea– Huevo, a lo sumo, con curvatura ligera (Figs. 24B-F) ............................................... 22(1) Huevo con collar característico subapical y constricción terminal a manera de

pezón en el extremo anterior (Fig. 24F) .............................. Stenoxenus y Paryphoconus– Huevo sin ornamentación en ninguno de los extremos, pequeñas espículas

pueden estar presentes (Figs. 24A-E) ............................................................................ 33(2) Huevo oval (Fig. 24B) ..................Leptoconops (en parte), Forcipomyiinae (en parte)– Huevo alargado, delgado (Figs. 24C-E) ........................................................................ 44(3) Huevos ordenados en una banda alargada, dispuestos en ángulo con respecto al

eje longitudinal de la banda (Fig. 24E) .................................................................................. mayoría de Sphaeromiini (conocidos para Johannsenomyia, Mallochohelea, Macropeza)

– Huevos colocados separadamente o en grupos; si están colocados en banda, loshuevos se hallan lado a lado (Figs. 24C, D) ................................................................. 5

5(4) Huevos colocados en banda (Fig. 24D) ............... Alluaudomyia (al menos en parte)– Huevos colocados separadamente o en grupos aislados (aunque no fijados uno

otro longitudinalmente) (Fig. 24C) ......................................... otros Ceratopogonidae

CLAVE PARA LAS LARVAS DE GÉNEROS DE CERATOPOGONIDAE DE LA REGIÓN NEOTROPICAL

Los inmaduros de Ceratopogonidae se hallan entre los más pobremente conocidos entretodas las familias de nematóceros (Table 2). Se brinda aquí una clave tentativa paraalgunos géneros comunes. El primer autor de esta contribución está preparando una clave,a nivel genérico, para larvas y pupas de géneros de todo el Mundo. Es importante destacarque sólo 11 de los 51 géneros conocidos de Ceratopogonidae Neotropicales han sidodescriptos como larvas sobre la base de material de esta región (Table 2), mientra que 11otros géneros se conocen como larvas a partir de material de otras regiones.

1 Pseudópodo presente en el protórax (Figs. 25B, C); cápsula cefálica hipognata aprognata (Figs. 25B, C); con setas fuertes en cápsula cefálica y cuerpo (Figs. 25B,C) (Forcipomyiinae) ........................................................................................................... 2

– Sin pseudópodo en el tórax (Figs. 25A, D-F); cápsula cefálica algo hipognata oprognata (Figs. 25A, D-F); sin setas fuertes en cápsula cefálica y cuerpoanteriormente al segmento anal (Figs. 25A, D-F) ....................................................... 3


2(1) Cuerpo algo aplanado dorsoventralmente, con tubérculos laterales (Fig. 25B) .................................................................................................................................. Atrichopogon

– Cuerpo cilíndrico en sección transversal, sin tubérculos laterales (Fig. 25C) ........................................................................................................................................ Forcipomyia

3(1) Cápsula cefálica dirigida anteroventralmente; finalización del abdomen con seriede ganchos, a veces dispuestos como dos pseudópodos (a menudo retraídos ydificil de observar en ejemplares preservados en alcohol), nunca con setasalargadas (Fig. 25D) ............................................................................................. Dasyhelea

– Cápsula cefálica prognata; finalización del abdomen simple, sin ganchos, con o sinsetas alargadas (Figs. 25A, E-F) ...................................................................................... 4

4(3) Cápsula cefálica con pigmentación pobremente desarrollada, con apodemas biendesarrollados dirigidos posteriormente y que se extienden en el protórax (Figs.26A, B); abdomen con segmentos divididos secundariamente (Fig. 25A) ............................................................................................................................................. Leptoconops

– Cápsula cefálica bien desarrollada, sin apodemas posteriores al margen de cápsulacefálica (Figs. 26C-F); abdomen con segmentos normales, no divididossecundariamente (Figs. 25E, F) (Ceratopogoninae) .................................................... 5

5(4) Cápsula cefálica con todas las setas simples (Figs. 26C, D) ............... Culicoides– Cápsula cefálica con una o más setas s, u y la posterior o plumosa (Figs.

26E, F, seta u no visible, generalmente lateral sobre la cápsula cefálica) .............. 66(5) Margen posteroventral de la cápsula cefálica con proceso que se proyecta

posteriormente (Fig. 26F) ........................................................ Sphaeromiini (en parte)– Margen posteroventral de la cápsula cefálica como una línea recta o algo curvada,

sin proceso que se proyecta posteriormente (Fig. 26E) ............................................................................................... Sphaeromiini (en parte), Palpomyiini, Ceratopogonini


8SYNOPSIS OF THE CERATOPOGONIDAE OF

THE NEOTROPICAL REGIONReferences providing available keys to the species of the various genera are also given inthe catalog in Table 4.

Allohelea KIEFFER. Only a single species occurs in the Neotropical Region, A. neotropicaWIRTH from Central America and Jamaica, but six species are known from the Nearc-tic (WIRTH, 1991a). Males are the only members in the family with massive, enlargedhind legs and a single hind claw, as in the predaceous female.

Alluaudomyia KIEFFER. Nineteen species occur in the Neotropical Region. Of these species,seventeen were reviewed by SPINELLI & WIRTH (1984c). These small, brightly pat-terned predaceous midges have larvae that inhabit ponds, lakes and streams wherethey are unique in swimming on the surface film.

Amerohelea GROGAN & WIRTH. Eleven species have been recorded from the NeotropicalRegion. Of these species, ten were reviewed by GROGAN & WIRTH (1981). Femalesdiffer from other members of the tribe Palpomyiini in having only a single spermath-eca and a single pair of internal abdominal tergal apodemes that arise near the lateralmargins of segment 7.

Atrichopogon KIEFFER. One hundred species are known from the Neotropical Region with18 recently described from Costa Rica (BORKENT & PICADO, 2004) and five from Pat-agonia (SPINELLI et al., 2006). Six new species from Peruvian Amazonia are being pres-ently described (SPINELLI & MARINO, in prep.). The genus is clearly very diverse in theNeotropical Region and we suspect that hundreds of species remain to be described.Adults are among the most commonly encountered diurnal biting midges and can befound on a variety of flowers (mostly small, white or green) or resting on vegetation.The adult males of at least some species form swarms. Larvae are aquatic to terrestri-al and have distinctive elongate dorsal setae (a feature shared with many Forcipomyia).The feeding habits of the adults are poorly known but some suck blood from blisterbeetles (WIRTH 1956a, b; BORKENT & ROCHA-FILHO, 2006), while others feed on deadinsects, are pollinovorous or are non-feeding (autogenous). The larvae and pupaehave important taxonomic features. EWEN & SAUNDERS (1958) reared nine speciesfrom several localities in the Neotropics.

Austrohelea WIRTH & GROGAN. Only a single species occurs in the Neotropical Region, A.shannoni WIRTH & BLANTON from the temperate Nothofagus forests of Argentina andChile. The genus exhibits a transantarctic distribution, with six other species known


from Australia and New Zealand (WIRTH & GROGAN, 1988). At least three unnamedspecies have been collected from Patagonia.

Austrosphaeromias SPINELLI. The genus is known only from two species from southern Ar-gentina and Chile (SPINELLI, 1997a). It is the only genus of the tribe Sphaeromiini inthe Neotropical Region inhabiting temperate Nothofagus forests.

Baeodasymyia CLASTRIER & RACCURT. Five species of these minute, highly reduced bitingmidges are known in the Neotropical Region (BORKENT & CRAIG, 1999). Adults walkwith a distinctive rapid, stuttering gait (best seen with a hand lens). The immatures arerestricted to the headwaters of springs and small seeps.

Baeohelea WIRTH & BLANTON. Only a single species (B. nana WIRTH & BLANTON) of thisminute, highly reduced predaceous genus is known in the Neotropical Region, distrib-uted from Costa Rica to Ecuador. Baeohelea and its sister group Baeodasymyia are theonly two genera in the family to have their palpi reduced to two segments.

Bezzia KIEFFER. Forty-six species occur in the Neotropical Region. Most species are shiningdark black, but many have striking banded leg patterns. This is the most commongenus in the tribe Palpomyiini in the region and the second most common predaceousgenus after Stilobezzia. The larva of at least one species in Costa Rica lives in brome-liads. Others likely occur in larger water bodies such as rivers and lakes.

Borkenthelea SPINELLI & GROGAN. The genus is known from four species from the temperateNothofagus forests of southern Argentina and Chile (SPINELLI & GROGAN, 2001). Malesare unique in the tribe Ceratopogonini in having flagellomeres 3-10 fused.

Brachypogon KIEFFER. Twenty-five species of these small, black midges are known from theNeotropical Region. Four new species from the Peruvian Amazonia are being described(SPINELLI & MARINO, in prep.). One undescribed species is common in some rocky coastalareas in Costa Rica and an additional 11 new species are known from Costa Rica. Thissuggests that numerous further species remain to be discovered in this region.

Cacaohelea WIRTH & GROGAN. Only a single described species of these tiny midges is known,and the original specimens came from a cacao plantation in Costa Rica (WIRTH &GROGAN, 1988). Further material, including that of a second undescribed species, hasbeen collected in Costa Rica in areas devoid of cacao.

Ceratoculicoides WIRTH & RATANAWORABHAN. Within the region, only one species is knownfrom Mexico, with a second, unnamed species from Costa Rica known from twofemale adults (HUERTA & BORKENT, 2005). All remaining species are restricted to theHolarctic Region (WIRTH & GROGAN, 1988).

Clastrieromyia SPINELLI & GROGAN. The genus is known from four Neotropical species (SPINEL-LI & GROGAN, 1985, 1986). Females differ from other members of the tribe Palpomyi-ini in having only a single pair of internal abdominal tergal apodemes arising from theanterior margin of tergite 7 and extending to about the anterior margin of tergite 6.

Clinohelea KIEFFER. Fourteen species of this predaceous genus occur in the NeotropicalRegion (SPINELLI & DURET, 1993). The elegant adults have long, slender legs and largedark maculations on their wings.

Culicoides LATREILLE. Two hundred and sixty-six species of this vertebrate-biting genus havebeen recorded in the region. Of these, only 16 are serious pests of humans or domes-tic animals. The most serious coastal species biting humans in Central America and


northern South America is C. furens, attacking on beaches and in homes near man-grove swamps. They may occur in such huge numbers that they provide the importantecological service of restricting humans from living in some areas. Various pest spe-cies are discussed above in the section on Bionomics. The adult males of at least somespecies form swarms. Larvae of species of Culicoides are semiaquatic (wet mud, moss-es, rotting vegetation) to aquatic and feed on microorganisms and small invertebrates.There are clearly a significant number of species remaining to be discovered, especial-ly from higher elevations in the Neotropical Region.

Dasyhelea KIEFFER. Fifty-eight species of this common, mainly diurnal genus are recordedfrom the Neotropics, but many more remain undescribed. At least eleven new speciesare known from Patagonia (DÍAZ & SPINELLI, in prep.). Adults have reduced, non-biting mouthparts but are common on many flowers (particularly those that are smalland white) where they feed on nectar. Adults may be common in some coastal habi-tats, particularly where there are seeps, pools and, for a few species, in flat rocky areas.The adult males of at least some species form swarms. The larvae are aquatic grazerson algae and detritus and are generally present in natural small aquatic habitats (e.g.tree holes, rock pools) or artificial ones (e.g. cemetery flower-holding containers).This very diverse and common genus is in urgent need of revision.

Diaphanobezzia INGRAM & MACFIE. The genus is known from four species from ArgentineanPatagonia. The peculiar halter, without a constriction below the knob, is unique withinthe family.

Downeshelea WIRTH & GROGAN. Eighteen species occur in the Neotropical Region (LANE &WIRTH, 1964). They have distinctive large blotches on their wings and may be reason-ably common in some Malaise trap samples.

Echinohelea MACFIE. Eleven species of this genus are known in the Neotropical Region (WIRTH,1994b). The distinctive adults are patterned and bright yellow to golden brown in color,have numerous spines on their legs and both sexes have long proboscis and elongateantennal flagella. The species are strikingly similar to one another and therefore some-what difficult to identify. Specimens are occasionally common in Malaise trap samples.

Fittkauhelea WIRTH & BLANTON. The genus is known from a single species, F. amazonicaWIRTH & BLANTON from Brazilian Amazonia. Females are similar to females of Para-bezzia except for the presence of tarsal claws with an basal inner tooth.

Forcipomyia MEIGEN. Two hundred and ten species of this common genus occur in theNeotropical Region. Adults of some species are ectoparasites of large insects (asdescribed in the section on Bionomics above) but those of the subgenus Lasioheleafeed on vertebrate blood. Although adult females are common and many have bitingmouthparts, the hosts for most of the species remain unknown. The adult males of atleast some species form swarms. Most larvae have elongate dorsal setae (also presentin Atrichopogon), are aquatic to terrestrial, and are found in a diverse array of habitatssuch as in mosses, under the bark of rotting trees, inside decaying fruits, and in aquat-ic epiphytes. Many species remain to be described.

Groganhelea SPINELLI & DIPPOLITO. This genus of the tribe Sphaeromiini is known from asingle species, G. rondoniensis SPINELLI & DIPPOLITO, from Brazilian Amazonia and Uru-guay. Males are unknown.


Heteromyia SAY. Eleven species occur in the Neotropical Region (DURET & LANE, 1955; WIRTH

& GROGAN, 1977). The distinctive, large adult females have greatly swollen fore femorawith dense ventral spines and extremely elongated hind legs, each with a single, verylong claw which they project upward when at rest. Occasionally adults may be abundant.

Isthmohelea INGRAM & MACFIE. The genus is known from a single species, I. disjuncta INGRAM

& MACFIE, from its type-locality, in the Valdivian rain forests of southern Chile. Themale has a strikingly elongate abdominal segment 9.

Johannsenomyia MALLOCH. Only two species are known from the Neotropical Region. Thelarvae inhabit rivers and streams.

Lanehelea WIRTH & BLANTON. This genus of the tribe Sphaeromiini is known from twospecies from Colombia. Males are unknown.

Leptoconops SKUSE. Twelve species of this vertebrate blood sucking genus occur in the Neo-tropical Region. Females, especially those of L. bequaerti, L. chilensis FORATTINI and L.petrocchiae SHANNON & DEL PONTE, are often serious daytime biting pests of humans.Adults of both sexes have the unusual habit of burrowing into sand to rest and forfemales to lay their eggs. The larvae are pink and primarily inhabit wet, protectedsands on beaches of oceans and lakes.

Leptohelea WIRTH & BLANTON. The genus is known from a single female specimen, L. micr-onyx WIRTH & BLANTON (WIRTH & BLANTON, 1970a), from Colombia and one malefrom Costa Rica. The maxillary palpus of adults is 3-segmented. Males are includedhere, in a key, for the first time.

Macrurohelea INGRAM & MACFIE. Twelve species occur in the Neotropical Region, particular-ly in Patagonia. SPINELLI & GROGAN (1990) provided a key to males and females ofeleven of these species. The genus exhibits a transantarctic distribution, being knownfrom 3 other species from Australia (GROGAN & WIRTH, 1985). Females are unique inthe family in having the tenth segment elongated and bent anteroventrally.

Mallochohelea WIRTH. Six species are known from the Neotropical Region. None are common.Monohelea KIEFFER. Eighteen species occur in the Neotropical Region, the majority of them

included in a key to males and females by LANE & WIRTH (1964). Adults have wingswith characteristic black and gray maculations, brightly banded legs and females havesingle large hind claws.

Nannohelea GROGAN & WIRTH. Only one named species of this minute predaceous genus,N. clastrieri GROGAN & WIRTH, occurs in the Neotropical Region, from Colombia (GRO-GAN & WIRTH, 1980a, 1990). Another undescribed species is known from Costa Rica,based on one male. Otherwise, three species are recorded from the Old World.

Neobezzia WIRTH & RATANAWORABHAN. Eight of these rare species occur in the NeotropicalRegion. WIRTH & RATANAWORABHAN (1972a) provided a key to females of seven ofthese species.

Nilobezzia KIEFFER. Three species occur in the Neotropical Region. LANE (1961b) provideda key to males and females. Adults may be locally common, especially around lowlandpermanent ponds.

Notiohelea GROGAN & WIRTH. The genus is known from two species from the temperateNothofagus forests of southern Argentina and Chile (GROGAN & WIRTH, 1979b; SPINELLI

& GROGAN, 1990). Males are unknown.


Pachyhelea WIRTH. Two species occur in the Neotropical Region, the wide-ranging P. pachym-era (WILLISTON) and P. albidiventris (KIEFFER) from Colombia (GROGAN & WIRTH, 1980c).It is similar to Palpomyia.

Palpomyia MEIGEN. Forty six species occur in the Neotropical. The adult females feed onsmall Diptera or mayflies (GROGAN & WIRTH, 1979c).

Parabezzia MALLOCH. Twenty-four species occur in the Neotropical region (SPINELLI & GRO-GAN, 1987).

Paradasyhelea MACFIE. Three species are restricted to southern Chile and Argentina (SPINEL-LI & GROGAN, 2003). Seven further species are known from the Australasian region,and a single species occurs in the Nearctic region.

Parastilobezzia WIRTH & BLANTON. This rare genus is known from a single species, P. leeiWIRTH & BLANTON, from Costa Rica and Colombia (WIRTH & GROGAN, 1988).

Paryphoconus ENDERLEIN. Forty species occur in the Neotropical Region. SPINELLI & WIRTH

(1984b) provided a key to females of thirty eight of these species. The adults aremedium-sized to large (female wing lengths 1.5-7 mm) and are generally rare. The fewabundant collections from the Neotropical Region are from near rivers.

Pellucidomyia MACFIE. Four species occur in the Neotropical Region (WIRTH & RATANA-WORABHAN, 1971b). Adults are rare and females resemble those of Heteromyia in hav-ing greatly elongated hind legs and a single elongate claw, but they lack the swollen,spinose forefemur of that genus.

Phaenobezzia HAESELBARTH. Three species are broadly distributed in the Neotropical Region(SPINELLI & WIRTH, 1986a).

Physohelea GROGAN & WIRTH. The genus is known only from two species from southernArgentina and Chile (GROGAN & WIRTH, 1979a). It is the only genus of the tribeHeteromyiini in the Neotropical Region inhabiting temperate Nothofagus forests.

Rhynchohelea WIRTH & BLANTON. A single species of this monotypic genus, R. monilicornisWIRTH & BLANTON, is known from Florida and Costa Rica (WIRTH & GROGAN, 1988).Within the Neotropical Region, this minute relative of Brachypogon is known from onlyone female specimen from Costa Rica (BORKENT, 1992).

Schizonyxhelea CLASTRIER. Two species of these small, reduced relatives of Stilobezzia areknown from the Neotropical Region (BORKENT, 2000b). The larvae of one specieswere collected from a moderately-sized spring in Costa Rica.

Sphaerohelea SPINELLI & FELIPPE-BAUER. A single species of this monotypic genus, S. biestroiSPINELLI & FELIPPE-BAUER, is known from northeastern Argentina (SPINELLI & FE-LIPPE-BAUER, 1990b). Males are unknown.

Stenoxenus COQUILLETT. Sixteen species occur in the Neotropical Region). WIRTH & RATANA-WORABHAN (1972b) provided a key to females of fourteen of these species. This genus isclosely related to Paryphoconus, the only two genera in the tribe Stenoxenini, and the eggsof both genera have a terminal nipple-like constriction with a fringed subapical collar.Female Stenoxenus have a strikingly different M2 wing vein (Fig. 13A) compared to theirmales and other members of the family. Adults may be common near rivers.

Stilobezzia KIEFFER. Sixty-four species of these common, often brightly colored and pat-terned predaceous midges are recorded from the Neotropical Region (LANE & FORAT-TINI, 1961). However, the genus is clearly diverse in the region. For example, we have


examined 87 species from Costa Rica alone and there are undoubtedly many more yetto be discovered in that small country. At least twelve new species in the subgenusAcanthohelea, from Nothofagus forest in Patagonia, are being described (CARZOLA &SPINELLI, in prep.).

Undescribed genus near Parabezzia MALLOCH. A single species is known as male and femaleadults from Costa Rica. It will be fully described in a future publication.


9REFERENCES

AITKEN, T.H.G., WIRTH, W.W, WILLIAMS, R.W, DAVIES, J.B & E.S. TIKASINGH (1975): A review of thebloodsucking midges of Trinidad and Tobago, West Indies (Diptera: Ceratopogonidae). - J.Entomol. (B) 44: 101-144.

ATCHLEY, W.R. & W.W. WIRTH (1979): A review of the Culicoides haematopotus group in North America(Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 52: 524-545.

BARBOSA, F.A.S. (1943): Descrição de “Culicoides recifensis” n. sp. e do macho de “Culicoides reticulatus”LUTZ (Diptera, Chironomidae). - Rev. Bras. Biol. 3: 261-264.

BARBOSA, F.A.S. (1947): Culicoides (Diptera: Heleidae) da região neotropical. - Ann. Soc. Biol.Pernambuco 7: 3-30 (Nov.).

BARBOSA, F.A.S. (1951): A change of specific name in the genus Culicoides (Diptera, Heleidae). - Proc.Entomol. Soc. Washington 53: 163.

BARBOSA, F.A.S. (1952): Novos subsidios para o conhecimento dos Culicoides neotrópicos (Diptera:Heleidae). - Thesis, Univ. Recife, Recife: Imprensa Industrial: 21 pp.

BARBOSA, F.A.S. (1953): Novos subsidios para o conhecimento dos Culicoides neotrópicos (Diptera:Heleidae). - Pub. Avulsas Inst. Aggeu Magalhães 2: 11-41.

BARRETTO, M.P. (1944): Sobre o género Culicoides LATREILLE, 1809, com a descrição de trés novasespécies (Diptera, Ceratopogonidae). - Ann. Fac. Med. Univ. São Paulo 20: 89-105, pls. 1-4.

BECK, E.C. (1951): A new Culicoides from Florida (Diptera, Ceratopogonidae). - Fla. Entomol. 34:135-136.

BECK, E.C. (1956): A new species of Culicoides from Florida with additional distribution data for thegenus (Diptera: Heleidae). - Fla. Entomol. 39: 133-138.

BLANTON, F.S. & W.W. WIRTH (1979): The sand flies (Culicoides) of Florida (Diptera: Ceratopogonidae).-Arthropods of Florida and neighboring land areas 10: xv + 204 pp.

BORKENT, A. (1991): The Ceratopogonidae (Diptera) of the Galápagos Islands, Ecuador with adiscussion of their phylogenetic relationships and zoogeographic origins. - Entomol. Scand.22: 97-122.

BORKENT, A. (1992): A new key to some genera of Ceratopogonini in the Holarctic (Diptera:Ceratopogonidae). - Entomol. Scand. 22: 433-436.

BORKENT, A. (1996): Biting midges (Ceratopogonidae: Diptera) feeding on a leatherback turtle inCosta Rica. - Brenesia 43-44: 25-30 (1995).

BORKENT, A. (2000a): Biting midges (Ceratopogonidae: Diptera) from Lower Cretaceous Lebaneseamber with a discussion of the diversity and patterns found in other ambers. - In: GRIMALDI, D.(ed.): Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey:355-452. Backhuys Publ., Leiden: viii + 498 pp.


BORKENT, A. (2000b): The larva and pupa of Schizonyxhelea forattinii WIRTH and GROGAN (Diptera:Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus. - Proc.Entomol. Soc. Washington 102: 862-868.

BORKENT, A. (2004): 10. Ceratopogonidae. - In: MARQUARDT, W.C. (ed.): Biology of disease vectors:113-126. 2nd edition. Elsevier Academic Press, Amsterdam: xxiii + 785 pp.

BORKENT, A. & D.A. CRAIG (1999): A revision of the Neotropical genus Baeodasymyia CLASTRIER andRACCURT (Diptera: Ceratopogonidae) with a discussion of their phylogenetic relationships. -Am. Mus. Novitates 3274: 1-26.

BORKENT, A. & D.A. CRAIG (2001): Submerged Stilobezzia rabelloi LANE (Diptera: Ceratopogonidae)pupae obtain oxygen from the aquatic fern Salvinia minima BAKER. - Proc. Entomol. Soc.Washington 103: 655-665.

BORKENT, A. & D.A. CRAIG (2004): Austroconops WIRTH and LEE, a Lower Cretaceous genus of bitingmidges yet living in Western Australia: a new species, first description of the immatures anddiscussion of their biology and phylogeny (Diptera: Ceratopogonidae). - Am. Mus. Novitates3449: 1-67.

BORKENT, A. & L. FORSTER (1986): Review of the Dasyhelea fasciigera species group (Diptera:Ceratopogonidae) with a revision of the Nearctic species. - Can. J. Zool. 64: 1280-1287.

BORKENT, A. & A. PICADO (2004): Distinctive new species of Atrichopogon KIEFFER (Diptera:Ceratopogonidae) from Costa Rica. - Zootaxa 637: 1-37.

BORKENT, A. & L.C. ROCHA-FILHO (2006): First record of female adult Atrichopogon KIEFFER (Diptera:Ceratopogonidae) biting in the Neotropical Region. - Proc. Entomol. Soc. Washington 108:998-1001.

BORKENT, A. & G.R. SPINELLI (2000): Catalog of New World biting midges south of the UnitedStates (Diptera: Ceratopogonidae). - Contrib. Entomol. Internat. 4: 1-107.

BORKENT, A. & W.W. WIRTH (1997): World species of biting midges (Diptera: Ceratopogonidae). -Bull. Am. Mus. Nat. Hist. 233: 1-257.

BRÈTHES, J. (1912): Descripción de un nuevo género y especie nueva de Chironomidae (Dipt). - An.Mus. Nac. Buenos Aires 15: 451-453.

BRÈTHES, J. (1914): Descripción de seis Cécidomyidae (Dipt.) de Buenos Aires. - An. Mus. Nac.Buenos Aires 26: 151-156.

BRICKLE, D.S. & D.V. HAGAN (1999): The Culicoides (Diptera: Ceratopogonidae) of Belize, CentralAmerica. - Insecta Mundi 13: 39-44.

BROWN, B.V. (2005): Malaise trap catches and the crisis in Neotropical Dipterology. - Am. Entomologist51: 180-183.

BROWNE, J.E. (1980): New species of biting midges of the genus Culicoides from Colombia and the firstdescription of the male of C. florenciae (Diptera: Ceratopogonidae). - J. Med. Entomol. 17: 533-544.

BYSTRAK, P.G. & W.W. WIRTH (1978): The North American species of Forcipomyia, subgenusEuprojoannisia (Diptera: Ceratopogonidae). - U.S. Dept. Agric., Tech. Bull. 1591: 1-51.

CARTER, H.F. (1921): A revision of the genus Leptoconops SKUSE. - Bull. Entomol. Res. 12: 1-28 (June).CAVALIERI, F. (1961a): Notas sobre Ceratopogonidae (Diptera, Nematocera) I. Forcipomyia (Forcipomyia)

wygodzinskyi sp. n. para Tierra del Fuego. - Revta. Soc. Entomol. Argent. 23: 17-19.CAVALIERI, F. (1961b): Notas sobre Ceratopogonidae (Diptera, Nematocera) II. Sobre dos nuevas

especies de Forcipomyia para Tierra del Fuego, Forcipomyia (F.) delpontei n. sp. y F. (F.) piroskyi. -Acta Zool. Lilloana 18: 169-175 (1962).


CAVALIERI, F. (1962): Notas sobre Ceratopogonidae (Dipt. Nematocera) III. Sobre un nuevo dípterohematófago para Argentina, Lasiohelea saltensis n. sp. y notas sobre las especies neotropicales deLasiohelea. - Acta Zool. Lilloana 18: 359-365.

CAVALIERI, F. (1966): Notas sobre Ceratopogonidae (Diptera, Nematocera) V. Descripción de unanueva especie de Venezuela: Culicoides (Oecacta) birabeni sp. n. - Physis 26: 59-63.

CAVALIERI, F. & I.F. CHIOSSONE (1966): Datos sobre la familia de dípteros hematófagos Leptoconopidaeen la Argentina, con la descripción de Leptoconops (Leptoconops) casali sp. n. y la redescripción deL. (L.) petrocchiae SHANNON y DEL PONTE, 1927. - Physis 26: 43-57.

CAVALIERI, F. & I.F. CHIOSSONE (1972): Notas sobre el género Atrichopogon KIEFFER, 1906, de la Argentina(Diptera, Ceratopogonidae). I. Consideraciones generales y descripción de Atrichpogon delponteisp. nov. - Revta. Soc. Entomol. Argent. 34: 119-129.

CAVALIERI, F. & I.F. CHIOSSONE (1973): Notas sobre el género Atrichopogon KIEFFER, 1906, de la Argentina(Diptera, Ceratopogonidae). II. Descripción de Atrichpogon casali sp. nov. - Physis 32: 151-159.

CAZORLA, C.G. & P.I. MARINO (2004): The pupa of Stilobezzia punctulata LANE from Peruvian Amazonia(Diptera: Ceratopogonidae). - Amazoniana 18: 75-80.

CAZORLA, C.G., DÍAZ, F. & M.M. RONDEROS (2006): Redescription of pupa and adult of Stilobezziafiebrigi KIEFFER 1917 (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 132: 111-119.

CAZORLA, C.G., SPINELLI, G.R. & F. DÍAZ (2005): Two new species of the subgenus Stilobezzia (Stilobezzia)KIEFFER from Peruvian Amazonia (Diptera: Ceratopogonidae). - Amazoniana 18: 289-297.

CHAN, K.L. & E.J. LEROUX (1965): Description of Forcipomyia (Neoforcipomyia) sp. n. and redescriptionof Forcipomyia (Neoforcipomyia) eques (JOHANNSEN) (Diptera: Ceratopogonidae), with an accountof the digestive and reproductive systems. - Phytoprotection 46: 74-104.

CHAN, K L. & E.J. LEROUX (1967): Ecological studies on three pond midges (Diptera:Ceratopogonidae) in Quebec. - Ann. Soc. Entomol. Québec 12: 14-68.

CHAN, K L. & E.J. LEROUX (1971): A new subgenus and species of midge (Diptera: Ceratopogonidae)from Singapore. - Can. Entomol. 103: 271-276.

CHAUDHURI, P.K., DAS GUPTA, S.K. & D.K. CHAUDHURI (1972): Biting midges of the genus AlluaudomyiaKIEFFER from India (Diptera: Ceratopogonidae). - Oriental Insects 6: 83-110.

CHAVEZ, R., COLAND, E. & I. PHILLIPS (1992): Fiebre de oropouche en Iquitos. Reporte preliminar de5 casos. - Rev. Farmacol. Terap. 2: 12-14.

CLASTRIER, J. (1968) : Deux Ceratopogonides nouveaux de la Guyane Française. - Arch. Inst. PasteurGuyane Franc. Territ.- L’Inini 21 : 85-92.

CLASTRIER, J. (1971) : Deux nouveaux Culicoides (Diptera, Ceratopogonidae) de la Guyane Française.- Ann. Parasitol. 46 : 285-294.

CLASTRIER, J. (1972) : Forcipomyia (F.) squamithorax n. sp. de la Guyane Française. - Bull. Soc. Entomol.France 77 : 170-176.

CLASTRIER, J. (1973) : Le genre Leptoconops, sous-genre Holoconops dans le midi de la France (Dipt.Ceratopogonidae). - Ann. Soc. Entomol. France 9: 895-920.

CLASTRIER, J. (1976): Alluaudomyia prima n. sp. De la Guyane Française (Diptera, Ceratopogonidae). -Nouv. Rev. Entomol. 6: 205-207.

CLASTRIER, J. (1979): Un nouvel Atrichopogon de la Guyane Française (Diptera: Ceratopogonidae). -Rev. Franc. Entomol. 1: 30-32.

CLASTRIER, J. (1984): Schizonyxhelea guyana n. g., n. sp. de la Guyane Française (Diptera, Ceratopogonidae).- Rev. Franc. Entomol. 6 : 1-4.


CLASTRIER, J. (1987) : Atrichopogon ornatipennis n. sp. de la Guyane Française. - Entomologiste 43 : 271-274.CLASTRIER, J. (1991) : Description de cinq nouveaux Stilobezzia apparentés à S. Insolita DAS GUPTA &

WIRTH, originaires des régions afrotropicale, paléarctique et néotropicale (Dipt.Ceratopogonidae). - Bull. Soc. Entomol. France 95: 297-310 (1990).

CLASTRIER, J. (1992): Description de Palpomyia guyana n. sp. de la Guyane Française (Diptera,Ceratopogonidae). - Rev. Franc. Entomol. 14: 117-121.

CLASTRIER, J. & J.-C. DELÉCOLLE (1994): Description de Forcipomyia (Phytohelea) musae n. sp. de laGuyane française (Diptera, Ceratopogonidae). - Rev. Franc. Entomol. 16: 51-56.

CLASTRIER, J. & J.-C. DELÉCOLLE (1997): Description de Forcipomyia (Trichohelea) roubaudi n. sp.,ectoparasite d’un hétéroptère réduvide capturé dans la canopée de la foret guyanaise (Diptera,Ceratopogonidae). - Bull. Soc. Entomol. France 102: 379-383.

CLASTRIER, J. & J. LEGRAND (1990): Forcipomyia (Pterobosca) incubans (MACFIE) et F. (Trichohelea) machetin. sp. Parasites des ailes de libellules en Guyane Française (Diptera, Ceratopogonidae; Odonata).- Rev. Franc. Entomol. 12: 167-170.

CLASTRIER, J. & C. RACCURT (1979): Quatre nouveaux Parabezzia de la Republique d’Haiti (Diptera,Ceratopogonidae). - Nouv. Rev. Entomol. 9: 165-175.

CLASTRIER, J. & C. RACCURT (1979b): Baeodasymyia modesta n. g., n. sp, de la République d’Haiti (Diptera,Ceratopogonidae). - Ann. Parasitol. 54: 99-104.

CLASTRIER, J. & W.W. WIRTH (1978): The Leptoconops kerteszi complex in North America (Diptera:Ceratopogonidae). - U.S. Dept. Agric. Tech. Bull. 1573: 1-58.

CLASTRIER, J. & W.W. WIRTH (1995) : Révision des Forcipomyia du sous-genre Microhelea de la région Néotropicale,parasites de phasmes (Diptera : Ceratopogonidae). - Ann. Soc. Entomol. France 31: 97-150.

COQUILLETT, D.W. (1899): A new dipterous family related to the Chironomidae. - Entomol. News 10:60-61.

COQUILLETT, D.W. (1901a): New Diptera in the U.S. National Museum. - Proc. U.S. Nat. Mus. 23:593-618.

COQUILLETT, D.W. (1901b): Papers from the Hopkins Stanford Galapagos Expedition, 1898-1899. II.Entomological Results (2): Dipteria. - Proc. Wash. Acad. Sci. 3: 371-379.

COQUILLETT, D.W. (1902): New Diptera from North America. - Proc. U.S. Nat. Mus. 25: 83-126.COQUILLETT, D.W. (1904a): New Diptera from Central America. - Proc. Entomol. Soc. Washington 6:

90-98.COQUILLETT, D.W. (1904b): A new Ceratopogon from Brazil. - J. New York Entomol. Soc. 12: 35-36.COQUILLETT, D.W. (1905): New nematocerous Diptera from North America. - J. New York Entomol.

Soc. 13: 56-69.COQUILLETT, D.W. (1910): The type-species of the North American genera of Diptera. - Proc. U.S.

Nat. Mus. 37: 499-647.CORDERO, E.H. (1929): Contribución al estudio de los Dipteros del Uruguay, I. Lophomyidium uruguayense

n. Gen., n. Sp. Nueva Ceratopogonina hematofaga. - An. Mus. Hist. Nat. Montevideo 3: 93-108.COSTA LIMA, A. DA (1928): Ceratopogonineos ectoparasitos de phasmideos. - Mem. Inst. Oswaldo

Cruz, Suppl. 3: 84-85, 2 pls.COSTA LIMA, A. DA (1937ª): Chave das espécies de Culicoides da região neotropical (Diptera:

Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 32: 411-422.COSTA LIMA, A. DA (1937b): Primeira espécie americana do gênero Pterobosca (Diptera: Ceratopogonidae).

- Mem. Inst. Oswaldo Cruz 32: 615-616, pls. 1-2.


CUNHA, R. G. (1990): Anticorpos neutralizantes em soros de ruminantes domésticos do Brasil frenteãos diferentes serotipos do virus da lingua azul. - Rev. Bras. Med. Vet. 12: 3-7.

DEBENHAM, M.L. (1987): The biting midge genus Forcipomyia (Diptera: Ceratopogonidae) in theAustralasian Region (exclusive of New Zealand). I. Introduction, key to subgenera, and theThyridomyia and Trichohelea groups of subgenera. - Invertebr. Taxon. 1 : 35-119.

DELÉCOLLE, J.-C. & J.-P. RIEB (1994) : Contribution à l’étude des Cératopogonidés de la Guadeloupe.Description de trois espèces nouvelles, appartenant aux genres Dasyhelea et Atrichopgon (Diptera,Nematocera). - Bull. Soc. Entomol. France 99 : 267-279.

DELECOLLE, J.-C., RACCURT, C.P. & C. REBHOLTZ (1986) : Culicoides haitiensis n. sp. Et Culicoides borinqueniFOX et HOFFMAN en Haiti. I. Étude morphologue (Diptera, Ceratopogonidae). - Nouv. Rev.Entomol. 3 : 107-115.

DE MEILLON, B. & W.W. WIRTH (1979) : A taxonomic review of the subgenus Phytohelea of Forcipomyia(Diptera : Ceratopogonidae). - Proc. Entomol. Soc. Washington 81: 178-206.

DE MEILLON, B. & W.W. WIRTH (1980): A new subgenus of Forcipomyia, with descriptions of eightnew species (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 82: 9-24.

DE MEILLON, B. & W.W. WIRTH (1991): The genera and subgenera (excluding Culicoides) of theAfrotropical biting midges (Diptera: Ceratopogonidae). - Ann. Natal Mus. 32: 27-147.

DÍAZ, F., RONDEROS, M.M. & G.R. SPINELLI (2005): The immatures of the Neotropical species Culicoidesvenezuelensis ORTÍZ and MIRSA (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 131:375-385.

DIPPOLITO, A., SPINELLI, G.R. & W.W. WIRTH (1995): A report on a collection of Ceratopogonidae(Diptera) from Rondonia (Brazil) and Iquitos (Peru) 1. Tribes Palpomyiini and Stenoxenini. -Insecta Mundi 9: 53-60.

DOVE, W.E., HALL, D.G. & J.B. HULL (1932): The salt marsh sand fly problem. - Ann. Entomol. Soc.Am. 25: 505-527.

DOW, M.I. & E.C. TURNER (1976): A revision of the Nearctic species of the genus Bezzia (Diptera:Ceratopogonidae). - Occassional Papers Dept. Entomol., Virginia Polytech. Inst. State Univ.,Res. Div. Bull. 103: 1-162.

DOW, M.I. & W.W. WIRTH (1972): Studies on the genus Forcipomyia, 2. The Nearctic species of thesubgenera Thyridomyia and Synthyridomyia (Diptera: Ceratopogonidae). - Ann. Entomol. Soc.Am. 65: 177-201.

DOWNES, J.A (1978): Feeding and mating in the insectivorous Ceratopogoninae (Diptera). - Mem.Entomol. Soc. Can. 104: 1-62.

DOWNES, J.A. & W.W. WIRTH (1981): 28. Ceratopogonidae. - In: MCALPINE, J.F., PETERSON, B.V., SHEWELL,G.E., TESKEY, H.J., VOCKEROTH, J.R. & D.M. WOOD (eds.): Manual of Nearctic Diptera: 393-421.Vol. 1. Agriculture Canada Monograph 27. Biosystematic Research Institute, Ottawa: 674 pp.

DURET, J. P. & J. LANE (1955): Novas Heteromyia da Argentina (Diptera. Ceratopogonidae). - Dusenia6: 35-40.

EDWARDS, F.W. (1922): On some Malayan and other species of Culicoides, with a note on the genusLasiohelea. - Bull. Entomol. Res. 13: 161-167.

EDWARDS, F.W. (1933): Die Ausbeute der deutschen Chaco-Expedition 1925/26. Diptera (Fortsetzung)XXXIX-XLI. Cecidomyiidae, Chironomidae, Ceratopogonidae. - Konowia 12: 86-88.

ENDERLEIN, G. (1912): Paryphoconus, eine neue Chironomidengattung aus Brasilien. - Stett. Entomol.Zeit. 73: 57-60.


ENDERLEIN, G. (1936): Ordnung: Zweiflügler, Diptera. Abt. 16. - In: BROHMER, P., EHRMANN, P. & G.ULMER (eds.): Die Tierwelt Mitteleuropas 6: Insekten III Teil. Quello & Meier, Leipzig.

EWEN, A.B. & L.G. SAUNDERS (1958): Contributions toward a revision of the genus Atrichopogonbased on characters of all stages (Diptera, Heleidae). - Can. J. Zool. 36: 671-724.

FELIPPE-BAUER, M.L. (1987): A new Neotropical Culicoides from Minas Gerais (Diptera:Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 82: 147-148.

FELIPPE-BAUER, M.L. & R. LOURENCO-DE-OLIVEIRA (1987): A new neotropical biting midge of theCulicoides debilipalpis group (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 82: 149-150.

FELIPPE-BAUER, M.L. & A.R. QUINTELAS (1993a): Two new Brazilian predaceous midges of the genusDowneshelea WIRTH and GROGAN (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 88: 33-38.

FELIPPE-BAUER, M.L. & A.R. QUINTELAS (1993b): Downeshelea bicornis, a new neotropical predaceousmidge from Brazil (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 88: 185-187.

FELIPPE-BAUER, M.L. & A.R. QUINTELAS (1994): Culicoides lobatoi, a new Brazilian biting midge of thelimai group (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 89: 25-27.

FELIPPE-BAUER, M.L. & G.R. SPINELLI (1991): New Brazilian predaceous midges of the genus MonoheleaKIEFFER, and redescription of M. guaimiesi (Diptera: Ceratopogonidae). - Mem. Inst. OswaldoCruz 86: 201-207.

FELIPPE-BAUER, M.L. & G.R. SPINELLI (1994): Two new Neotropical species of Monohelea KIEFFER

and Downeshelea WIRTH & GROGAN (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz89: 161-165.

FELIPPE-BAUER, M.L. & G.R. SPINELLI (1998): A new Neotropical species of Monohelea KIEFFER fromUruguay (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 93: 63-64.

FELIPPE-BAUER, M.L. & W.W. WIRTH (1987): A new Culicoides of the stigmalis group (Diptera:Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 82: 415-419.

FELIPPE-BAUER, M.L. & W.W. WIRTH (1988): Culicoides tavaresi, a new species from the state of Rio deJaneiro, Brazil (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 83: 261-263.

FELIPPE-BAUER, M.L., HUERTA, H. & S. IBAÑEZ BERNAL (2000a): A new species of predaceous midgeof the genus Monohelea KIEFFER from Mexico (Diptera: Ceratopogonidae). - Mem. Inst. OswaldoCruz 95: 815-818.

FELIPPE-BAUER, M.L., QUINTELAS, A.R. & G.R. SPINELLI (1995): A new Neotropical predaceous midge,Downeshelea deani, and redescription of Downeshelea guianae (WIRTH) (Diptera: Ceratopogonidae).- Mem. Inst. Oswaldo Cruz 90: 395-399.

FELIPPE-BAUER, M.L., VERAS, R. S., CASTELLON, E. G. & N.A. MOREIRA (2000b): A new Culicoides from theAmazonian Region, Brazil (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 95: 35-37.

FELIPPE-BAUER, M.L., CACERES, A.G., SILVA, C.S., VALDERRAMA-BAZAN, W. & A. GONZALES-PEREZ

(2003): Two new Culicoides of the paraensis species group (Diptera: Ceratopogonidae) from theAmazonian region of Peru. Mem. - Inst. Oswaldo Cruz 98: 1051-1058.

FELIPPE-BAUER, M.L., CACERES, A.G., SILVA, C.S., VALDERRAMA-BAZAN, W. & A. GONZALES-PEREZ

(2005): A new Culicoides (Diptera: Ceratopogonidae) of the subgenus Diphaomyia from Peru. -Mem. Inst. Oswaldo Cruz 100: 51-53.

FIEBRIG-GERTZ, C. (1928): Un Diptère ectoparasite sur un Phasmide: Ceratopogon ixodoides n. sp. -Ann. Parasit. Hum. Comp. 6: 284-290.

FLOCH, H. & E. ABONNENC (1942a): Cératopogonides hématophages de la Guyane Française. - Pub.Inst. Pasteur Guyane Territ. L’Inini 37: 1-10.


FLOCH, H. & E. ABONNENC (1942b): Cératopogonidés hématophages de la Guyane Française - II. -Pub. Inst. Pasteur Guyane Territ. L’Inini 49: 1-5.

FLOCH, H. & E. ABONNENC (1949): Sur les Cératopogonidés du Venezuela. Description de deux espècesnouvelles: Culicoides lichyi et Lasiohelea danaisi. - Pub. Inst. Pasteur Guyane Territ. L’Inini 196: 1-5.

FLOCH, H. & E. ABONNENC (1950a): Ceratopogonides nouveaux du Venezuela Culicoides lichyi etLasiohelea danaisi n. sp. - Bol. Entomol. Venez. 8: 69-75 (1949).

FLOCH, H. & E. ABONNENC (1950b): Les Culicoides de la Guadeloupe. Description de Culicoidesguadeloupensis n. sp. - Pub. Inst. Pasteur Guyane Territ. L’Inini 203: 1-5.

FORATTINI, O.P. (1954a): Nova espécie de Culicoides (Diptera, Ceratopogonidae) do Brasil Central. -Arq. Fac. Hig. Saude Pub. Univ. São Paulo 8: 189-192.

FORATTINI, O.P. (1954b): Nova espécie de Culicoides (Diptera, Ceratopogonidae) do estado de Goiaz,Brasil. - Folia Clin. Biol. 21: 315-320.

FORATTINI, O.P. (1956a): Nova espécie de Culicoides (Diptera, Ceratopogonidae) de São Vicente, estadode São Paulo, Brasil. - Arq. Fac. Hig. Saude Pub. Univ. São Paulo 10: 81-84.

FORATTINI, O.P. (1956b): A new Culicoides from Panama (Diptera, Ceratopogonidae). - Proc. Entomol.Soc. Washington 58: 35-36.

FORATTINI, O.P. (1957): Culicoides da região neotropical (Diptera, Ceratopogonidae). - Arq. Fac. Hig.Saude Pub. Univ. São Paulo 11: 159-526.

FORATTINI, O.P. (1958): Nova espécie de Leptoconops s. str. da regiao neotropical (Diptera,Ceratopogonidae). - Rev. Bras. Entomol. 8: 37-41.

FORATTINI, O.P. & J. LANE (1955): Sobre uma Forcipomyia que suga Phasmidae (Diptera, Nematocera,Ceratopogonidae). - Rev. Bras. Entomol. 4: 1-6.

FORATTINI, O.P. & E.X. RABELLO (1956): As formas imaturas de Culicoides guyanensis FLOCH e ABONNENC, 1942e de algumas espécies de Stilobezzia (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 6: 43-50.

FORATTINI, O.P. & E.X. RABELLO (1957): Dados sobre Dasyhelea KIEFFER, 1911 do Brasil (Diptera,Ceratopogonidae). - Rev. Bras. Entomol. 7: 241-250.

FORATTINI, O.P., RABELLO, E.X. & D. PATOLLI (1956): Nota sobre a larva e pupa de Culicoides insignisLUTZ, 1913 (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 4: 195-198.

FORATTINI, O.P., RABELLO, E.X. & R.D. PATOLLI (1960): Sobre as formas imaturas de Culicoides reticulatus LUTZ

e de Stilobezzia panamensis LANE e FORATTINI (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 9:1-4.FOX, I. (1942): The respiratory trumpet and anal segment of the pupae of some species of Culicoides

(Diptera: Ceratopogonidae). - Puerto Rico J. Pub. Health Trop. Med. 17: 412-425 [Spanishversion, pp. 426-434].

FOX, I. (1946): A review of the species of biting midges or Culicoides from the Caribbean Region(Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 39: 248-258.

FOX, I. (1947): Two new Central American biting midges or Culicoides (Diptera: Ceratopogonidae). -Kuba 3: 90-91 (June).

FOX, I. (1948): Hoffmania, a new subgenus in Culicoides (Diptera: Ceratopogonidae). - Proc. Biol. Soc.Washington 61: 21-28.

FOX, I. (1952): Six new Neotropical species of Culicoides (Diptera: Ceratopogonidae). - Ann. Entomol.Soc. Am. 45: 364-368.

FOX, I. (1955): A catalogue of the bloodsucking midges of the Americas (Culicoides, Leptoconops, andLasiohelea) with keys to the subgenera and Nearctic species, a geographic index, and bibliography.- J. Agric. Univ. Puerto Rico 39: 214-285.


FOX, I. & W.A. HOFFMAN (1944): New Neotropical biting sand flies of the genus Culicoides (Diptera:Ceratopogonidae). - Puerto Rico J. Pub. Health Trop. Med. 20: 108-111.

FRANA, M.F., GASPARICH, G.E. & W.L. GROGAN (2001): First isolation of a Spiroplasma (Mollicutes:Spiroplasmataceae) from biting midges (Diptera: Ceratopogonidae). - Entomol. News 112: 64-70.

GARRETT, C.B.D. (1925): Seventy new Diptera. - Cranbrook, British Columbia: 16 pp.GERRY, B.I. (1933): Four new species of Chironomidae from the Greater Antilles. - Psyche 40: 94-97.GLUKHOVA, V.M. (1979): Larvae of the biting midges of the subfamilies Palpomyiinae and

Ceratopogoninae of the Fauna of the USSR (Diptera, Ceratopogonidae = Heleidae) [in Russian].- AN SSSR, Opredeliteli po faune SSSR, 121, Nauk, SSSR, Leningrad: 230 pp.

GLUKHOVA, V.M. (1989): Blood-sucking midges of the genera Culicoides and Forcipomyia(Ceratopogonidae) [in Russian]. - Fauna of the USSR 139, Vol. 3(5a): 1-408.

GOELDI, E. (1905): Os mosquitos no Pará. Reunião de quatro trabalhos sobre os mosquitos indígenas,principalmente as espécies que molestam o homem. - Mem Mus. Goeldi (Museu Paraense)Hist. Nat. Ethnogr. 4: 1-154, 5 pls.

GOETGHEBUER, M. (1920) : Ceratopogoninae de Belgique. - Mem. Mus. R. Hist. Nat. Belgique 8: 1-116.GORCH, C., VAGNOZZI, A., DUFFY, S., MIQUET, J., PACHECO, J., BOLONDI, A., DRAGHI, G., CETRA, B.,

SONI, C., RONDEROS, M., RUSSO, S., RAMIREZ, V. & I. LAGER (2002): Lengua azul: aislamiento ycaracterización del virus e identificación de vectores en el noreste argentino. - Rev. Argent.Microbiol. 34: 150-156.

GRIFFITHS, G.C.D. (1980): Preface to flies of the Nearctic region (Pp. v-vii in Vol. I, part 1). -Schweizerbart, Stuttgart.

GROGAN, W.L. & J.A. WIENERS (2006): A new species of the biting midge genus Dasyhelea KIEFFER

(Diptera: Ceratopogonidae) from the Bahamas. - Proc. Entomol. Soc. Washington 108: 467-473.GROGAN, W.L. & W.W. WIRTH (1975): A revision of the Nearctic species of Clinohelea KIEFFER (Diptera:

Ceratopogonidae). - Great Basin Natur. 35: 275-287.GROGAN, W.L. & W.W. WIRTH (1977): A revision of the Nearctic species of Parabezzia MALLOCH

(Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 50: 49-83.GROGAN, W.L. & W.W. WIRTH (1979a): A new neotropical genus of predaceous midges, with a key to the

genera of Heteromyiini (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 81: 51-59.GROGAN, W.L. & W.W. WIRTH (1979b): Notiohelea, a new genus of biting midges of the tribe

Ceratopogonini from Chile (Diptera: Ceratopogonidae). - Pan-Pac. Entomol. 54: 283-286 (1978).GROGAN, W.L. & W.W. WIRTH (1979c): The North American predaceous midges of the genus Palpomyia

MEIGEN (Diptera: Ceratopogonidae). - Mem. Entomol. Soc. Washington 8: 1-125.GROGAN, W.L. & W.W. WIRTH (1980a): Nannohelea, a new genus of biting midges of the tribe

Ceratopogonini, related to Baeohelea WIRTH and BLANTON (Diptera: Ceratopogonidae). - J. KansasEntomol. Soc. 53: 373-385.

GROGAN, W.L. & W.W. WIRTH (1980b): Two new species of Macrurohelea from Chile with a key to theNeotropical species (Diptera: Ceratopogonidae). - Pan-Pac. Entomol. 56: 137-143.

GROGAN, W.L. & W.W. WIRTH (1980c): The taxonomic status of the predaceous midge Pachyhelea pachymera(WILLISTON) (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 82: 74-80.

GROGAN, W.L. & W.W. WIRTH (1981): A new American genus of predaceous midges related to Palpomyiaand Bezzia (Diptera: Ceratopogonidae). - Proc. Biol. Soc. Washington 94: 1279-1305.

GROGAN, W.L. & W.W. WIRTH (1985): Two new australian species of Macrurohelea with a descriptionof the male of M. commoni (Diptera: Ceratopogonidae). - Internat. J. Entomol. 27: 128-135.


GROGAN, W.L. & W.W. WIRTH (1990): A new species of the minute predaceous midge genus Nannoheleafrom Sri Lanka (Diptera, Ceratopogonidae). - Proc. Entomol. Soc. Washington 92: 347-350.

GUTSEVICH, A.V. & V.M. GLUKHOVA (1970): Methods for collecting and studying blood-sucking midges[in Russian]. - Methods in Parasitological investigations 3: 1-103, Akademia Nauk SSSR, Leningrad.

HAESELBARTH, E. (1965): Phaenobezzia, a new genus of biting midges (Diptera: Ceratopogonidae),with a review of the African species. - Zeitsch. Ang. Zool. 52: 297-324.

HOCH, A.L., ROBERTS, D.R. & F.P. PINHEIRO (1990): Host-seeking behavior and seasonal abundance ofCulicoides paraensis (Diptera: Ceratopogonidae) in Brazil. - J. Am. Mosquito Control Assoc. 6: 110-114.

HOFFMAN, W.A. (1925): A review of the species of Culicoides of North and Central America and theWest Indies. - Am. J. Hyg. 5: 274-301.

HOFFMAN, W.A. (1926): Two new species of American Leptoconops (Diptera, Chironomidae). - Bull.Entomol. Res. 17: 133-136.

HOFFMAN, W.A. (1939): Culicoides filariferus, new species. - Puerto Rico J. Public Health Trop. Med. 15: 172-176.HOGUE, C.L. & W.W. WIRTH (1968): A new Central American sand fly breeding in crab holes (Diptera,

Ceratopogonidae). - Los Angeles County Mus., Contrib. Sci. 152: 1-7.HRIBAR, L.J. (1993): Mouthpart morphology and feeding behavior of biting midge larvae (Diptera:

Ceratopogonidae). - In: SCHAEFER, C.W. & R.A.B. LESCHEN (eds.): Functional morphology ofinsect feeding. 43-58. Thomas SAY Publications in Entomology, Entomological Society ofAmerica, Lanham, Maryland.

HRIBAR, L.J. & G.R. MULLEN (1991): Comparative morphology of the mouthparts and associatedfeeding structures of biting midge larvae (Diptera: Ceratopogonidae). - Contrib. Am. Entomol.Inst. 26: 3-71.

HUERTA, H. (1996): Los Ceratopogónidos colectados por Alfonso DAMPF en Chiapas, Mexico (Diptera:Ceratopogonidae). - Thesis. Universidad Nacional Autónoma de Mexico: xi + 199 pp.

HUERTA, H. (2001): A new species of the genus Atrichopogon KIEFFER (Diptera: Ceratopogonidae)from Mexico. - Proc. Entomol. Soc. Washington 103: 373-375.

HUERTA, H. & A. BORKENT (2005): A new species and first record of Ceratoculicoides WIRTH andRATANAWORABHAN from the Neotropical region and new species and records of BrachypogonKIEFFER from Mexico (Diptera: Ceratopogonidae).- Folia Entomol. Mex. 44 (Supl. 1): 111-119.

HUERTA, H. & S. IBAÑEZ-BERNAL (1996): Especie nueva de Forcipomyia (Lasiohelea) de Chiapas, México(Diptera: Ceratopogonidae). - An. Inst. Biol., Univ. Nac. Auton. México, Ser. Zool. 67: 349-355.

HUNT, G. J. & E.T. SCHMIDTMANN (2004): 54. Care, maintenance, and experimental infection of bitingmidges. - In: MARQUARDT, W.C. (ed.): Biology of disease vectors: 741-745. 2nd edition. ElsevierAcademic Press, Amsterdam: xxiii + 785 pp.

ICHES, L. (1906): Sobre cinco dípteros nuevos del Chaco Austral. - Bol. Min. Agric. Gan. Argent. 6:262-273.

INGRAM, A. & J.W.S. MACFIE (1921): West African Ceratopogonidae. - Ann. Trop. Med. Parasitol. 15:313-376.

INGRAM, A. & J.W.S. MACFIE (1924): Notes on some African Ceratopogonidae-species of the genusLasiohelea. - Ann. Trop. Med. Parasitol. 18: 377-392, pl. 22.

INGRAM, A. & J.W.S. MACFIE (1931): Ceratopogonidae. - In: Diptera of Patagonia and South Chile.Part II. Fasc. 4: 155-232.

JAMNBACK, H. (1965): The Culicoides of New York State (Diptera: Ceratopogonidae). - New York St.Mus. Sci. Serv. Bull. 399: viii + 154 pp.


JAMNBACK, H., WALL, W.J. & D.L. COLLINS (1958): Control of Culicoides melleus (COQ.) (Diptera: Heleidae)in small plots, with brief descriptions of the larvae and pupae of two coastal Culicoides. -Mosquito News 18: 64-70.

JOHANNSEN, O.A. (1905): Aquatic nematocerous Diptera II. - In: NEEDHAM, J.G., MORTON, K.J. &O.A. JOHANNSEN (eds.): May flies and midges of New York. Third report on aquatic insects:76-315. - New York St. Mus. Bull. 86: 7-352 (= New York St. Educ. Dept. Bull. 343). Alsopublished in New York St. Mus. Ann. Report (1904) 58 (Vol. 5 = app. 7): 76-315.

JOHANNSEN, O.A. (1907): Some new species of Kansas Chironomidae. - In: TUCKER, E.S.: Someresults of desultory collecting of insects in Kansas and Colorado:109-112. Bull. Univ. Kansas,Sci. Bull. 4: 51-112.

JOHANNSEN, O.A. (1927): The genus Stenoxenus (Chironomidae, Diptera). - Entomol. News 38:70-72.

JOHANNSEN, O.A. (1938): New species of Nemocera from Puerto Rico. - J. Agri. Univ. Puerto Rico22: 219-225.

JOHANNSEN, O.A. (1943): Two new species of American Ceratopogonidae (Diptera). - Ann. Entomol.Soc. Am. 36: 761-762.

JOHNSON, C.W. (1908): The Diptera of the Bahamas, with notes and description of one new species.- Psyche 15: 69-80.

JOHNSON, C.W. (1913): The Diptera fauna of Bermuda. - Ann. Entomol. Soc. Am. 6: 443-452.JONES, R.H. (1961): Descriptions of pupae of thirteen North American species of Culicoides (Diptera:

Ceratopogonidae). - Ann. Entomol. Soc. Am. 54: 729-746.KARSCH, F. (1886): Ein neues märkisches Dipteron (Ceratopogon crudelis n. sp.). - Berl. Entomol. Zeitsch.

30: xvii-xviii.KHALAF, K.T. (1952): The Culicoides of the Wichita Refuge, Oklahoma. Taxonomy and seasonal

incidence (Diptera, Heleidae). - Ann. Entomol. Soc. Am. 45: 348-358 (June).KIEFFER, J.J. (1899): Description d’un nouveau genre et tableau des genres européens de la famille

des Chironomides (Dipt.). - Bull. Soc. Entomol. France 1899: 66-70.KIEFFER, J.J. (1906a): Diptera. Fam. Chironomidae. - In: WYTSMAN, P. (ed.): Genera Insectorum: Fasc.

42: 78 pp., 4 pls. Bruxelles.KIEFFER, J.J. (1906b): Description d’un genre nouveau et de quelques espèces nouvelles de diptères

de l’Amérique de Sud. - Ann. Soc. Scient. Bruxelles 30: 349-358.KIEFFER, J.J. (1909): Considérations sur le genre Stenoxenus COQ. avec description d’une espèce nouvelle.

- Ann. Hist.-Nat. Mus. Nat. Hungarici 7: 46-49.KIEFFER, J.J. (1911a): Description de nouveaux chironomides de l’Indian Museum de Calcutta. - Rec.

Indian Mus. 6: 113-177, pls. 6-7.KIEFFER, J.J. (1911b): Les chironomides de Tendipedidae de l’Himalaya et d’Assam. - Rec. Indian

Mus. 6: 319-349.KIEFFER, J.J. (1911c): Nouvelles descriptions de chironomides obtenus d’éclosion. - Bull. Soc. Hist.

Nat. Metz 27: 1-60 (Jan.).KIEFFER, J.J. (1913): Chironomidae et Cecidomyidae. - In: ALLUAUD, C.A. & R. JEANNEL (eds.): Voyage

de Ch. ALLUAUD et R. JEANNEL en Afrique orientale (1911-1912). Resultats scientifiques. Diptera(= Vol. 5): 1-43, Paris: 351 pp.

KIEFFER, J.J. (1917a): Chironomides d’Australie conservés au Musée National Hongrois de Budapest.- Ann. Hist.-Nat. Mus. Nat. Hungarici 15: 175-228.


KIEFFER, J.J. (1917b): Chironomides d’Amérique conservés au Musée National Hongrois de Budapest.- Ann. Hist.-Nat. Mus. Nat. Hungarici 15: 292-364.

KIEFFER, J.J. (1918): Chironomides d’Afrique et d’Asie conservés au Museum National Hongrois deBudapest. - Ann. Hist.-Nat. Mus. Nat. Hungarici 16: 31-136.

KIEFFER, J.J. (1919a): Chironomides d’Europe conservés au Musée National Hongrois de Budapest.- Ann. Hist.-Nat. Mus. Nat. Hungarici 17: 1-160. (Dec. 29).

KIEFFER, J.J. (1919b): Observations sur les chironomides (Dipt.) décrits par J.R. MALLOCH. - Bull. Soc.Entomol. France 1919: 191-194.

KIEFFER, J.J. (1921a): Chironomides de l’Afrique Équatoriale. - Ann. Soc. Entomol. France 90: 1-56.KIEFFER, J.J. (1921b): Description de quelques Chironomides exotiques. - Ann. Soc. Scientif. Bruxelles

40: 181-186.KIEFFER, J.J. (1921c): Sur quelques Diptères piqueurs de la tribu des Ceratopogoninae. - Arch. Inst.

Pasteur Afrique du Nord 1: 107-115 (March).KIEFFER, J.J. (1925): Chironomides de la République Argentine. - Ann. Soc. Scient. Bruxelles 44: 73-92.KNAB, F. (1914): Ceratopogoninae sucking the blood of caterpillars. - Proc. Entomol. Soc. Washington

16: 63-66.KNAB, F. (1915): New Ceratopogoninae from Peru. - Insec. Insc. Menstr. 3: 109-111.KNAUSENBERGER, W.I. (1987): Contributions to the autecology and ecosystematics of immature

Ceratopogonidae (Diptera), with emphasis on the tribes Heteromyiini and Sphaeromiini in theMiddle Atlantic United States. - Ph.D.-thesis, Virginia Polytechnic Institute and State University,Blacksburg, Virginia: xxix + 729 pp.

KRIVOSHEINA, N.P. (1968): A contribution to the biology and morphology of little studied bitingmidges of the genus Forcipomyia Meigen (Diptera, Ceratopogonidae) [in Russian, Englishsummary]. - Zool. Zhur. 47: 578-590.

LAMBERSON, C., PAPPAS, C.D. & L.G. PAPPAS (1992): Pupal taxonomy of the tree-hole Culicoides (Diptera:Ceratopogonidae) in eastern North America. - Ann. Entomol. Soc. Am. 85: 111-120.

LANE, J. (1944): As espécies neotropicais do gênero Clinohelea KIEFFER, 1917. (Diptera,Ceratopogonidae). - Rev. Entomol. 15: 249-261.

LANE, J. (1946a): New Neotropical Ceratopogonidae (Heleidae). (Diptera, Nemocera). - Rev. Entomol.17: 202-215.

LANE, J. (1946b): Duas espécies novas de Ceratopogonídeos Brasileiros (Diptera Ceratopogonidae(Heleidae)). - Livro de Homenagem a R.F. d’ALMEIDA 22: 219-225.

LANE, J. (1947a): Espécies Brasileiras de Stilobezzia (Dipt. Ceratopogonidae) e Zygoneura stonei nov.nom. (Dipt. Mycetophilidae). - Rev. Entomol. 18: 197-214.

LANE, J. (1947b): Novas espécies de Palpomyia do Brasil (Diptera, Ceratopogonidae). - Rev. Entomol.18: 438-447.

LANE, J. (1947c): A biologia e taxonomia de algumas espécies dos grupos Forcipomyia e Culicoides(Diptera, Ceratopogonidae) (Heleidae). - Arq. Fac. Hig. Saude Pub. Univ. São Paulo 1: 159-170.

LANE, J. (1948): Novas Ceratopogonídeos do Brasil (Diptera, Ceratopogonidae) (Heleidae). - Arq.Fac. Hig. Saude Pub. Univ. São Paulo 1: 225-239, Figs 1-8.

LANE, J. (1956a): On “Paryphoconous” and Stenoxenus (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 16:299-308.

LANE, J. (1956b): The genus “ Macfiehelea” LANE, 1946 (Diptera, Ceratopogonidae). - Rev. Bras. Biol.16: 435-438.


LANE, J. (1958): On Neotropical Bezzia. - Rev. Bras. Entomol. 8: 25-36.LANE, J. (1960): Additional data on “Palpomyia” (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 20:

381-389.LANE, J. (1961a): Further notes on neotropical Ceratopogonidae (Diptera). - Rev. Bras. Biol. 21: 37-44.LANE, J. (1961b): Insecta Amapaensia, Diptera: Ceratopogonidae and Anisopodidae. - Studia Entomol.

4: 449-452.LANE, J. (1961c): The genera Stenoxenus and Paryphoconus in the Neotropics (Diptera: Ceratopogonidae).

- Studia Entomol. 4: 452-458.LANE, J. & J.P. DURET (1954): Clinohelea da Argentina e do Brasil (Diptera, Ceratopogonidae). - Dusenia

5: 247-253.LANE, J. & O.P. FORATTINI (1956): Neotropical Stilobezzia KIEFFER, 1911 I. Nine new Panamanian species.

(Diptera, Nematocera, Ceratopogonidae). - Rev. Bras. Malariol. Doencas Trop. 8: 207-226.LANE, J. & O.P. FORATTINI (1958): Neotropical Stilobezzia II. Fourteen new species, chiefly from

Panama (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 8: 203-224.LANE, J. & O.P. FORATTINI (1961): Neotropical Stilobezzia, 1911 III. Key for the the adults of this genus

and description of one new species (Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 10: 83-94.LANE, J. & W.W. WIRTH (1961): A new Neotropical species of Dicrohelea (Diptera, Ceratopogonidae).

- Rev. Bras. Entomol. 10: 81-82.LANE, J. & W.W. WIRTH (1964): The biting midge genus Monohelea KIEFFER in the Neotropical region

(Diptera, Ceratopogonidae). - Studia Entomol. 7: 209-236.LANE, J., FORATTINI, O.P. & E.X. RABELLO (1955): Biologia e espécies novas de Palpomyia e Stilobezzia

(Diptera, Nematocera, Ceratopogonidae). - Dusenia 6: 81-88.LATREILLE, P.A. (1809): Genera crustaceorum et insectorum secundum ordinem naturalem in familias

disposita, iconibus exemplisque plurimis explicata. - Vol. 4. Paris and Strasbourg: 399 pp., 4 pls.LEDUC, J.W. & F.P. PINHEIRO (1989): Oropouche fever. - In: The arboviruses: epidemiology and

ecology: 1-14. Volume 4. Chapter 36. CRC Press, Florida: 243 pp.LENZ, F. (1934): 13a. Heleidae (Ceratopogonidae). - In: LINDNER, E. (ed.): Die Fliegen der

palaearktischen Region 3: 95-133 (Lfg. 78). Schweizerbart, Stuttgart.LINLEY, J.R. (1965a): The ovarian cycle and egg stage in Leptoconops (Holoconops) becquaerti (sic) (KIEFF.)

(Diptera, Ceratopogonidae). - Bull. Entomol. Res. 56: 37-56.LINLEY, J.R. (1965b): Descriptions of the pupae of Culicoides barbosai, C. insignis, and C. borinqueni

(Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 58: 57-63.LINLEY, J.R. (1968): Autogeny and polymorphism for wing length in Leptoconops becquaerti (KIEFF.)

(Diptera: Ceratopogonidae). - J. Med. Entomol. 5: 53-66.LINLEY, J.R. (1970): Description of the pupa of Culicoides floridensis. - Ann. Entomol. Soc. Am. 63:

1016-1019.LINLEY, J.R. & J.B. DAVIES (1971): Sand flies and tourism in Florida and the Bahamas and Caribbean

area. - J. Econ. Entomol. 64: 264-278.LINLEY, J.R. & D.S. KETTLE (1964): A description of the larvae and pupae of Culicoides furens POEY, and

Culicoides hoffmani FOX (Diptera: Ceratopogonidae). - Ann. Mag. Nat. Hist., Ser. 13, 7: 129-149.LOEW, H. (1861): Diptera Americae septentrionalis indigena. - Berl. Entomol. Zeitsch. 5: 307-359.LOEW, H. (1866): Diptera Americae septentrionalis indigena. - Berl. Entomol. Zeitsch. 9: 127-186.LUTZ, A. (1913): Contribuição para o estudo das Ceratopogoninas hematófagas do Brazil. - Mem.

Inst. Oswaldo Cruz 5: 45-73, pls. 6-8.


LUTZ, A. (1914): Contribuição para o conhecimento das “Ceratopogoninas” do Brasil. - Mem. Inst.Oswaldo Cruz 6: 81-99, pls. 8-9.

LYNCH ARRIBÁLZAGA, F. (1893): Dipterología Argentina (Chironomidae). - Bol. Acad. Nac. Ciencias(Córdoba) 13: 211-258.

MACFIE, J.W.S. (1932): Ceratopogonidae from the wings of dragonflies. - Tijdsch. Entomol. 75: 265-283.MACFIE, J.W.S. (1934): A new British Forcipomyia (Diptera, Ceratopogonidae). - Entomol. Mo. Mag.

70: 144 (June).MACFIE, J.W.S. (1935a): Ceratopogonidae (Dipt.) from the river Amazon. - Stylops 4: 49-56.MACFIE, J.W.S. (1935b): A new ceratopogonid (Dipt.) from British Guiana. - Stylops 4: 265.MACFIE, J.W.S. (1936): Four species of Ceratopogonidae (Diptera) from the wings of insects. - Proc.

R. Entomol. Soc. London (B) 5: 227-230.MACFIE, J.W.S. (1937a): Ceratopogonidae from Trinidad. - Ann. Mag. Nat. Hist. Ser. 10, 20: 1-18.MACFIE, J.W.S. (1937b): Notes on Ceratopogonidae (Diptera). - Proc. R. Entomol. Soc. London (B)

6: 111-118.MACFIE, J.W.S. (1938): Notes on Ceratopogonidae (Diptera). - Proc. R. Entomol. Soc. London (B) 7:

157-166.MACFIE, J.W.S. (1939a): 5. Ceratopogonidae. - In: Ruwenzori Expedition 1934-5 (British Mus. Nat.

Hist.) 1(5): 81-102, pls. 25, 26.MACFIE, J.W.S. (1939b): A report on a collection of Brazilian Ceratopogonidae. - Rev. Entomol. 10:

137-219.MACFIE, J.W.S. (1939c): A key to the species of Ceratopogonidae akin to Macropeza MG. (Diptera). -

Trans. R. Entomol. Soc. London 89: 1-12, pl. 1.MACFIE, J.W.S. (1940a): Ceratopogonidae (Diptera) from north-eastern Brazil. - Proc. R. Entomol.

Soc. London (B) 9: 73-79.MACFIE, J.W.S. (1940b): Ceratopogonidae (Diptera) from British Guiana and Tinidad. Part 1. - Proc.

R. Entomol. Soc. London (B) 9: 179-186.MACFIE, J.W.S. (1940c): Ceratopogonidae (Diptera) from British Guiana and Tinidad. Part 2. - Proc.

R. Entomol. Soc. London (B) 9: 187-195.MACFIE, J.W.S. (1940d): A report on a collection of Ceratopogonidae (Diptera) from British Guiana.

- Entomol. Mo. Mag. 76: 23-32.MACFIE, J.W.S. (1940e): Forcipomyia furcifera, sp. n. (Dipt. Ceratopogonidae). - Rev. Entomol. 11: 920-922.MACFIE, J.W.S. (1940f): The genera of Ceratopogonidae. - Ann. Trop. Med. Parasitol. 34: 13-30.MACFIE, J.W.S. (1943): A new species of Dasyhelea (Diptera) from the Bahama Islands. - Proc. R.

Entomol. Soc. London (B) 12: 119-120.MACFIE, J.W.S. (1944): Ceratopogonidae collected in Trinidad from cacao flowers. - Bull. Entomol.

Res. 35: 297-300.MACFIE, J.W.S. (1948): Some species of Culicoides (Diptera, Ceratopogonidae) from the state of Chiapas,

Mexico. - Ann. Trop. Med. Parasitol. 42: 67-87.MACFIE, J.W.S. (1949): Notes on Ceratopogonidae. - Proc. R. Entomol. Soc. London (B) 18: 109-115.MACFIE, J.W.S. (1953): Ceratopogonidae from Costa Rica. - Beitr. zur Entomol. 3: 95-105.MALLOCH, J.R. (1915a): The Chironomidae, or midges, of Illinois, with particular reference to the

species occurring in the Illinois River. - Bull. Illinois St. Lab. Nat. Hist. 10: 275-543, 24 pls.MALLOCH, J.R. (1915b): Some additional records of Chironomidae for Illinois and notes on other

Illinois Diptera. - Bull. Illinois St. Lab. Nat. Hist. 11: 305-363, pls. 80-84.


MARINO, P.I. & G.R. SPINELLI (1999a): The subgenus Forcipomyia (Metaforcipomyia) in Argentina (Diptera,Ceratopogonidae). - Iheringia, sér. Zool. 86: 3-8.

MARINO, P.I. & G.R. SPINELLI (1999b): The species groups of the subgenus Forcipomyia (Forcipomyia)in the Neotropics, with a description of a new species of the genualis group (Diptera:Ceratopogonidae). - Trans. Am. Entomol. Soc. 125: 445-452 (Dec.).

MARINO, P. I. & G.R. SPINELLI (2001a): Las especies del subgénero Forcipomyia (Forcipomyia) en laPatagonia (Diptera, Ceratopogonidae). - Revta. Soc. Entomol. Argent. 60: 99-124.

MARINO, P.I. & G.R. SPINELLI (2001b): El subgenero Forcipomyia (Euprojoannisia) en la Patagonia (Diptera,Ceratopogonidae). - Gayana 65: 11-18.

MARINO, P.I. & G.R. SPINELLI (2001c): Los subgéneros de Forcipomyia, Thyridomyia y Synthyridomyia, enla Patagonia argentina, con la descipción de F. (S.) soibelzoni (Diptera, Ceratopogonidae). -Neotrópica 47: 13-16.

MARINO, P.I. & G.R. SPINELLI (2002): A revision of the Forcipomyia squamitibia group in the Neotropics withthe description of three new species (Diptera: Ceratopogonidae). - Ins. Sci. Appl. 22: 307-319.

MARINO, P.I. & G.R. SPINELLI (2003): The Patagonian species of the subgenus Forcipomyia(Metaforcipomyia) (Diptera: Ceratopogonidae), with a key to the New World species. - Ins. Syst.Evol. 34: 21-28.

MARINO, P.I. & G.R. SPINELLI (2004a): Further notes on the subgenus Forcipomyia (Forcipomyia) fromArgentinean Patagonia (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 130: 147-154.

MARINO, P.I. & G.R. SPINELLI (2004b): Descriptions of the Patagonian species of the subgenusTrichohelea of Forcipomyia, with a key to the Neotropical species (Diptera: Ceratopogonidae). - J.Nat. Hist. 38: 2251-2262.

MARINO, P.I. & G.R. SPINELLI (2004c): A new species of Atrichopogon KIEFFER from northern Argentina(Diptera: Ceratopogonidae). - Entomol. News 114: 156-159 (2003).

MARINO, P.I. & N. VON ELLENRIEDER (1999): New records of Forcipomyia (Pterobosca) incubans (MACFIE)(Diptera: Ceratopogonidae) on libellulids (Anisoptera). - Not. Odonatol. 5: 38-39.

MARINO, P.I., SPINELLI, G.R. & C.G. CAZORLA (2002): Type-specimens of Ceratopogonidae (Insecta:Diptera) in the collection of the Museo de La Plata, Argentina. - Fac. Cs. Nat. Mus. Univ. Nac.La Plata. Pub. Tec. Did. 42: 1-37.

MATTA, J.F. (1967): A new species of Culicoides (Diptera, Ceratopogonidae) from Honduras. - Fla.Entomol. 50: 75-77.

MAYER, K. (1934): Die Metamorphose der Ceratopogonidae (Dipt.). Ein Beitrag zur Morphologie,Systematik, Ökologie und Biologie der Jugendstadien dieser Dipterenfamilie. - Arch. Natur. 3:205-288.

MAYER, K. (1937): Beitrag zur Kenntnis der deutschen Ceratopogoniden. (Dipt.) IX. - Stett. Entomol.Zeit. 98: 301-304.

MAYER, K. (1952): Die Macropeza-Gruppe der Heleiden (Diptera: Heleidae). - Beitr. zur Entomol. 2:582-585.

MAYER, K. (1959): Die Puppen brasilianischer Heleiden (Diptera). - Deut. Entomol. Zeitsch. 6: 230-233.MCALPINE, J.F., PETERSON, B.V., SHEWELL, G.E., TESKEY, H.J., VOCKEROTH, J.R. & D.M. WOOD (1981)

(eds.): Manual of Nearctic Diptera. - Volume 1. Agriculture Canada Monograph 27.Biosystematic Research Institute, Ottawa: 674 pp.

MEIGEN, J.G. (1818): Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten.- Vol. 1 : xxxvi + 333 pp., pls. 1-11. Aachen.


MELLOR, P.S., BOORMAN, J. & M. BAYLIS (2000): Culicoides biting midges: their role as arbovirus vectors.- Ann. Rev. Entomol. 45: 307-340.

MERCER, D.R., SPINELLI, G.R., WATTS, D.M. & R.B. TESH (2003): Biting rates and developmentalsubstrates for biting midges (Diptera: Ceratopogonidae) in Iquitos, Peru. - J. Med. Entomol.40: 807-812.

MESSERSMITH, D.H. (1972): A new species of Culicoides from Colombia (Diptera: Ceratopogonidae).- Proc. Entomol. Soc. Washington 74: 165-169.

MINAYA, G. (1978): Dasyhelea mediomunda, sp. n. (Diptera: Ceratopogonidae) de la costa central delPerú. - Rev. Per. Entomol. 21: 79-81.

MO, C.L., THOMSON, L.H., HOMAN, E.J., OVIEDO, M.T., GREINER, E.C., GONZÁLEZ, J. & M.R. SÁENZ.(1994): Bluetongue virus isolations from vectors and ruminants in Central America and theCaribbean. - Am. J. Vet. Res. 55: 211-215.

MURPHREE, C.S. & G.R. MULLEN (1991): Comparative larval morphology of the genus CulicoidesLATREILLE (Diptera: Ceratopogonidae) in North America with a key to species. - Bull. Soc.Vector Ecol. 16: 269-399.

NEWMAN, E. (1834): Attempted division of British insects into natural orders. - Entomol. Mag. 2: 379-431.NIELSEN, A. (1951): Contributions to the metamorphosis and biology of the genus Atrichopogon

KIEFFER (Diptera, Ceratopogonidae) with remarks on the evolution and taxonomy of the genus.- Det Kong. Dan. Viden. Selsk., Biol. Skr. 6: 1-95, pls. 1-2.

NOÈ, G. (1905): Un nuovo genere appartenente alla famiglia Chironomidae. - Rendiconti, Accad.Naz. Lincei 14: 114-120.

NOÈ, G. (1907): Due nuove specie di ditteri appartenenti ad un genere nuovo. - Arch. Zool. Italiano:Pubb. Sotto Ausp. Un. Zool. 3: 101-164, 3 pls.

OIE (Office International des Epizooties) (2001): Lengua azul en Brasil. - Inf. Sanit. 14: 10-11.ORTÍZ, I. (1950a): Estudios en Culicoides II. Diptera, Ceratopogonidae. Culicoides lanei n. sp. de Panamá.

- Rev. San. Asist. Soc. 15: 431-433.ORTÍZ, I. (1950b): Estudios en Culicoides IV. Revisión de las especies americanas del sub-género Hoffmania

FOX 1948, con la descripción de dos nuevas especies. - Rev. San. Asist. Soc. 15: 437-460.ORTÍZ, I. (1950c): Estudios en Culicoides V. Informes sobre una nueva especie y lista de los machos

cuyas genitalias son conocidas. - Rev. San. Asist. Soc. 15: 461-465.ORTÍZ, I. (1950d): Descripción de Stilobezzia glauca venezuelensis n. vr. y Monohelea mayeri n. sp. - Mem.

Soc. Cs. Nat. La Salle 10: 199-204.ORTÍZ, I. (1951a): Estudios en Culicoides (Diptera, Ceratopogonidae). 6. Culicoides bricenoi n. sp. - Bol.

Lab. Clin. “Luis Razetti” 11: 442-448.ORTÍZ, I. (1951b): Estudios sobre Culicoides (Diptera, Ceratopogonidae) VII.- Culicoides dominicii n. sp.

y clave para el reconocimiento de las especies venezolanas. - Nov. Cient., Contrib. Ocasion.Mus. Hist. Nat. La Salle, Ser. Zool. 5: 1-12.

ORTÍZ, I. (1951c): Estudios en Culicoides (Diptera, Ceratopogonidae) 9. Sobre los caracteres diferencialesentre Culicoides paraensis (GOELDI, 1905), C. stellifer (COQUILLETT, 1901), y C. lanei (ORTÍZ, 1950).Descipción de cuatro nuevas especies con la redescripción de algunas otras poco conocidas. -Rev. San. Asist. Soc. 16: 573-591.

ORTÍZ, I. (1952a): Los insectos del género Leptoconops SKUSE, 1889 en Venezuela. Descripción de unanueva especie: Leptoconops (L.) venezuelensis y redescripción de la hembra de Leptoconops (L.)torrens) (TOWNSEND, 1893). - Rev. San. Asist. Soc. 17: 163-171.


ORTÍZ, I. (1952b): Apuntes en Ceratopogonidae (Diptera Nematocera). 3. Sobre una nueva especiede Monohelea KIEFFER 1917 (Monohelea ocumare n. sp.). - Rev. San. Asist. Soc. 17: 253-256.

ORTÍZ, I. (1953a): Nueva contribución al conocimiento de los caracteres morfológicos externos delas hembras americanas del género Culicoides LTR. (Diptera, Ceratopogonidae) con unaespermateca. Descripción de dos nuevas especies de Venezuela: Culicoides transferrans (C.oublepharus [sic] ORTÍZ, 1952) y Culicoides mirsae. - Rev. San. Asist. Soc. 18: 797-806.

ORTÍZ, I. (1953b): Sobre una nueva especie del género Culicoides (Diptera, Ceratopogonidae) próximade C. horticola LUTZ 1913. - Rev. San. Asist. Soc. 18: 807-812.

ORTÍZ, I. (1954): Sobre dos nuevos dípteros hematófagos del género Culicoides (Nematocera,Ceratopogonidae). - Arch. Venez. Patol. Trop. Parasitol. Med. 2: 221-226.

ORTÍZ, I. (1956): Sobre un pequeño díptero de la región del Auyantepuy. - Bol. Venez. Lab. Clin. 1: 93-96.ORTÍZ, I. (1957): Nuevos representantes hematófagos de los géneros Culicoides (Diptera: Ceratopogonidae)

y Simulium (Diptera: Simuliidae) de Venezuela. - Bol. Venez. Lab. Clin. 2: 161-168.ORTÍZ, I. (1959): Contribución al estudio de los Ceratopogonidos (Jejenes) hematófagos de Venezuela

(Leptoconops SKUSE, 1890, Lasiohelea KIEFFER, 1921 y Culicoides LATREILLE, 1809) (Diptera:Ceratopogonidae). - Rev. San. Asist. Soc. 24: 349-369.

ORTÍZ, I. (1961): Descripción de una nueva especie de Culicoides (C. forattinii sp. n.) del subgéneroOecacta POEY, 1851, de Venezuela. - Rev. Bras. Entomol. 10: 211-215.

ORTÍZ, I. (1968): Los Dípteros hematófagos del género Culicoides en Venezuela (Diptera,Ceratopogonidae). Part I. Hembras da una sola espermateca con la descripción de una nuevaespecie (C. rodriguezi n. sp.). - Rev. Inst. Nac. Hig. 1: 65-71.

ORTÍZ, I. & L.A. LEÓN (1955): Los Culicoides (Diptera: Ceratopogonidae) de la República del Ecuador.- Bol. Inf. Cient. Nac. 7: 564-594 (January-February).

ORTÍZ, I. & M. MIRSA (1950): Estudios en Culicoides (Diptera, Ceratopogonidae) VIII.- Sobre unanueva especie del grupo haematopotus: Culicoides venezuelensis n. sp. - Arch. Venez. Patol. Trop.Parasitol. Med. 2: 137-143.

ORTÍZ, I. & M. MIRSA (1951): Estudios en Culicoides. Descripción de dos nuevas especies: Culicoidesavilaensis y C. discrepans, y del macho de C. leopoldoi ORTÍZ, 1951. Redescripción de C. limaiBARRETTO, 1944, C. baueri HOFFMAN 1925, C. lichyi FLOCH and ABONNENC, 1949, y C. pusillusLUTZ, 1913. - Rev. San. Asist. Soc. 16: 593-605.

ORTÍZ, I. & M. MIRSA (1952a): Culicoides de Venezuela - Redescripción de 10 especies con la descripciónde algunos sexos no conocidos. - Rev. San. Asist. Soc. 17: 257-279.

ORTÍZ, I. & M. MIRSA (1952b): Sobre las especies Americanas del género “Culicoides” LATR., (Diptera,“Ceratopogonidae”) con una espermateca. - Acta Cient. Venez. 2: 125-128.

PAINTER, R.H. (1927): The biology, immature stages, and control of the sand flies (bitingCeratopogoninae) at Puerto Castilla, Honduras. - 1926 Ann. Rep. Med. Dept. Unit. Fruit Comp.15: 245-262.

PALCHICK, S.M. (1981): Morphological descriptions of a new species of Bezzia and pupae of sixspecies of Bezzia and three species of Nilobezzia (Diptera: Ceratopogonidae). - M.Sc.-thesis,University of Wisconsin, Madison: viii + 28 pp., 12 plates.

PERRUOLO, G. (1990): Culicoides (Diptera, Ceratopogonidae) de Venezuela. Descripción de una nuevaespecie: Culicoides malariologiensis n. sp. - Bol. Dir. Malar. San. Amb. 28: 27-31 (1988).

PERRUOLO, G. (2001): Nueva especie de Culicoides (Diptera: Ceratopogonidae), de la región Andina,Venezuela. - Bol. Dir. Malar. San. Amb. 41: 34-37.


PHILIPPI, R.A. (1865): Aufzählung der chilenischen Dipteren. - Verh. Kaiser.-König. Zool.-Botan.Gesell. Wien 15: 595-782.

PIERCE, W.D. (1966): Fossil arthropods of California. 29. Silicified Miocene pupae of ceratopogonidflies. - Bull. South. Calif. Acad. Sci. 65: 81-98.

PINHEIRO, F.P., TRAVASSOS DA ROSA, A.P.A. & F.C. VASCONCELOS (1998): An overview of oropouche feverepidemics in Brazil and neighbour countries. - In: TRAVASSOS DA ROSA, A.P.A. et al. (eds.): An overviewof arbovirology in Brazil and neighbouring countries. Instituto Evandro Chagas, Belém: 296 pp.

POEY, F. (1853): Memorias sobre la historia natural de la Isla de Cuba, acompañadas de sumarioslatinos y extractos en francés. - Vol. 1 (pt. 4): 201-280, Havanna.

RAFINESQUE, C.S. (1815): Analyse de la nature ou tableau de l’univers et des corps organisés. Lenature est mon guide, et LINNÉUS mon maitre.- Palmero, privately published: 224 pp.

REMM, H. (1971): On the fauna of Ceratopogonidae of South Primorye (Ussuri Land) [in Russian,English summary]. - In: Living Nature of the Far East: 182-220, Akademiia nauk EstonskoiSSR: 240 pp.

REMM, H. (1975): On the classification of the biting midges (Diptera, Ceratopogonidae) (in Russian).- Parazitol. 9: 393-397.

ROBACK, S.S. (1957): A new Palpomyia from Peru (Diptera: Heleidae). - Not. Nat. Acad. Nat. Sci.Philadelphia 297: 1-2.

RODRÍGUEZ, M.C. DE & W.W. WIRTH (1986): A new species of man-biting Culicoides from the highAndes of Colombia (Diptera: Ceratopogonidae). - Fla. Entomol. 69: 311-314.

RONDEROS, M.M. (1988): Estudio sistemático de los géneros hematófagos de Ceratopogonidae(Diptera: Nematocera) de la República Argentina. - Ph.D.-thesis, Univ. Nac. La Plata, Argentina:135 pp. + 49 pls.

RONDEROS, M.M. (1990a): Dos especies nuevas del género Culicoides para la Argentina y Uruguay(Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 48: 115-120.

RONDEROS, M.M. (1990b): A new species of Lepteconops [sic] from Argentina (Diptera,Ceratopogonidae). - Rev. Bras. Entomol. 34: 423-426.

RONDEROS, M.M. & G.R. SPINELLI (1989): The pupa of the biting midge Culicoides venezuelensis ORTÍZ

and MISRA 1950 (Diptera: Ceratopogonidae). - Biota 5: 77-81.RONDEROS, M.M. & G.R. SPINELLI (1992): A key to the Neotropical Leptoconops (Diptera:

Ceratopogonidae), with the description of a new species from Argentina. - Revta. Soc. Entomol.Argent. 51: 41-45.

RONDEROS, M.M. & G.R. SPINELLI (1993): Dos especies nuevas de Leptoconops neotropicales, ydescripcion del macho de Leptoconops (Leptoconops) chilensis (Diptera: Ceratopogonidae). - GayanaZool. 57: 305-308.

RONDEROS, M.M. & G.R. SPINELLI (1994): Dos especies nuevas de Culicoides de la Argentina (Diptera:Ceratopogonidae). - Revta. Soc. Entomol. Argent. 53: 47-50.

RONDEROS, M.M. & G.R. SPINELLI (1995a): Two new neotropical species of Culicoides from the ParanáRiver Basin of Argentina and Paraguay (Diptera: Ceratopogonidae). - Trans. Am. Entomol.Soc. 121: 59-63.

RONDEROS, M.M. & G.R. SPINELLI (1995b): Redescripción de Culicoides lahillei y descripción de Culicoides ichesi,n. sp. de la Argentina, Paraguay y Uruguay (Diptera: Ceratopogonidae). - Neotrópica 41: 77-81.

RONDEROS, M.M. & G.R. SPINELLI (1997): Culicoides patagoniensis n. sp. del sur de Argentina y Chile(Diptera: Ceratopogonidae). - Gayana Zool. 61: 33-39.


RONDEROS, M.M. & G.R. SPINELLI (1998): Las especies de Culicoides en el área de influencia de larepresa de Yacyretá (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 57: 79-83.

RONDEROS, M.M. & G.R. SPINELLI (1999): On the subgenus Forcipomyia (Lasiohelea) in the NeotropicalRegion (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc. 125: 151-161(June).

RONDEROS, M.M. & G.R. SPINELLI (2000): The larva and pupa of Culicoides bambusicola LUTZ observedwith SEM, and additional notes on the adult (Diptera Ceratopogonidae). - Trans. Am. Entomol.Soc. 126: 133-144.

RONDEROS, M.M., DÍAZ, F. & G.R. SPINELLI (2004a): A new species of Dasyhelea KIEFFER fromArgentina described as adult and pupa (Diptera: Ceratopogonidae). - Trans. Am. Entomol.Soc. 103: 193-200.

RONDEROS, M.M., SPINELLI, G.R. & F. DÍAZ (2004b): Description of larva and redescription of pupa andadult of Palpomyia guarani (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 63: 45-54.

RONDEROS, M.M., SPINELLI, G.R. & F. DÍAZ (2006): Observations on the immatures of Dasyheleanecrophila SPINELLI & RODRÍGUEZ in laboratory (Diptera: Ceratopogonidae). - Trans. Am. Entomol.Soc. 132: 291-297.

RONDEROS, M.M., SPINELLI, G.R., HUERTA, H. & F. DÍAZ (2003): Immature stages of two Neotropicalspecies of Dasyhelea KIEFFER, 1911 (Diptera: Ceratopogonidae). - Trans. Am. Entomol. Soc.129: 295-308.

RONDEROS, M.M., SPINELLI, G.R. & P. SARMIENTO (2000): Preparation and mounting of biting midgesof the genus Culicoides LATREILLE (Diptera: Ceratopogonidae) to be observed with a scanningelectron microscope. - Trans. Am. Entomol. Soc. 126: 125-132.

ROOT, F.M. & W.A. HOFFMAN (1937): The North American species of Culicoides. - Am. J. Hyg. 25:150-176, pls. 1-8.

SÁENZ, M.R. & E.C. GREINER (1994): Culicoides aspirated from cattle in Costa Rica, Honduras, Panamaand Puerto Rico, and their role as potential vectors of bluetongue viruses. - Med. Vet. Entomol.8: 15-19.

SAUNDERS, L.G. (1925): On the life history, morphology and systematic position of Apelma KIEFF.and Thyridomyia n. g. (Diptera, Nemat. Ceratopogoninae). - Parasitol. 17: 252-277.

SAUNDERS, L.G. (1957): Revision of the genus Forcipomyia based on characters of all stages (Diptera,Ceratopogonidae). - Can. J. Zool. 34: 657-705 (1956).

SAUNDERS, L.G. (1959): Methods for studying Forcipomyia midges, with special reference to cacao-pollinating species (Diptera, Ceratopogonidae). - Can. J. Zool. 37: 33-51.

SAUNDERS, L.G. (1964): New species of Forcipomyia in the Lasiohelea complex described in all stages(Diptera, Ceratopogonidae). - Can. J. Zool. 42: 463-482.

SAUNDERS, L.G. & G.F. BOWMAN (1956): Cacao pollination in Costa Rica. - Banco de Costa Rica, SanJose: 13 pp.

SAY, T. (1825): American entomology, or descriptions of the insects of North America. - Vol. 2.Philadelphia: 121 pp., pls. 19-36.

SÉGUY, E. (1941): Quelques Cératopogonides vulnérants parasites des insectes. - Rev. Franc. Entomol.8: 82-88.

SHANNON, R.C. & E. DEL PONTE (1927): Cuatro notas sobre especies nuevas de Dipteros, Nematoceros,hematofagos o no, de la Republica Argentina. - Rev. Inst. Bacterial. Dept. Nac. Hig. 4: 724-736.

SKUSE, F.A.A. (1889): Diptera of Australia. Part VI. - The Chironomidae. - Proc. Linn. Soc. N. S. W.4: 215-311, pls. 11-14.


SORIA, S. & P.G. BYSTRAK (1975): A new species of Forcipomyia (Diptera, Ceratopogonidae) describedin all stages, with an account of its role as a cacao pollinator. - Rev. Theobroma (Brasil) 5: 3-11.

SPINELLI, G.R. (1982): Cuatro especies nuevas del género Artichopogon [sic] KIEFER (Diptera:Ceratopogonidae) en la República Argentina. - Revta. Soc. Entomol. Argent. 41: 201-210.

SPINELLI, G.R. (1983a): Notas sobre Ceratopogonidae de la República Argentina (Diptera, Nematocera)II. Nuevos aportes al conocimiento del género Forcipomyia MEIGEN. - Neotrópica 29: 121-129.

SPINELLI, G.R. (1983b): Notas sobre Ceratopogonidae (Diptera, Nematocera) de la República ArgentinaI. Una nueva especie del género Alluaudomyia KIEFFER, redescripción de Dasyhelea penthesileaeMACFIE, y nuevas citas para el género Stilobezzia KIEFFER. - Limnobios 2: 403-411.

SPINELLI, G.R. (1983c): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina.III. Nuevos aportes al conocimiento de la tribu Palpomyiini. - Limnobios 2:17-24.

SPINELLI, G.R. (1984): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina.IV. Descripción de adulto y pupa de Neobezzia termophila sp. nov. - Neotrópica 30: 197-200.

SPINELLI, G.R. (1987): Notas sobre Ceratopogonidae (Diptera: Nematocera) de la República Argentina.VI. Las hembras de Paradasyhelea brevipalpis y de Macrurohelea paracaudata. - Limnobios 2: 667-670.

SPINELLI, G.R. (1988): Two new species of Neotropical Alluaudomyia KIEFER, 1913 (Insecta, Diptera,Certopogonidae [sic]). - Iheringia, Ser. Zooll. 68: 129-136.

SPINELLI, G.R. (1989): Amerohelea similis n. sp., from Argentina and Uruguay (Diptera: Ceratopogonidae).- Rev. Asoc. Cs. Nat. Litoral 20: 25-28.

SPINELLI, G.R. (1990): The genus Brachypogon in Argentina, with a key to the Neotropical species(Diptera, Ceratopogonidae). - Rev. Bras. Entomol. 34: 743-755.

SPINELLI, G.R. (1994): The male of Physohelea turgidipes (Diptera: Ceratopogonidae). - Neotrópica 40:73-74.

SPINELLI, G.R. (1996): A new species in the genus Diaphanobezzia from subantarctic Argentina (Diptera:Ceratopogonidae). - Neotrópica 42: 77-79.

SPINELLI, G.R. (1997a): A new genus of the tribe Sphaeromiini from subantarctic Argentina andChile related to Mackerrasomyia DEBENHAM (Diptera: Ceratopogonidae). - Mem. Entomol. Soc.Washington 18: 224-229.

SPINELLI, G.R. (1997b): Further notes on Neotropical Ceratopogonini (Diptera: Ceratopogonidae). -Revta. Soc. Entomol. Argent. 56:123-124.

SPINELLI, G.R. (1998): Three new species and new records of Neotropical Stenoxenini (Diptera:Ceratopogonidae). - Neotrópica 44: 51-55.

SPINELLI, G.R. & A. BORKENT (2004a): New species of Central American Culicoides LARTREILLE (Diptera:Ceratopogonidae) with a synopsis of species from Costa Rica. - Proc. Entomol. Soc. Washington106: 361-395.

SPINELLI, G.R. & A. BORKENT (2004b): A new species and first record of the subgenus Forcipomyia(Schizoforcipomyia) CHAN and LEROUX from the Neotropical Region (Diptera: Ceratopogonidae).- Zootaxa 572: 1-8.

SPINELLI, G.R. & C.G. CAZORLA (2004): A new species of Brachypogon (Isohelea) from arid zones ofArgentina (Diptera: Ceratopogonidae). - Zootaxa 570: 1-6.

SPINELLI, G.R. & C.G. CAZORLA (2006): A new predaceous midge of the Palpomyia tibialis group fromnortheastern Argentina (Diptera: Ceratopogonidae). - Rev. Biol. Trop. 54: 1067-1070.

SPINELLI, G.R. & A. DIPPOLITO (1995): Two new neotropical species of the Forcipomyia (F.) argenteolaspecies group (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 54: 155-158.


SPINELLI, G.R. & J.P. DURET (1993): Las especies neotropicales de Clinohelea (Diptera: Ceratopogonidae).- Graellsia 49: 39-50.

SPINELLI, G.R. & M.L. FELIPPE-BAUER (1990a): Two new Neotropical species of predaceous midgesof the tribe Sphaeromiini (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 85: 87-90.

SPINELLI, G.R. & M.L. FELIPPE-BAUER (1990b): Sphaerohelea, a new Neotropical predaceous midgegenus of the tribe Sphaeromiini (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 85:195-198.

SPINELLI, G.R. & W.L. GROGAN (1984): Three new species of Macrurohelea from Argentina with a keyto the Neotropical species (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 86:961-967.

SPINELLI, G.R. & W.L. GROGAN (1985): Clastrieromyia, a new Neotropical genus of predaceous midgesrelated to Palpomyia and Bezzia (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington87: 329-334.

SPINELLI, G.R. & W.L. GROGAN (1986): Two new species of the predaceous midge genus Clastrieromyiafrom Uruguay with a new record of C. schnacki for Argentina (Diptera: Ceratopogonidae). -Proc. Entomol. Soc. Washington 88: 455-460.

SPINELLI, G.R. & W.L. GROGAN (1987): A revision of the Neotropical species of Parabezzia (Diptera:Ceratopogonidae). - Biol. Acuat. 11: 1-45.

SPINELLI, G.R. & W.L. GROGAN (1989): Dos especies nuevas de Palpomyia, grupo distincta de la regiónNeotropical (Diptera: Ceratopogonidae). - Neotrópica 35: 3-8.

SPINELLI, G.R. & W.L. GROGAN (1990): New species of predaceous midges of the tribe Ceratopogoninifrom subantarctic Argentina (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington92: 127-134.

SPINELLI, G.R. & W.L. GROGAN (1993): Borkenthelea, a new predaceous midge genus from subantarcticArgentina (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 95: 321-326.

SPINELLI, G.R. & W.L. GROGAN (1994): Two new Neotropical species of the subgenus Isohelea ofBrachypogon, with the description of the female of B. (I.) misionensis and a key to the Neotropicalspecies (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 53: 1-8.

SPINELLI, G.R. & W.L. GROGAN (1998): A revision of the Neotropical predaceous midges of Brachypogon(Brachypogon) KIEFFER (Diptera: Ceratopogonidae). - Insecta Mundi 12: 59-79.

SPINELLI, G.R. & W.L. GROGAN (1999): A new species of Macrurohelea INGRAM and MACFIE, and newrecords of biting midges of the tribes Culicoidini and Ceratopogonini (Diptera: Ceratopogonidae)from Tierra del Fuego and the Magallanes. - Proc. Entomol. Soc. Washington 101: 708-713.

SPINELLI, G.R. & W.L. GROGAN (2001): A revision of the Patagonian predaceous midge genusBorkenthelea SPINELLI and GROGAN (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington103: 147-156.

SPINELLI, G.R. & W.L. GROGAN (2003): A revision of the Neotropical biting midges of the genusParadasyhelea INGRAM and MACFIE (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington105: 568-577.

SPINELLI, G.R. & P.I. MARINO (1997): Two new species of the subgenus Thyridomyia of Forcipomyiafrom Argentina and new records of F. (Synthyridomyia) sanctaeclarae (Diptera: Ceratopogonidae).- Trans. Am. Entomol. Soc. 123: 187-190.

SPINELLI, G.R. & M.E. MARTINEZ (1992): The genus Culicoides in Uruguay (Diptera: Ceratopogonidae).- Insecta Mundi 5: 175-179 (1991).


SPINELLI, G.R. & E.A. RODRÍGUEZ (1999): A new species of Dasyhelea from Argentina described asadult, pupae and fourth instar larvae (Diptera: Ceratopogonidae). - Neotrópica 45: 59-62.

SPINELLI, G.R. & M.M. RONDEROS (1987): Notas sobre Ceratopogonidae (Diptera: Nematocera) de laRepública Argentina. V. Nuevos aportes al conocimiento del género Dasyhelea KIEFFER. -Neotrópica 33: 11-17.

SPINELLI, G.R. & M.M. RONDEROS (1991): Los polvorines del género Culicoides en area de influenciade la represa de Salto Grande (Diptera: Ceratopogonidae). - Neotrópica 37: 83-94.

SPINELLI, G.R. & M.M. RONDEROS (2001): First record of the genus Bezzia in Chile, with a description ofa new species of the venustula group (Diptera: Ceratopogonidae). - Rev. Chil. Hist. Nat. 74: 751-754.

SPINELLI, G.R. & W.W. WIRTH (1981): A new species of predaceous midge of the genus Bezzia KIEFFER

from Argentina (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 40: 187-192.SPINELLI, G.R. & W.W. WIRTH (1984a): Ocho especies nuevas del género Culicoides LATREILLE de la

región Neotropical. Primera descripción del macho de C. flinti WIRTH, y de la hembra de C. lentiTAVARES y LUNA DIAS (Diptera: Ceratopogonidae). - Revta. Soc. Entomol. Argent. 43: 171-185.

SPINELLI, G.R. & W.W. WIRTH (1984b): A review of the Neotropical predaceous midge genusParyphoconus (Diptera: Ceratopogonidae). - Proc. Biol. Soc. Washington 97: 882-908.

SPINELLI, G.R. & W.W. WIRTH (1984c): The Neotropical predaceous midges of the genus Alluaudomyia(Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 86: 673-702.

SPINELLI, G.R. & W.W. WIRTH (1984d): Las especies Neotropicales del género Dasyhelea, grupo cincta(Diptera: Ceratopogonidae). - Limnobios 2: 586-608.

SPINELLI, G.R. & W.W. WIRTH (1986a): The Neotropical species of Phaenobezzia (Diptera:Ceratopogonidae). - Fla. Entomol. 69: 231-236.

SPINELLI, G.R. & W.W. WIRTH (1986b): Clave para la identificación de las especies del género CulicoidesLATREILLE presentes al sur de la cuenca Amazónica. Nuevas citas y notas sinonímicas (Diptera:Ceratopogonidae). - Revta. Soc. Entomol. Argent. 44: 49-73.

SPINELLI, G.R. & W.W. WIRTH (1989a): Las especies Neotropicales del género Bezzia (Diptera:Ceratopogonidae). II. Los grupos nobilis y punctipennis. - Revta. Soc. Entomol. Argent. 45: 109-129.

SPINELLI, G.R. & W.W. WIRTH (1989b): The Neotropical predaceous midges of the genus Bezzia(Diptera: Ceratopogonidae) Part I. The glabra and brevicornis groups. - Limnobios 2: 762-778.

SPINELLI, G.R. & W.W. WIRTH (1990): Neotropical predaceous midges of the genus Bezzia (Diptera:Ceratopogonidae) Part III. The gibbera group of species. - Insecta Mundi 4: 11-32.

SPINELLI, G.R. & W.W. WIRTH (1991): The Neotropical predaceous midges of the genus Bezzia (Diptera:Ceratopogonidae) Part IV. The dentifemur and venustula groups. - Insecta Mundi 5: 1-17.

SPINELLI, G.R. & W.W. WIRTH (1993): Los Ceratopogonidae de la Argentina (Insect: Diptera). - In:CASTELLANOS, Z. (ed.). Fauna de agua dulce de la República Argentina. Volumen 38. DipteraFascículo 3. Ceratopogonidae: 124 pp.

SPINELLI, G.R., DIPPOLITO, A. & W.W. WIRTH (1995): A report on a collection of Ceratopogonidae (Diptera)from Rondonia, Brazil. 2. Tribes Heteromuiini [sic] and Sphaeromiini. - Insecta Mundi 9: 165-169.

SPINELLI, G.R., GREINER, E.C. & W.W. WIRTH (1993): The Neotropical bloodsucking midges of theCulicoides guttatus group of the subgenus Hoffmania (Diptera: Ceratopogonidae). - Contrib. Am.Entomol. Inst. 27: 1-91.

SPINELLI, G.R., MARINO, P.R. & P. POSADAS (2006): The Patagonian species of the genus AtrichopogonKIEFFER, with a biogeographic analysis based on Forcipomyiinae (Diptera: Ceratopogonidae).- Ins. Syst. and Evol. 37: 301-324.


SPINELLI, G.R., MARINO, P.I. & M.M. RONDEROS (2005a): The fourth instar larva and pupa of theNeotropical biting midge Forcipomyia (Forcipomyia) rioplatensis MARINO and SPINELLI (Diptera:Ceratopogonidae). - Proc. Entomol. Soc. Washington 107: 108-114.

SPINELLI, G.R., RONDEROS, M.M. & E.G. BALSEIRO (1989): Análisis de la actividad diaria de dípterosnematocéros en Punta Lara (Pdo. de Ensenada, Prov. de Buenos Aires). II. Ceratopogonidae,con la descripción de una especie nueva en el género Atrichopogon. - Limnobios 2: 733-737.

SPINELLI, G.R., RONDEROS, M.M., DÍAZ, F. & P.I. MARINO (2005b): The bloodsucking biting midgesof Argentina (Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 100: 137-150.

SPINELLI, G.R., SCHAEFER, E.F. & A.I. KEHR (2002): First record of biting midges (Diptera: Ceratopogonidae)attacking frogs in the Neotropical Region. - Proc. Entomol. Soc. Washington 104: 527-528.

STAEGER, R.C. (1839): Systematisk fortegnelse over de i Danmark hidtil fundne Diptera. - Naturhist.Tids. 2: 549-600.

SZADZIEWSKI, R. (1996): Biting midges from Lower Cretaceous amber of Lebanon and UpperCretaceous Siberian amber of Taimyr (Diptera, Ceratopogonidae). - Studia Dipterol. 3: 23-86.

SZADZIEWSKI, R., KRZYWINSKI, J. & W. GILKA (1997): Diptera Ceratopogonidae, biting midges. In: NILSSON,A.N. (ed.). Aquatic insects of North Europe - A taxonomic handbook. Vol. 2: 243-263.

TAVARES, O. & A.P. LUNA DIAS (1980): Quatro novas espécies do gênero Culicoides LATREILLE, 1809,do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 40: 393-399.

TAVARES, O. & R.A. RUIZ (1980): Sobre duas espécies novas do gênero Culicoides LATREILLE, 1809, doBrasil (Diptera, Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 75: 27-32.

TAVARES, O. & M.A. DE SOUZA (1978): Sobre três espécies novas do gênero Culicoides LATREILLE, 1809do Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 38: 619-624.

TAVARES, O. & M.A. DE SOUZA (1979): Sobre algumas espécies do gênero Culicoides LATREILLE, 1809do Estado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 39: 611-613.

TAVARES, O. & M.A. DE SOUZA (1980): Duas espécies novas do gênero Monohelea KIEFFER, 1917, doEstado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 40: 95-100.

TAVARES, O. & A.J. SILVA PEREIRA (1978): Duas espécies novas do gênero Monohelea KIEFFER, 1917, doEstado do Rio de Janeiro, Brasil (Diptera, Ceratopogonidae). - Rev. Bras. Biol. 38: 157-160.

THIENEMANN, A. (1928): Chironomiden - Metamorphosen I. - Arch. Hydrobiol. 19: 585-623.THOMSEN, L.C. (1937): Aquatic Diptera Part V. Ceratopogonidae. - Cornell Univ. Agric. Exp. Sta.

Mem. 210: 57-80, pls. 10-18.TOKUNAGA, M. (1961): Notes on biting midges III. - Pub. Ent. Lab., Univ. Osaka Pref. 6: 115-122.TOWNSEND, C.H.T. (1893): An interesting blood-sucking gnat of the family Chironomidae. - Psyche

6: 369-371.UTMAR, J.A. & W.W. WIRTH (1976): A revision of the new world species of Forcipomyia, subgenus

Caloforcipomyia (Diptera: Ceratopogonidae). - Fla. Entomol. 59: 109-133.VARGAS, L. (1953a): Beltranmyia n. subgen. de Culicoides (Insecta: Heleidae). - Rev. Inst. Sal. Enf. Trop.

13: 33-36.VARGAS, L. (1953b): Culicoides wirthomyia n. sp. y Culicoides stigmalis WIRTH, 1952 (Insecta, Diptera). -

Rev. Inst. Sal. Enf. Trop. 13: 227-233.VARGAS, L. (1954): Dos nuevas especies de Culicoides Mexicanos (Diptera, Heleidae). - Rev. Inst. Sal.

Enf. Trop. 14: 25-32.VARGAS, L. (1960): The subgenera of Culicoides of the Americas (Diptera, Ceratopogonidae). - Rev.

Biol. Trop. 8: 35-47.


VARGAS, L. & W.W. WIRTH (1955): Culicoides blantoni n. sp. (Diptera, Heleidae). - Rev. Inst. Sal. Enf.Trop. 15: 33-35.

VILLALOBOS, C. DE & M. RONDEROS (2003): Dasyhelea necrophila SPINELLI et RODRÍGUEZ, 1999 (Diptera:Ceratopogonidae) a new potential paratenic host of Paragordius varius (LEIDY, 1851) (Gordiida,Nematomorpha). - Acta Parasitol. 48: 218-221.

VITALE, G.C., WIRTH, W.W. & T.H.G. AITKEN (1981): New species and records of Culicoides rearedfrom arboreal habitats in Panama, with a synopsis of the debilipalpis group (Diptera:Ceratopogonidae). - Proc. Entomol. Soc. Washington 83: 140-159.

WALKER, F. (1856): Vol. 1. Diptera. - In: Insecta Saundersiana: or characters of undescribed insects inthe collection of William Wilson SAUNDERS: 415-474, Esq., F.R.S., F.L.S. & C. Van Voorst,London: 474 pp.

WALTL, J. (1837): Neue Gattungen von Mücken bei Passau. - Isis (Oken’s) 1837: 279-283.WARMKE, H.E. (1951): Studies on pollination of Hevea brasiliensis in Puerto Rico. - Science 113: 646-648.WARMKE, H.E. (1952): Studies on natural pollination of Hevea brasiliensis in Brazil. - Science 116: 474-475.WATTS, D., PHILLIPS, M.I., CALLAHAN, D.J., GRIEBENOW, W., HYAMS, K.C. & C.G. HAYESL (1997): Oropouche

virus transmission in the Amazon river basin of Peru. - Am. J. Trop. Med. Hyg. 56: 148-152.WAUGH, W.T. & W.W. WIRTH (1976): A revision of the genus Dasyhelea KIEFFER of the eastern United

States north of Florida (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 69: 219-247.WESENBERG-LUND, C. (1914): Bidrag til nogle myggeslaegters, saerlig Mochlonyx og Corethra biologi. -

Mindeskrift i Anledning of Hundredaaret for Japetus Steenstrups Fodsel, no. 34: 25 pp.WESTWOOD, J.O. (1840): Order XIII. Diptera Aristotle (Antliata FABRICIUS. Halteriptera Clairv.). An

introduction to the modern classification of insects; founded on the natural habits andcorresponding organisation of the different families: 125-154. - Synopsis of the genera ofBritish insects, London: 158 pp.

WEYENBERGH, H. (1883): Die Gattung Didymophleps m. - Stett. Entomol. Zeit. 1883: 108-111.WILLIAMS, F.X. (1944): Biological studies in Hawaiian water-loving insects. Part III. Diptera or flies. D.

Culicidae, Chironomidae and Ceratopogonidae. - Proc. Hawaiian Entomol. Soc. 12: 149-180.WILLIAMS, R.W. (1956): The biting midges of the genus Culicoides found in the Bermuda Islands

(Diptera, Heleidae) I. A description of C. bermudensis n. sp. with a key to the local fauna. - J.Parasitol. 42: 297-300.

WILLISTON, S.W. (1896): On the Diptera of St. Vincent (West Indies). - Trans. Entomol. Soc. London1896: 253-446, pls. 8-14.

WILLISTON, S.W. (1900): Supplement (Part). - In: GODMAN, F.D. & O. SALVIN (eds.): Biologia Centrali-Americana. Zoologia-Insects-Diptera: 217-248, Vol. 1. London: 378 pp., 6 pls.

WINDER, J.A. (1977a): Field observations on Ceratopogonidae and other Diptera: Nematoceraassociated with cocoa flowers in Brazil. - Bull. Entomol. Res. 67: 57-63.

WINDER, J.A. (1977b): Biology, activity, and behavior of cocoa pollinators (abstract). - MosquitoNews 37: 289.

WINNERTZ, J. (1852): Beitrag zur Kenntniss der Gattung Ceratopogon MEIGEN. - Linn. Entomol. 6: 1-80.WIRTH, W.W. (1951a): A new biting midge of the genus Leptoconops from Florida, with new records

of other American species (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 53: 281-284.WIRTH, W.W. (1951b): New species and records of Virginia Heleidae (Diptera). - Proc. Entomol. Soc.

Washington 53: 313-326.WIRTH, W.W. (1952a): The Heleidae of California. - Univ. Calif. Pub. Entomol. 9: 95-266.


WIRTH, W.W. (1952b): Los insectos de las Islas Juan Fernández. 7. Heleidae and Tendipedidae (Diptera).- Rev. Chil. Entomol. 2: 87-104.

WIRTH, W.W. (1952c): Two new species of anthropophilic Culicoides from Guatemala (Diptera:Heleidae). - J. Parasitol. 38: 245-247.

WIRTH, W.W. (1952d): The immature stages of two species of Florida salt marsh sand flies (Diptera,Heleidae). - Fla. Entomol. 35: 91-100.

WIRTH, W.W. (1953a): Biting midges of the heleid genus Stilobezzia in North America. - Proc. U.S.Nat. Mus. 103: 57-85.

WIRTH, W.W. (1953b): American biting midges of the heleid genus Monohelea. - Proc. U.S. Nat. Mus.103: 135-154.

WIRTH, W.W. (1955): Three new species of Culicoides from Texas (Diptera, Heleidae). - J. WashingtonAcad. Sci. 45: 355-359.

WIRTH, W.W. (1956a): New species and records of biting midges ectoparasitic on insects (Diptera,Heleidae). - Ann. Entomol. Soc. Am. 49: 356-364.

WIRTH, W.W. (1956b): The biting midges ectoparasitic on blister beetles (Diptera, Heleidae). - Proc.Entomol. Soc. Washington 58: 15-23.

WIRTH, W.W. (1956c): The heleid midges involved in the pollination of rubber trees in America(Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 241-250.

WIRTH, W.W. (1959a): Pachyhelea, a new genus of American Ceratopogonidae related to Palpomyia(Diptera). - Bull. Brooklyn Entomol. Soc. 54: 50-52.

WIRTH, W.W. (1959b): New species and records of Heleidae from Brazil (Diptera). - Deut. Entomol.Zeitsch. 6: 234-237.

WIRTH, W.W. (1960): The genus Pellucidomyia MACFIE (Diptera, Ceratopogonidae). - Bull. BrooklynEntomol. Soc. 55: 1-3.

WIRTH, W.W. (1962): A reclassification of the Palpomyia-Bezzia-Macropeza groups, and a revision of theNorth American Sphaeromiini (Diptera, Ceratopogonidae). - Ann. Entomol. Soc. Am. 55: 272-287.

WIRTH, W.W. (1965a): A new Johannsenomyia from Brazil (Diptera: Ceratopogonidae). - Proc. Entomol.Soc. Washington 67: 4.

WIRTH, W.W. (1965b): A revision of the genus Parabezzia MALLOCH (Diptera, Ceratopogonidae). -Proc. Entomol. Soc. Washington 67: 215-230.

WIRTH, W.W. (1965c): Two new species of Macrurohelea from Chile (Diptera, Ceratopogonidae). -Pan-Pac. Entomol. 41: 46-50.

WIRTH, W.W. (1966): A new Jamaican blood-sucking midge from lacewings (Diptera, Ceratopogonidae).- Proc. Entomol. Soc. Washington 68: 29-32.

WIRTH, W.W. (1969): New species and records of Galapagos Diptera. - Proc. Calif. Acad. Sci. 36: 571-594.WIRTH, W.W. (1970): The neotropical Forcipomyia midges of the subgenus Thyridomyia SAUNDERS

(Diptera, Ceratopogonidae). - Studia Entomol. 13: 429-440.WIRTH, W.W. (1971): A review of the “stick-ticks”, neotropical biting midges of the Forcipomyia

subgenus Microhelea parasitic on walking stick insects (Diptera: Ceratopogonidae). - Entomol.News 82: 229-245.

WIRTH, W.W. (1972): The neotropical Forcipomyia (Microhelea) species related to the caterpillar parasiteF. fuliginosa (Diptera: Ceratopogonidae). - Ann. Entomol. Soc. Am. 65: 564-577.

WIRTH, W.W. (1974): Family Ceratopogonidae. - In: A catalog of the diptera of the Americas southof the United States, Fasc. 14: 89 pp.


WIRTH, W.W. (1976): Forcipomyia pictoni MACFIE and descriptions of two new related species fromFlorida (Diptera: Ceratopogonidae). - Fla. Entomol. 59: 77-84.

WIRTH, W.W. (1977): A review of the pathogens and parasites of the biting midges (Diptera:Ceratopogonidae). - J. Washington Acad. Sci. 67:60-75.

WIRTH, W.W. (1978): New species and records of intertidal biting midges of the genus DasyheleaKIEFFER from the Gulf of California (Diptera: Ceratopogonidae). - Pacific Insects 18: 191-198.

WIRTH, W.W. (1982a): The cacao-pollinating midges of the Forcipomyia argenteola group (Diptera:Ceratopogonidae). - Proc. Entomol. Soc. Washington 84: 568-585.

WIRTH, W.W. (1982b): New species of neotropical Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol.65: 248-253.

WIRTH, W.W. (1983a): The North American predaceous midges of the Bezzia bicolor group (Diptera:Ceratopogonidae). - Fla. Entomol. 66: 292-310.

WIRTH, W.W. (1983b): A review of the American predaceous midges of the Bezzia nobilis group(Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 85:670-685.

WIRTH, W.W. (1991a): The predaceous midge genus Allohelea KIEFFER in the western hemisphere(Diptera: Ceratopogonidae). - Fla. Entomol. 74: 491-505.

WIRTH, W.W. (1991b): New and little-known species of Forcipomyia (Diptera: Ceratopogonidae)associated with cocoa pollination in Brazil. - Proc. Entomol. Soc. Washington 93: 163-175.

WIRTH, W.W. (1991c): Forcipomyia bicolor and related species of the subgenus Lepidohelea in Brazil(Diptera: Ceratopogonidae). - Fla. Entomol. 74: 506-517.

WIRTH, W.W. (1992): A new genus of Ceratopogonini (Diptera: Ceratopogonidae) from Brazil. -Proc. Entomol. Soc. Washington 94: 276-281.

WIRTH, W.W. (1994a): The subgenus Atrichopogon (Lophomyidium) with a revision of the Nearcticspecies (Diptera: Ceratopogonidae). - Insecta Mundi 8: 17-36.

WIRTH, W.W. (1994b): The western hemisphere species of the predaceous midge genus Echinohelea,with descriptions of six new species (Diptera: Ceratopogonidae). - Insecta Mundi 8: 227-242.

WIRTH, W.W. & W.R. ATCHLEY (1973): A review of the North American Leptoconops (Diptera:Ceratopogonidae). - Grad. Stud. Texas Tech Univ. 5: 1-57.

WIRTH, W.W. & P. BARRETO (1978): New species of Culicoides biting midges (Diptera: Ceratopogonidae)from Colombia. - J. Med. Entomol. 14: 553-564.

WIRTH, W.W. & F.S. BLANTON (1953a): Studies in Panama Culicoides (Diptera: Heleidae): I, Descriptionsof six new species. - J. Washington Acad. Sci. 43: 69-77.

WIRTH, W.W. & F.S. BLANTON (1953b): Studies in Panama Culicoides (Diptera, Heleidae). II. Descriptionsof six additional new species. - J. Parasitol. 39: 229-236.

WIRTH, W.W. & F.S. BLANTON (1953c): Studies in Panama Culicoides (Diptera, Heleidae). III. A newspecies related to phlebotomus (WILLISTON). - Entomol. News 64: 113-120.

WIRTH, W.W. & F.S. BLANTON (1955a): Studies in Panama Culicoides (Diptera, Heleidae) IV. Descriptionsof three new species. - Bull. Brooklyn Entomol. Soc. 50: 100-106 (October).

WIRTH, W.W. & F.S. BLANTON (1955b): Studies in Panama Culicoides (Diptera, Heleidae) V.Descriptions of three new species of the subgenus Oecacta POEY. - Bull. Brooklyn Entomol.Soc. 50: 121-127.

WIRTH, W.W. & F.S. BLANTON (1956a): A new species of salt-marsh sand fly from Florida, the Bahamas,Panama and Ecuador: its distribution and taxonomic differentiation from Culicoides furens (POEY)(Diptera, Heleidae). - Fla. Entomol. 39: 157-162.


WIRTH, W.W. & F.S. BLANTON. (1956b): Studies in Panama Culicoides VII. The species of the pulicarisand cova-garciai group (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 211-227.

WIRTH, W.W. & F.S. BLANTON (1956c): Studies in Panama Culicoides (Diptera, Heleidae). IX. Two newspecies related to leoni BARBOSA and reevesi WIRTH. - Bull. Brooklyn Entomol. Soc. 51: 45-52.

WIRTH, W.W. & F.S. BLANTON (1959): Biting midges of the genus Culicoides from Panama (Diptera:Heleidae). - Proc. U.S. Nat. Mus. 109: 237-482.

WIRTH, W.W. & F.S. BLANTON (1967): The North American Culicoides of the guttipennis group (Diptera:Ceratopogonidae). - Fla. Entomol. 50: 207-232.

WIRTH, W.W. & F.S. BLANTON (1968a): A revision of the neotropical biting midges of the hylas groupof Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol. 51: 201-215.

WIRTH, W.W. & F.S. BLANTON (1968b): A new Culicoides from Guyana (Diptera: Ceratopogonidae). -Fla. Entomol. 51: 251-252.

WIRTH, W.W. & F.S. BLANTON (1970a): New genera of Neotropical Ceratopogonidae (Diptera). - Fla.Entomol. 53: 7-14.

WIRTH, W.W. & F.S. BLANTON (1970b): New species of Neotropical Culicoides (Diptera:Ceratopogonidae). - Fla. Entomol. 53: 39-45.

WIRTH, W.W. & F.S. BLANTON (1970c): Notes on Brachypogon KIEFFER (Diptera, Ceratopogonidae), a newspecies, and two new Neotropical genera of the tribe Ceratopogonini. - Fla. Entomol. 53: 93-104.

WIRTH, W.W. & F.S. BLANTON (1970d): A review of the Culicoides nigrigenus group, with two newspecies (Diptera: Ceratopogonidae). - Entomol. News 81: 141-151.

WIRTH, W.W. & F.S. BLANTON (1971): New Neotropical sandflies of the Culicoides debilipalpis group(Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 34-43.

WIRTH, W.W. & F.S. BLANTON (1972a): Lanehelea, a new neotropical genus of Sphaeromiini (Diptera:Ceratopogonidae). - Studia Entomol. 15: 433-438.

WIRTH, W.W. & F.S. BLANTON (1972b): A new Patagonian biting midge of the the genus Monohelea,Isthmohelea (Diptera: Ceratopogonidae). - Fla. Entomol. 55: 173-176.

WIRTH, W.W. & F.S. BLANTON (1973): A review of the maruins or biting midges of the genus Culicoides(Diptera: Ceratopogonidae) in the Amazon Basin. - Amazoniana 4: 405-470.

WIRTH, W.W. & F.S. BLANTON (1974a): A new Florida sand fly closely related to Culicoides haematopotusMALLOCH (Diptera: Ceratopogonidae). - Fla. Entomol. 57: 23-26.

WIRTH, W.W. & F.S. BLANTON (1974b): The West Indian sandflies of the genus Culicoides (Diptera:Ceratopogonidae). - Agri. Res. Serv., U.S. Dept. Agric., Tech. Bull. 1474: iv + 98 pp.

WIRTH, W.W. & F.S. BLANTON (1978): Two new species of neotropical Culicoides (Diptera:Ceratopogonidae). - Pan-Pac. Entomol. 54: 236-240.

WIRTH, W.W. & J.L. CASTNER (1990): New neotropical species of “stick-tick” (Diptera:Ceratopogonidae) from katydids. - Fla. Entomol. 73: 157-160.

WIRTH, W.W. & M.I. DOW (1971): Studies on the genus Forcipomyia III. Blantonia, a new subgenus inthe Trichohelea complex (Diptera: Ceratopogonidae). - Fla. Entomol. 54: 289-295.

WIRTH, W.W. & M.I. DOW (1972): Studies on the genus Forcipomyia 4. Rhynchoforcipomyia, a newneotropical subgenus in the Trichohelea complex (Diptera: Ceratopogonidae). - Ann. Entomol.Soc. Am. 65: 862-872.

WIRTH, W.W. & A.L. DYCE (1985): The current taxonomic status of the Culicoides vectors of bluetongueviruses. - In: BARBER, T.L. & M.M. JOCHIM (eds.): 151-162. Bluetongue and related orboviruses.Alan R. Liss, New York: 746 pp.


WIRTH, W.W. & M.L. FELIPPE-BAUER (1989): The neotropical biting midges related to Culicoides paraensis(Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 84:551-565.

WIRTH, W.W. & W.L. GROGAN (1977): Taxonomic notes on the genus Heteromyia SAY, and a newspecies from Nicaragua (Diptera: Ceratopogonidae). - Fla. Entomol. 60: 177-185.

WIRTH, W.W. & W.L. GROGAN (1981): Natural History of Plummer’s Island, Maryland XXV. Bitingmidges (Diptera: Ceratopogonidae). 3. The species of the tribe Stilobezziini. - Bull. Biol. Soc.Washington 5: 1-102.

WIRTH, W.W. & W.L. GROGAN (1982): The predaceous midges of the genus Phaenobezzia in NorthAmerica (Diptera: Ceratopogonidae). - Mem. Entomol. Soc. Washington 10:179-192.

WIRTH, W.W. & W.L. GROGAN (1983): The Nearctic species of the Bezzia bivittata group (Diptera:Ceratopogonidae). - Proc. Biol. Soc. Washington 96: 489-523.

WIRTH, W.W. & W.L. GROGAN (1988): The predaceous midges of the world (Diptera: Ceratopogonidae;Tribe Ceratopogonini). - Flora and Fauna Handbook 4. E.J. Brill, Leiden: xv + 160 pp.

WIRTH, W.W. & A.A. HUBERT (1960): Ceratopogonidae (Diptera) reared from cacti, with a review ofthe copiosus group of Culicoides. - Ann. Entomol. Soc. Am. 53: 639-658.

WIRTH, W.W. & V.H. LEE (1967): New species of Culicoides from high altitudes in the ColombianAndes (Diptera: Ceratopogonidae). - Proc. U.S. Nat. Mus. 124: 1-22.

WIRTH, W.W. & D.H. MESSERSMITH (1971): Studies on the genus Forcipomyia. 1. The North Americanparasitic midges of the subgenus Trichohelea (Diptera: Ceratopogonidae). - Ann. Entomol. Soc.Am. 64: 15-26.

WIRTH, W.W. & B.A. MULLENS (1992): Culicoides boydi (Diptera: Ceratopogonidae): a potential vectorof hemorrhagic disease viruses to desert bighorn sheep in southern California. - J. Med. Entomol.29: 1006-1010.

WIRTH, W.W. & N.C. RATANAWORABHAN (1971a): Ceratoculicoides, a new genus related to CeratopogonMEIGEN (Diptera, Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 170-177.

WIRTH, W.W. & N.C. RATANAWORABHAN (1971b): Notes on Neotropical Pellucidomyia (Diptera,Ceratopogonidae). - Ann. Entomol. Soc. Am. 64: 446-448.

WIRTH, W.W. & N.C. RATANAWORABHAN (1972a): Neobezzia, a new neotropical biting midge genus ofthe tribe Sphaeromiini (Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 45: 476-490.

WIRTH, W.W. & N.C. RATANAWORABHAN (1972b): A revision of the tribe Stenoxenini (Diptera:Ceratopogonidae). - Ann. Entomol. Soc. Am. 65: 1368-1388.

WIRTH, W.W. & N.C. RATANAWORABHAN (1978): Studies on the genus Forcipomyia. V. Key to subgeneraand description of a new subgenus related to Euprojoannisia BRÈTHES (Diptera: Ceratopogonidae).- Proc. Entomol. Soc. Washington 80: 493-507.

WIRTH, W.W. & S.J. SORIA (1975): A new Neotropical Forcipomyia midge closely related to F. (F.)genualis (LOEW) (Diptera: Ceratopogonidae). - Rev. Theobroma 5: 19-27.

WIRTH, W.W. & S.J. SORIA (1980): Studies on the genus Forcipomyia VI. The Neotropical species of thesubgenus Warmkea (Diptera: Ceratopogonidae). - Rev. Theobroma 9: 137-161.

WIRTH, W.W. & S.J. SORIA (1981): Two Culicoides biting midges reared from inflorescences of Calatheain Brazil and Colombia, and a key to the species of the discrepans groups (Diptera:Ceratopogonidae). - Rev. Theobroma 11: 107-117.

WIRTH, W.W. & G.R. SPINELLI (1992a): Immature stages of Forcipomyia seminole WIRTH and a relatednew neotropical species (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 349-356.


WIRTH, W.W. & G.R. SPINELLI (1992b): American predaceous midges of the subgenus Eukraioheleaof Stilobezzia (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 342-349.

WIRTH, W.W. & G.R. SPINELLI (1993a): The American species of the annulatipes group of the subgenusLepidohelea, genus Forcipomyia (Diptera: Ceratopogonidae). - Insecta Mundi 6: 109-125 (1992).

WIRTH, W.W. & G.R. SPINELLI (1993b): The North American species of the Forcipomyia (Lepidohelea)bicolor subgroup (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 95: 611-634.

WIRTH, W.W. & W.T. WAUGH (1976): Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae)associated with culture of Cocoa. - Studia Entomol. 19: 223-236.

WIRTH, W.W. & R.W. WILLIAMS (1957): The biting midges of the Bermuda Islands, with descriptionsof five new species. - Proc. Entomol. Soc. Washington 59: 5-14.

WIRTH, W.W., DYCE, A.L. & G.R. SPINELLI (1988): An atlas of wing photographs, with a summary ofthe numerical characters of the Neotropical species of Culicoides (Diptera: Ceratopogonidae). -Contrib. Am. Entomol. Inst. 25: 1-72.

YOUNG, A.M. (1982): Effects of shade cover and availability of midge breeding sites on pollinatingmidge populations and fruit set in two cocoa farms. - J. Appl. Ecol. 19: 47-63.

YOUNG, A.M. (1983): Seasonal differences in abundance and distribution of cocoa-pollinating midgesin relation to flowering and fruit set between shaded and sunny habitats of the La Lola cocoafarm in Costa Rica. - J. Appl. Ecol. 20: 801-831.

YOUNG, A.M. (1986a): Cocoa Pollination. - Cocoa Grower’s Bull. 37: 5-23.YOUNG, A.M. (1986b): Notes on the distribution and abundance of midges (Diptera: Ceratopogonidae

and Cecidomyiidae) in some Central American cacao plantations. - Brenesia 24: 273-286.


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

10SUBJECT INDEX *

aliciae HUERTA & BORKENT (Ceratoculicoides) 80alleni CLASTRIER & WIRTH (Forcipomyia) 54Allohelea KIEFFER 32, 76, 101, 115-116, 139,

141, 148Alluaudomyia KIEFFER 32, 76, 101, 115-116, 129,

140-141, 146, 148almeidai (LANE) (Palpomyia) 95almirantei WIRTH & BLANTON (Culicoides) 74altivolans MACFIE (Atrichopogon) 44alvarezi ORTÍZ (Culicoides) 75amapaensis LANE (Paryphoconus) 97amazonica CLASTRIER (Stilobezzia) 85amazonica SPINELLI & WIRTH (Alluaudomyia) 76amazonica WIRTH & BLANTON (Fittkauhelea) 81,

104, 150amazonica WIRTH (Forcipomyia) 54amazonius MACFIE (Culicoides) 69ameliae BROWNE (Culicoides) 62americana KIEFFER (Stilobezzia) 86Amerohelea GROGAN & WIRTH 32, 41, 92, 102, 114,

122, 128, 143, 145, 148amnicola (MACFIE) (Neobezzia) 30, 91amnigena (MACFIE) (Stilobezzia) 86Amossovia GLUKHOVA 62ancora (COQUILLETT) (Dasyhelea) 59andensis INGRAM & MACFIE (Dasyhelea) 59andicola WIRTH & LEE (Culicoides) 63andinus WIRTH & LEE (Culicoides) 73angustipennis ENDERLEIN (Paryphoconus) 31, 97, 113,

127, 131-132Anilomyia VARGAS 40, 62anitae HUERTA & IBAÑEZ-BERNAL (Forcipomyia) 52

* Figures in bold


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

54, 101, 103, 105, 109, 111, 129, 134, 136,147-148, 150

atrichopogon LANE & FORATTINI (Stilobezzia) 85atripalpis WIRTH & BLANTON (Culicoides) 71attenuata SAUNDERS (Forcipomyia) 26, 52aureus ORTÍZ (Culicoides) 74auricoma KIEFFER (Atrichopogon) 44australis WIRTH (Dasyhelea) 59

Austrohelea WIRTH & GROGAN 32, 40, 77, 101,116-117, 140, 142, 148

austroparaensis SPINELLI (Culicoides) 66Austrosphaeromias SPINELLI 40, 90, 102, 122-123,

143-144, 149Avaritia FOX 63avilaensis ORTÍZ & MIRSA (Culicoides) 70azureus WIRTH & BLANTON (Culicoides) 70bachmanni SPINELLI (Culicoides) 66Baeodasymyia CLASTRIER & RACCURT 32, 40-41, 77,

101, 107, 138, 149Baeohelea WIRTH & BLANTON 40-41, 77, 101, 107,

138, 149bahamensis (JOHNSON) (Dasyhelea) 27, 59Bahiahelea WIRTH 40, 78, 101, 110, 115, 138-139bahiensis BARBOSA (Culicoides) 64bahiensis WIRTH & SPINELLI (Forcipomyia) 52bajensis WIRTH (Dasyhelea) 59balboa (LANE & WIRTH) (Downeshelea) 80balsapambensis ORTÍZ & LEÓN (Culicoides) 72balseiroi SPINELLI & GROGAN (Parabezzia) 30, 84balseiroi SPINELLI (Atrichopogon) 44bambusicola LUTZ (Culicoides) 27, 64banksi (GERRY) (Nilobezzia) 92barbatus BORKENT & PICADO (Atrichopogon) 44barbosai WIRTH & BLANTON (Culicoides) 17, 27, 70barrettoi LANE & DURET (Clinohelea) 88barrettoi LANE (Palpomyia) 95barrettoi LANE (Paryphoconus) 97barthi TAVARES & SOUZA (Culicoides) 70basifemoralis WIRTH & SPINELLI (Forcipomyia) 52bassoi RONDEROS & SPINELLI (Leptoconops) 43batesi LANE (Paryphoconus) 98batesi WIRTH & BLANTON (Culicoides) 67baueri WIRTH (Forcipomyia) 58bayano WIRTH (Culicoides) 27, 66beaveri WIRTH & BARRETO (Culicoides) 71beccus BORKENT & PICADO (Atrichopogon) 45beckae WIRTH (Stilobezzia) 86beebei FOX (Culicoides) 62bejaranoi DURET & LANE (Heteromyia) 89belemensis CLASTRIER & WIRTH (Forcipomyia) 54belemensis WIRTH & BLANTON (Culicoides) 71bella (COQUILLETT) (Alluaudomyia) 29, 76, 112, 118


Beltranmyia VARGAS 63benarrochi ORTÍZ & MIRSA (Culicoides) 73bequaerti (KIEFFER) (Leptoconops) 17, 25, 33, 43,

109, 130, 151bermudae WIRTH & WILLIAMS (Dasyhelea) 60bermudensis WILLIAMS (Culicoides) 27, 63Bezzia KIEFFER 32, 86, 91-93, 97-98, 102, 122-

123, 128, 133, 143-145, 149bicellii (LANE) (Mallochohelea) 91bicinctipes INGRAM & MACFIE (Stilobezzia) 85bicolor LANE (Stilobezzia) 30, 86bicolor LUTZ (Forcipomyia) 52, 59bicolor SAUNDERS (Forcipomyia) 26, 58-59bicornis FELIPPE-BAUER & QUINTELAS (Downeshelea) 80bicuspis BORKENT & PICADO (Atrichopogon) 45bidentata FELIPPE-BAUER & SPINELLI (Monohelea) 83biestroi SPINELLI & FELIPPE-BAUER (Sphaerohelea)

92, 152biestroi SPINELLI & RONDEROS (Culicoides) 67biestroi SPINELLI (Alluaudomyia) 29, 76bifida WIRTH & SPINELLI (Forcipomyia) 52bifidus EWEN (Atrichopogon) 25, 45bifidus SPINELLI & GROGAN (Brachypogon) 78bimaculata (LOEW) (Clinohelea) 113, 121bimaculata LANE & FORATTINI (Stilobezzia) 86bimaculatus FLOCH & ABONNENC (Culicoides) 68bimaculatus SPINELLI & GROGAN (Brachypogon) 78birabeni CAVALIERI (Culicoides) 70bispinosa KIEFFER (Stilobezzia) 86bivittata (COQUILLETT) (Bezzia) 30, 93blantoni (LANE & WIRTH ) (Johannsenomyia) 90blantoni (LANE & WIRTH) (Downeshelea) 80blantoni (LANE) (Pellucidomyia) 89blantoni LANE & FORATTINI (Stilobezzia) 86blantoni SORIA & BYSTRAK (Forcipomyia) 25, 49blantoni SPINELLI & WIRTH (Bezzia) 30, 93blantoni VARGAS & WIRTH (Culicoides) 64blantoni WIRTH & RATANAWORABHAN (Neobezzia)

91, 114blantoni WIRTH & RATANAWORABHAN (Stenoxenus) 99blantoni WIRTH (Echinohelea) 81blantoni WIRTH (Parabezzia) 84Blantonia WIRTH & DOW 48boliviensis (KIEFFER) (Palpomyia) 96

boliviensis KIEFFER (Palpomyia) 95-96boliviensis SPINELLI & WIRTH (Culicoides) 73bonaerensis SPINELLI (Brachypogon) 78borgmeieri WIRTH & WAUGH (Dasyhelea) 60borinqueni FOX & HOFFMAN (Culicoides) 28, 65Borkenthelea SPINELLI & GROGAN 40, 78, 101, 116-

117, 140, 142, 149borkenti SPINELLI & CAZORLA (Brachypogon) 79bourioni (CLASTRIER) (Nannohelea) 83, 108Brachyconops WIRTH & ATCHLEY 43Brachypogon KIEFFER 32, 78, 101, 107, 115-116,

138-139, 141, 149, 152brachyrhyncha WIRTH & DOW (Forcipomyia) 56brasiliae (LANE) (Neobezzia) 91brasiliana CLASTRIER & WIRTH (Forcipomyia) 54brasilianum FORATTINI (Culicoides) 68brasiliensis (LANE) (Nilobezzia) 92brasiliensis (LUTZ) (Leptoconops) 43-44brasiliensis LANE (Monohelea) 83brasiliensis MACFIE (Atrichopogon) 45brasiliensis MACFIE (Forcipomyia) 26, 52brasiliensis MACFIE (Palpomyia) 95brasiliensis MACFIE (Stenoxenus) 99brasiliensis SPINELLI & GROGAN (Parabezzia) 84brasiliensis WIRTH (Bahiahelea) 78, 113, 125bredini WIRTH & BLANTON (Culicoides) 65breelandi CLASTRIER & WIRTH (Forcipomyia) 54brevicornis (KIEFFER) (Bezzia) 30, 93brevilabellata CLASTRIER & WIRTH (Forcipomyia) 54brevipalpis (INGRAM & MACFIE) (Paradasyhelea) 75brevipalpis MACFIE (Atrichopogon) 45bricenoi ORTÍZ (Culicoides) 70broadheadi CLASTRIER & WIRTH (Forcipomyia) 54bromeliae SAUNDERS (Forcipomyia) 25, 49bromeliae SPINELLI & WIRTH (Bezzia) 30, 93bromelicola (LUTZ) (Forcipomyia) 26, 55-56brownei SPINELLI (Culicoides) 68brunnea WIRTH (Parabezzia) 84brunneipennis SPINELLI & WIRTH (Paryphoconus) 98bystraki GROGAN & WIRTH (Parabezzia) 109Cacaohelea WIRTH & GROGAN 40-41, 79, 101, 115,

117, 126, 140-141, 149cacaoi WIRTH & WAUGH (Dasyhelea) 60cacaophila RONDEROS & SPINELLI (Forcipomyia) 52


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

catarinensis MARINO & SPINELLI (Forcipomyia) 51catarinensis SPINELLI & WIRTH (Alluaudomyia) 76catarinensis SPINELLI & WIRTH (Bezzia) 93caucaensis WIRTH & LEE (Culicoides) 72caudata INGRAM & MACFIE (Macrurohelea) 82caudispina (Dipetalonema) 33cayoensis SPINELLI & GROGAN (Parabezzia) 84cayoensis SPINELLI & WIRTH (Bezzia) 93cebacoi (LANE & WIRTH) (Downeshelea) 80Centrorhynchus LUTZ 51-52, 61, 64Ceratobezzia KIEFFER 41, 88, 98Ceratoculicoides WIRTH & RATANAWORABHAN 41,

101, 107, 109, 138, 149Ceratopogon MEIGEN 27, 44, 46-47, 49-51, 54,

56, 59-62, 67-70, 75-76, 79, 83, 86-87, 92-98, 118

cerifera SAUNDERS (Forcipomyia) 26, 53cervicalis (Onchocerca) 33chacoensis SPINELLI & WIRTH (Culicoides) 65chaconi MACFIE (Stilobezzia) 30, 86chaquensis DURET & LANE (Heteromyia) 89charrua FELIPPE-BAUER & SPINELLI (Downeshelea) 80charruus SPINELLI & Martinez (Culicoides) 28, 68chaverrii SPINELLI & BORKENT (Culicoides) 28, 62chiapasi (LANE & WIRTH) (Downeshelea) 80chilensis (PHILIPPI) (Forcipomyia) 51chilensis FORATTINI (Leptoconops) 44, 151chilensis GROGAN & WIRTH (Notiohelea) 83chilensis INGRAM & MACFIE (Atrichopogon) 46chilensis INGRAM & MACFIE (Dasyhelea) 60chilensis INGRAM & MACFIE (Palpomyia) 96chilensis SPINELLI & RONDEROS (Bezzia) 93chirusi (LANE & WIRTH) (Downeshelea) 80christopheri BORKENT (Baeodasymyia) 29, 77chrysonotus WIRTH & BLANTON (Culicoides) 62cincta (COQUILLETT) (Dasyhelea) 27, 59, 60claripennis LYNCH ARRIBÁLZAGA (Ceratopogon) 99clastrieri DESSART (Forcipomyia) 55clastrieri GROGAN & WIRTH (Nannohelea) 83, 151clastrieri SPINELLI & GROGAN (Parabezzia) 84Clastrieromyia SPINELLI & GROGAN 32, 41, 95, 102,

122, 128, 143, 145, 149clavata WILLISTON (Heteromyia) 30, 89clavipennis SPINELLI & WIRTH (Bezzia) 93


clavipes WIRTH & RATANAWORABHAN (Neobezzia) 91Clinohelea KIEFFER 32, 41, 88, 101, 117, 119,

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

Culicoides LATREILLE 17-18, 22, 24, 32-34, 40, 42-43, 60-65, 67, 69, 100-101, 103, 105, 133-134, 137, 147, 149, 150

cummingi SPINELLI & BORKENT (Culicoides) 71cunasi LANE & WIRTH (Monohelea) 83cylindrica WIRTH & DOW (Forcipomyia) 56cylindricornis WIRTH & BLANTON (Culicoides) 74daedaloides WIRTH & BLANTON (Culicoides) 71daedalus MACFIE (Culicoides) 71dalcyi GROGAN & WIRTH (Amerohelea) 92dalessandroi WIRTH & BARRETO (Culicoides) 70damascenoi LANE & DURET (Clinohelea) 88danaisi (FLOCH & ABONNENC) (Forcipomyia) 58darlingtonae WIRTH & BLANTON (Culicoides) 66darwini MARINO & SPINELLI (Forcipomyia) 53Dasyhelea KIEFFER 15, 22, 24, 32, 40, 59, 75, 101,

103, 105, 129, 133-134, 137, 146-147, 150dasyophrus MACFIE (Culicoides) 72davidi SPINELLI (Culicoides) 68daviesi WIRTH & BLANTON (Culicoides) 70deanei FELIPPE-BAUER & QUINTELAS (Downeshelea) 80deanei FELIPPE-BAUER & WIRTH (Culicoides) 75debilipalpis LUTZ (Culicoides) 17, 28, 66decor (WILLISTON) (Culicoides) 62delpontei CAVALIERI & CHIOSSONE (Atrichopogon) 45delpontei CAVALIERI (Forcipomyia) 50denisae CLASTRIER (Culicoides) 66dentifemur SPINELLI & WIRTH (Bezzia) 93depilis MACFIE (Atrichopogon) 45desutterae CLASTRIER & WIRTH (Forcipomyia) 54diabolicus HOFFMAN (Culicoides) 68Diaphanobezzia INGRAM & MACFIE 40, 80, 101,

110, 139, 150Dicrohelea KIEFFER 90dicrourus WIRTH & BLANTON (Culicoides) 28, 70Didymophleps WEYENBERGH 99, 102didymothecae MACFIE (Atrichopogon) 45diffusus SPINELLI (Culicoides) 68diminuta LANE & FORATTINI (Stilobezzia) 87diminutus BARBOSA (Culicoides) 69dimorphus KIEFFER (Stenoxenus) 99Diphaomyia VARGAS 64discoloripes MACFIE (Forcipomyia) 52discrepans ORTÍZ & MIRSA (Culicoides) 70


disjuncta INGRAM & MACFIE (Isthmohelea) 82, 126, 151distispinulosa SPINELLI & WIRTH (Alluaudomyia) 29, 76diversus FELIPPE-BAUER (Culicoides) 66dominicana DE MEILLON & WIRTH (Forcipomyia) 26, 56dominicanus WIRTH & BLANTON (Culicoides) 62dominicii ORTÍZ (Culicoides) 71domizii SPINELLI (Atrichopogon) 45donajii VARGAS (Culicoides) 72donskoffi CLASTRIER & WIRTH (Forcipomyia) 54dorsalis WIRTH & DOW (Forcipomyia) 56dorsofasciata (LUTZ) (Stilobezzia) 30, 86dowi BYSTRAK & WIRTH (Forcipomyia) 49Downeshelea WIRTH & GROGAN 80, 101, 110, 113,

115-116, 118, 126, 138, 140-141, 150dryadum MACFIE (Stilobezzia) 87Drymodesmyia VARGAS 65duartei TAVARES & LUNA DIAS (Culicoides) 73dubia MACFIE (Forcipomyia) 52dubitans LANE (Stilobezzia) 30, 87dunklei CLASTRIER & WIRTH (Forcipomyia) 54dunni WIRTH & BLANTON (Culicoides) 71dureti LANE & FORATTINI (Stilobezzia) 87dureti RONDEROS & SPINELLI (Culicoides) 66dycei SPINELLI & GROGAN (Clastrieromyia) 31, 95,

127, 131eadsi WIRTH & BLANTON (Culicoides) 66echinodes MACFIE (Atrichopogon) 45Echinohelea MACFIE 32, 40, 81, 101, 104, 122, 127,

131, 142, 150Echinoideshelea WIRTH 81ecuadorensis SPINELLI & GROGAN (Brachypogon) 78ecuadorensis SPINELLI & WIRTH (Paryphoconus) 98edeni WIRTH & BLANTON (Culicoides) 28, 64edmistoni WIRTH & SPINELLI (Forcipomyia) 26, 52edwardsi (SAUNDERS) (Forcipomyia) 26, 50, 56edwardsi INGRAM & MACFIE (Forcipomyia) 50edwardsi INGRAM & MACFIE (Stilobezzia) 85edwardsiana WIRTH (Forcipomyia) 50efferus FOX (Culicoides) 62eldridgei WIRTH & BARRETO (Culicoides) 66elegantula (JOHANNSEN) (Stilobezzia) 86elutus MACFIE (Culicoides) 64endemicus SPINELLI & MARINO (Atrichopogon) 45enderleini LANE (Paryphoconus) 98

equatoriensis BARBOSA (Culicoides) 66eriophora (WILLISTON) (Forcipomyia) 54esmeralda LANE & FORATTINI (Stilobezzia) 87espinolai FELIPPE-BAUER & LOURENCO-DE-

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

115, 139-140, 150fittkaui SPINELLI & WIRTH (Alluaudomyia) 76fittkaui SPINELLI & WIRTH (Paryphoconus) 98


fittkaui WIRTH & BLANTON (Culicoides) 74fittkaui WIRTH & RATANAWORABHAN (Neobezzia) 91flava (WILLISTON) (Forcipomyia) 51flavicauda MACFIE (Dasyhelea) 27, 60flavicaudae MACFIE (Atrichopogon) 45flavidus (JOHANNSEN) (Paryphoconus) 31, 98flavifemoris MACFIE (Forcipomyia) 53flavifrons (GUÉRIN) (Dasyhelea) 132flavipes LUTZ (Atrichopogon) 45, 95flavivenulus COSTA LIMA (Culicoides) 68flinti SPINELLI & WIRTH (Bezzia) 93flinti SPINELLI & WIRTH (Paryphoconus) 98flinti WIRTH (Culicoides) 66flochabonnenci ORTÍZ & MIRSA (Culicoides) 72florenciae MESSERSMITH (Culicoides) 72floridensis BECK (Culicoides) 28, 75floridensis DOW & WIRTH (Forcipomyia) 57floridensis WIRTH (Leptoconops) 44, 104fluminensis FELIPPE-BAUER & QUINTELAS

(Downeshelea) 80fluminensis LANE (Bezzia) 93fluminensis LANE (Stilobezzia) 87flumineus MACFIE (Atrichopogon) 45fluvialis MACFIE (Culicoides) 72fluviatilis (LUTZ) (Culicoides) 75forattinii ORTÍZ (Culicoides) 74forattinii WIRTH & GROGAN (Schizonyxhelea) 30,

85, 112Forcipomyia MEIGEN 15, 17, 18, 19-21, 22, 24, 32-

33, 40, 47-49, 53, 55-58, 101-103, 105, 118,133, 134, 136, 147-148, 150

formosa (LOEW) (Nilobezzia) 135foxi ORTÍZ (Culicoides) 17, 68franklini CLASTRIER & WIRTH (Forcipomyia) 54franklini SPINELLI (Culicoides) 68freitasi WIRTH & BLANTON (Culicoides) 64frontispina (DOW & TURNER) (Amerohelea) 92, 127fuliginosa (MEIGEN) (Forcipomyia) 26, 53-54fulvus JOHANNSEN (Stenoxenus) 99fur (JOHNSON) (Forcipomyia) 56furcifera MACFIE (Forcipomyia) 48furens (POEY) (Culicoides) 17, 28, 33, 70, 150furva INGRAM & MACFIE (Stilobezzia) 85fusca (PHILIPPI) (Forcipomyia) 40, 50

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


granditibialis BORKENT & PICADO (Atrichopogon) 45griseola WIRTH (Dasyhelea) 60Groganhelea SPINELLI & DIPPOLITO 41, 90, 102-

103, 122, 128, 134, 143, 145, 150grogani SPINELLI & WIRTH (Bezzia) 94guadeloupensis DELÉCOLLE & RIEB (Dasyhelea) 60guadeloupensis FLOCH & ABONNENC (Culicoides) 72guaimiesi LANE & WIRTH (Monohelea) 83guamai WIRTH & BLANTON (Culicoides) 74guamai WIRTH & DOW (Forcipomyia) 57guarani LANE (Palpomyia) 31, 96guarani MARINO & SPINELLI (Forcipomyia) 50guarani RONDEROS & SPINELLI (Culicoides) 72guarani RONDEROS & SPINELLI (Forcipomyia) 52guarani SPINELLI (Alluaudomyia) 76guerrai WIRTH & BLANTON (Culicoides) 72guianae (MACFIE) (Stilobezzia) 85guianae (WIRTH) (Downeshelea) 80guianae MACFIE (Paryphoconus) 98guianensis MACFIE (Atrichopogon) 45gustavoi BORKENT (Baeodasymyia) 77guttatus (COQUILLETT) (Culicoides) 67-68gutturosa (Onchocerca ) 33guyana CLASTRIER & WIRTH (Forcipomyia) 54guyana CLASTRIER (Palpomyia) 96guyana CLASTRIER (Schizonyxhelea) 85guyanensis FLOCH & ABONNENC (Culicoides) 28, 74Haematomyidium GOELDI 66-67haematopotus MALLOCH (Culicoides) 28, 64, 125haitiensis CLASTRIER & RACCURT (Parabezzia) 84haitiensis DELÉCOLLE (Culicoides) 65harii SPINELLI & GROGAN (Borkenthelea) 78, 108harpa SPINELLI & BORKENT (Forcipomyia) 57harpegonata WIRTH & SORIA (Forcipomyia) 25, 50harrisi MACFIE (Atrichopogon) 45Hartomyia MALLOCH 85hatoensis UTMAR & WIRTH (Forcipomyia) 48hayesi MATTA (Culicoides) 62heliconiae FOX & HOFFMAN (Culicoides) 68herediae WIRTH & SPINELLI (Forcipomyia) 26, 53hermani SPINELLI & BORKENT (Culicoides) 63hermosa UTMAR & WIRTH (Forcipomyia) 48hertigi WIRTH & BLANTON (Culicoides) 65Heteromyia SAY 32, 89, 101, 117, 142, 151-152

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


iquitosensis CLASTRIER & WIRTH (Forcipomyia) 54iriartei FOX (Culicoides) 64irwini GROGAN & WIRTH (Macrurohelea) 82irwini SPINELLI & WIRTH (Culicoides) 64Isoecacta GARRETT 76Isohelea KIEFFER 79Isthmohelea INGRAM & MACFIE 40, 82, 101, 115,

117, 140, 142, 151ixodoides (FIEBRIG-GERTZ) (Forcipomyia) 18, 53-54jamaicensis EDWARDS (Culicoides) 65jamaicensis SPINELLI & WIRTH (Bezzia) 94jamaicensis SPINELLI & WIRTH (Dasyhelea) 60jamaicensis WIRTH (Echinohelea) 81jamaicensis WIRTH (Forcipomyia) 58jamaicensis WIRTH (Parabezzia) 84jipajapae WIRTH (Forcipomyia) 58jocosa SAUNDERS (Forcipomyia) 26, 56johannseni (LANE) (Palpomyia) 96johannseni (WIRTH) (Allohelea) 112Johannsenomyia MALLOCH 32, 90-91, 95, 102, 122-

123, 129, 143-144, 146, 151johnsoni COQUILLETT (Stenoxenus) 31, 99jubata SPINELLI & WIRTH (Bezzia) 94jurbergi FELIPPE-BAUER (Culicoides) 64kampa FELIPPE-BAUER (Culicoides) 67kawensis CLASTRIER & WIRTH (Forcipomyia) 54keilini (SAUNDERS) (Forcipomyia) 26, 56kiefferi LANE (Heteromyia) 89kiefferi LANE (Paryphoconus) 98kiefferi LANE (Stilobezzia) 87kintzi WIRTH & BLANTON (Culicoides) 74knowltoni BECK (Culicoides) 63knowltoni CLASTRIER & WIRTH (Leptoconops) 43koenigi DELÉCOLLE & RIEB (Dasyhelea) 60kremeri SPINELLI & GROGAN (Clastrieromyia) 95kuanoskeles MACFIE (Forcipomyia) 53kuscheli WIRTH & BLANTON (Culicoides) 74kuscheli WIRTH (Macrurohelea) 82lacajae MACFIE (Atrichopogon) 46lacrimatorii MACFIE (Forcipomyia) 53lacustris INGRAM & MACFIE (Dasyhelea) 27, 60lacustris LANE (Palpomyia) 31, 96lacustris RONDEROS (Culicoides) 71lahillei (ICHES) (Culicoides) 17, 29, 33, 67

lamprogaster EDWARDS (Heteromyia) 89Lanehelea WIRTH & BLANTON 41, 90, 102-103, 122-

123, 134, 143-144, 151lanei ORTÍZ (Culicoides) 74lanei WIRTH & RATANAWORABHAN (Pellucidomyia) 89lanei WIRTH & RATANAWORABHAN (Stenoxenus) 99lanei WIRTH (Echinohelea) 29, 81, 112, 118lanei WIRTH (Johannsenomyia) 90lanei WIRTH (Monohelea) 83lanei WIRTH (Paryphoconus) 31, 98Lasiohelea KIEFFER 17, 20, 22, 26, 51-52, 58-59, 150latifemoris (INGRAM & MACFIE) (Pachyhelea) 95latipennis SPINELLI & WIRTH (Paryphoconus) 98leei SPINELLI & WIRTH (Alluaudomyia) 76leei SPINELLI & WIRTH (Bezzia) 94leei SPINELLI & WIRTH (Paryphoconus) 98leei WIRTH & BLANTON (Lanehelea) 90-91leei WIRTH & BLANTON (Parastilobezzia) 84, 113,

119, 152leei WIRTH & RATANAWORABHAN (Forcipomyia) 57leei WIRTH (Echinohelea) 81lenkoi LANE (Bezzia) 94lenti TAVARES & LUNA DIAS (Culicoides) 70leoni BARBOSA (Culicoides) 73leopoldoi ORTÍZ (Culicoides) 72Lepidohelea KIEFFER 26, 48, 52Leptoconops SKUSE 16-17, 22, 25, 32-33, 43-44, 101,

103, 111, 129, 134, 136, 146-147, 151Leptohelea WIRTH & BLANTON 40-41, 82, 101, 110,

138-139, 151lesliei WIRTH (Forcipomyia) 26, 58-59levis (COQUILLETT) (Atrichopogon) 44, 111, 118lichyi FLOCH & ABONNENC (Culicoides) 73limai BARRETTO (Culicoides) 73limitrofe SPINELLI & FELIPPE-BAUER (Mallochohelea) 91limnetis INGRAM & MACFIE (Forcipomyia) 58limnochares (MACFIE) (Palpomyia) 96limonensis ORTÍZ & LEÓN (Culicoides) 67limpidus WIRTH & RATANAWORABHAN (Stenoxenus) 99lineata (MEIGEN) (Palpomyia) 131lituratus (WILLISTON) (Atrichopogon) 46lobatoi FELIPPE-BAUER (Culicoides) 73lobatus BORKENT & PICADO (Atrichopogon) 46longicornis (WILLISTON) (Palpomyia) 97


longicornis EWEN (Atrichopogon) 25, 46longispina SAUNDERS (Forcipomyia) 25, 49lopesi BARRETTO (Culicoides) 73Lophomyidium CORDERO 47lota (WILLISTON) (Forcipomyia) 49loughnani EDWARDS (Culicoides) 65louriei (MACFIE) (Forcipomyia) 59lutea (MALLOCH) (Stilobezzia) 112, 119lutealaris WIRTH & BLANTON (Culicoides) 63luteigenua WIRTH & SPINELLI (Forcipomyia) 26, 53luteisquamosa WIRTH (Forcipomyia) 54luteogrisea WIRTH & WILLIAMS (Dasyhelea) 60luteovenus ROOT & HOFFMAN (Culicoides) 29, 64lutzi COSTA LIMA (Culicoides) 68lutzi LANE (Palpomyia) 96lydiae BORKENT (Baeodasymyia) 77lyrinotatus WIRTH & BLANTON (Culicoides) 74Macfiehelea LANE 89macfiei (COSTA LIMA) (Forcipomyia) 56macfiei LANE (Echinohelea) 81macfiei LANE (Paryphoconus) 98macfiei LANE (Stilobezzia) 87macfiei SPINELLI & GROGAN (Paradasyhelea) 75Macfiella FOX 40, 69macheti CLASTRIER & LEGRAND (Forcipomyia) 58macieli TAVARES & RUIZ (Culicoides) 29, 70macrostigma WIRTH & BLANTON (Culicoides) 74Macrurohelea INGRAM & MACFIE 40, 82, 101, 116,

130, 141, 151maculata LANE (Stilobezzia) 87maculata MACFIE (Dasyhelea) 60maculicrus INGRAM & MACFIE (Palpomyia) 96maculipennis (COQUILLETT) (Monohelea) 83maculipennis CLASTRIER (Atrichopogon) 46maculithorax (WILLISTON) (Culicoides) 70maculitibia LANE & FORATTINI (Stilobezzia) 86maculosa INGRAM & MACFIE (Forcipomyia) 53magna (SAUNDERS) (Forcipomyia) 26, 56magnipalpis WIRTH & BLANTON (Culicoides) 73magnus BORKENT & PICADO (Atrichopogon) 46, 95maia LANE & FORATTINI (Stilobezzia) 85malariologiensis PERRUOLO (Culicoides) 75Mallochohelea WIRTH 32, 91, 102, 122, 128-129,

143, 145-146, 151

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

112, 119, 126, 151microtoma (KIEFFER) (Forcipomyia) 54minasensis FELIPPE-BAUER (Culicoides) 65minisquamosa WIRTH (Forcipomyia) 54minitheca MARINO & SPINELLI (Forcipomyia) 50


minor (WIRTH) (Nilobezzia) 130mirsae ORTÍZ (Culicoides) 65misionensis SPINELLI (Brachypogon) 79misionensis SPINELLI (Paryphoconus) 98miyamotoi WIRTH & BLANTON (Culicoides) 74modesta CLASTRIER & RACCURT (Baeodasymyia) 77, 108modesta LANE (Stilobezzia) 87mojingaensis WIRTH & BLANTON (Culicoides) 69-70monicae SPINELLI & BORKENT (Culicoides) 63monicae SPINELLI & GROGAN (Brachypogon) 79monilicornis (COQUILLETT) (Forcipomyia) 57, 106monilicornis WIRTH & BLANTON (Rhynchohelea) 84-

85, 104, 106, 112, 152Monohelea KIEFFER 32, 46, 77, 80-82, 101, 110,

127, 138, 151monotheca SPINELLI & GROGAN (Macrurohelea) 82monticola INGRAM & MACFIE (Dasyhelea) 60monticola WIRTH & LEE (Culicoides) 73moorei CLASTRIER & WIRTH (Forcipomyia) 55morenoi MARINO & SPINELLI (Forcipomyia) 53morrisoni GROGAN & WIENERS (Dasyhelea) 60mortuifolii SAUNDERS (Forcipomyia) 25, 49multidentata RONDEROS & SPINELLI (Forcipomyia) 52multilineata (LUTZ) (Downeshelea) 80multipicta INGRAM & MACFIE (Forcipomyia) 50multispinosa SPINELLI & GROGAN (Palpomyia) 96musae CLASTRIER & DELÉCOLLE (Forcipomyia) 26, 56mutabilis (COQUILLETT) (Dasyhelea) 27, 59-60muzoni MARINO & SPINELLI (Forcipomyia) 50muzoni SPINELLI & DURET (Clinohelea) 88Mycterotypus NOÈ 43nana (MACFIE) (Forcipomyia) 58nana WIRTH & BLANTON (Baeohelea) 77-78, 108-

109, 112, 149Nannohelea GROGAN & WIRTH 83, 101, 107, 109,

138, 151nanus MACFIE (Atrichopogon) 46narthekophora MACFIE (Forcipomyia) 59navaiae BYSTRAK & WIRTH (Forcipomyia) 49nebulosus MACFIE (Atrichopogon) 46necrophila SPINELLI & RODRÍGUEZ (Dasyhelea) 24,

27, 60needhami THOMSEN (Alluaudomyia) 132neivai LANE (Clinohelea) 88

nelsoni GROGAN & WIRTH (Amerohelea) 92nemoralis (MACFIE) (Mallochohelea) 91Neobezzia WIRTH & RATANAWORABHAN 31, 41, 91,

102, 120, 122, 128, 143, 145, 151Neoceratopogon MALLOCH 76neoparaensis TAVARES & SOUZA (Culicoides) 67neopulicaris WIRTH (Culicoides) 29, 64neotropica (MACFIE) (Nilobezzia) 92neotropica CLASTRIER & WIRTH (Forcipomyia) 55neotropica WIRTH (Allohelea) 76, 118, 148neotropica WIRTH (Echinohelea) 81neotropicalis (LANE) (Paryphoconus) 98nerudai SPINELLI & GROGAN (Borkenthelea) 78neunguis GROGAN & WIRTH (Parabezzia) 84niger LANE (Stenoxenus) 99nigerrima INGRAM & MACFIE (Stilobezzia) 85nigra KIEFFER (Heteromyia) 89nigrescens MACFIE (Forcipomyia) 48nigrigenus WIRTH & BLANTON (Culicoides) 62-63nigrimaxillata CLASTRIER & WIRTH (Forcipomyia) 55nigripes (MACFIE) (Mallochohelea) 91nigripes MACFIE (Clinohelea) 30, 88nigripes MACFIE (Paryphoconus) 98nigritibialis SPINELLI & WIRTH (Bezzia) 94nigroflava LANE & FORATTINI (Stilobezzia) 87nigroflava LANE (Palpomyia) 96nigroscutellata LANE (Palpomyia) 96Nilobezzia KIEFFER 32, 91, 102, 122, 128, 143,

145, 151nobilis (WINNERTZ) (Bezzia) 31, 93-94, 114, 124, 130nodosa SAUNDERS (Forcipomyia) 26, 58nothofagus SPINELLI & GROGAN (Borkenthelea) 78, 112Notiohelea GROGAN & WIRTH 40, 83, 101, 103,

117, 134, 142, 151novaeteutoniae CLASTRIER & WIRTH (Forcipomyia) 55novaeteutoniae MACFIE (Atrichopogon) 46nubeculosa SPINELLI & WIRTH (Alluaudomyia) 76nubeculosus MACFIE (Atrichopogon) 46nubifer MACFIE (Paryphoconus) 98nuncupata MACFIE (Forcipomyia) 50obesa COSTA LIMA (Forcipomyia) 55obfuscatus INGRAM & MACFIE (Atrichopogon) 46obnoxius FOX (Culicoides) 74obnubilus INGRAM & MACFIE (Atrichopogon) 46


obscura LANE & FORATTINI (Stilobezzia) 87obscurus EWEN (Atrichopogon) 25, 46ocumare (ORTÍZ) (Atrichopogon) 46ocumarensis ORTÍZ (Culicoides) 69Oecacta POEY 70oedidactyla (INGRAM & MACFIE) (Physohelea) 90, 113oklahomensis KHALAF (Culicoides) 29, 62oligarthra SAUNDERS (Forcipomyia) 26, 56oliveirai (LANE) (Pellucidomyia) 89oliveirai LANE (Palpomyia) 96oliveirai LANE (Paryphoconus) 31, 98oliveri FOX & HOFFMAN (Culicoides) 69opilionivora (LANE) (Forcipomyia) 58orellana (ROBACK) (Heteromyia) 89orjuelai WIRTH & LEE (Culicoides) 63ornata LANE & FORATTINI (Stilobezzia) 85ornata WIRTH (Monohelea) 83ornaticrus INGRAM & MACFIE (Stilobezzia) 85ornatipennis CLASTRIER (Atrichopogon) 46ornatipennis MACFIE (Echinohelea) 81ornatipennis MACFIE (Forcipomyia) 51ortizi FOX (Culicoides) 71ozzardi (Dipetalonema) 33pabloi BROWNE (Culicoides) 72pachycnemus MACFIE (Atrichopogon) 46Pachyhelea WIRTH 32, 41, 95, 102, 123, 128, 144-

145, 152Pachyleptus WALKER 89pachymera (WILLISTON) (Pachyhelea) 31, 95, 124, 152pachymerus LUTZ (Culicoides) 74Padrosia RAFINESQUE 61painteri FOX (Culicoides) 68pallescens LANE & FORATTINI (Stilobezzia) 87pallida SPINELLI & GROGAN (Parabezzia) 84pallidipennis SPINELLI & GROGAN (Brachypogon) 79pallidipes KIEFFER (Atrichopogon) 46palpalis MACFIE (Atrichopogon) 46palpalis MACFIE (Culicoides) 69Palpomyia MEIGEN 23, 30, 32, 79-80, 86, 89, 90,

92, 95, 102, 122-123, 128, 143-145, 152pampaensis SPINELLI & WIRTH (Culicoides) 67pampana LANE (Palpomyia) 96pampoikilus MACFIE (Culicoides) 71panamensis (LANE & WIRTH) (Downeshelea) 80

panamensis BARBOSA (Culicoides) 65panamensis LANE & FORATTINI (Stilobezzia) 30, 87panamensis RONDEROS & SPINELLI (Leptoconops) 44panamensis WIRTH (Echinohelea) 81panamensis WIRTH (Parabezzia) 84panamericanus FOX (Culicoides) 71pancensis BROWNE (Culicoides) 75Parabezzia MALLOCH 32, 83, 101, 104, 109, 110,

115-116, 139-141, 150, 152-153paracaudata GROGAN & WIRTH (Macrurohelea) 82paracincta WIRTH (Dasyhelea) 27, 60Paradasyhelea MACFIE 32, 40, 75, 101, 105, 137, 152paraensis (GOELDI) (Culicoides) 17, 24, 29, 33, 66-67paraensis (LANE) (Nilobezzia) 92paraensis LANE (Palpomyia) 96paraensis LANE (Stenoxenus) 99paraensis WIRTH & BLANTON (Brachypogon) 79, 112parahybae MACFIE (Dasyhelea) 60paraignacioi SPINELLI (Culicoides) 69paramaruim WIRTH & BLANTON (Culicoides) 69paranaensis SPINELLI & WIRTH (Paryphoconus) 98parascopus WIRTH & BLANTON (Culicoides) 65Parastilobezzia WIRTH & BLANTON 40-41, 84, 101,

110, 139Paryphoconus ENDERLEIN 32, 41, 97, 102, 123, 128-

129, 144-146, 152patagonica INGRAM & MACFIE (Dasyhelea) 60patagonica INGRAM & MACFIE (Forcipomyia) 50patagonica INGRAM & MACFIE (Palpomyia) 96patagonica INGRAM & MACFIE (Stilobezzia) 85patagonica SPINELLI & GROGAN (Diaphanobezzia) 80patagoniensis RONDEROS (Leptoconops) 43patagoniensis RONDEROS & SPINELLI (Culicoides) 17, 64patulipalpis WIRTH & BLANTON (Culicoides) 72paucienfuscatus BARBOSA (Culicoides) 74paulistana FORATTINI & RABELLO (Dasyhelea) 27, 61paulistensis LANE (Palpomyia) 96paulistensis LANE (Paryphoconus) 98paulistensis LANE (Stilobezzia) 87pectinatus MACFIE (Atrichopogon) 46Pedilohelea DE MEILLON & WIRTH 48, 55pellucida INGRAM & MACFIE (Diaphanobezzia) 80Pellucidomyia MACFIE 32, 89, 101, 117, 142, 152penicillatus DELÉCOLLE & RIEB (Atrichopogon) 46


penthesileae MACFIE (Dasyhelea) 27, 60penultimata WIRTH (Forcipomyia) 55perstans (Dipetalonema ) 33peruviana CLASTRIER & WIRTH (Forcipomyia) 55peruvianus FELIPPE-BAUER (Culicoides) 67peruvianus KIEFFER (Atrichopogon) 46petiolata MALLOCH (Parabezzia) 83, 113petrocchiae SHANNON & DEL PONTE (Leptoconops)

44, 151Phaenobezzia HAESELBARTH 32, 97, 102, 120, 122,

128, 143, 145, 152phaeonotus WIRTH & BLANTON (Culicoides) 71Phasmidohelea MAYER 53-55philotherma MACFIE (Dasyhelea) 61phlebotomus (WILLISTON) (Culicoides) 17-18, 29, 33, 69Physohelea GROGAN & WIRTH 40, 90, 102, 117,

142, 152Phytohelea REMM 26, 55picadoae SPINELLI & BORKENT (Culicoides) 72piceiventris KIEFFER (Atrichopogon) 46pichindensis BROWNE (Culicoides) 73picta (COQUILLETT) (Stilobezzia) 87pictipennis (PHILIPPI) (Culicoides) 70-71pictipennis CLASTRIER (Atrichopogon) 46pictoni MACFIE (Forcipomyia) 26, 51pifanoi ORTÍZ (Culicoides) 72, 74pilosa SPINELLI & GROGAN (Notiohelea) 83pilosus WIRTH & BLANTON (Culicoides) 65pinamarensis SPINELLI (Forcipomyia) 51piroskyi CAVALIERI (Forcipomyia) 50pistiae (INGRAM & MACFIE) (Phaenobezzia) 97, 114, 130platensis (BRÈTHES) (Forcipomyia) 49plaumanni CLASTRIER & WIRTH (Forcipomyia) 55plaumanni SPINELLI & WIRTH (Alluaudomyia) 76plaumanni SPINELLI (Culicoides) 69plebeja (LOEW) (Palpomyia) 114, 124poikilonotus MACFIE (Culicoides) 65pollex BORKENT & FORSTER (Dasyhelea) 61polypori WIRTH & BLANTON (Culicoides) 69polystictus KIEFFER (Culicoides) 75poncai LANE & WIRTH (Monohelea) 83popayanensis WIRTH & LEE (Culicoides) 63poulaineae INGRAM & MACFIE (Forcipomyia) 50-51prima CLASTRIER (Alluaudomyia) 76

propinqua (WILLISTON) (Forcipomyia) 50propinquus MACFIE (Culicoides) 75propriipennis MACFIE (Culicoides) 72Protersesthes KIEFFER 43pseudocrescentis TAVARES & LUNA DIAS (Culicoides) 72Pseudoculicoides MALLOCH 59pseudodiabolicus FOX (Culicoides) 17, 69pseudofasciata GROGAN & WIRTH (Amerohelea) 92pseudofulvus SPINELLI (Stenoxenus) 99pseudogibbera SPINELLI & WIRTH (Bezzia) 94pseudoincisurata WAUGH and WIRTH (Dasyhelea) 106,

111, 118, 125pseudolacustris DIPPOLITO & SPINELLI (Palpomyia) 96pseudoparaensis SPINELLI & GROGAN (Brachypogon) 79pseudovenustula SPINELLI & WIRTH (Bezzia) 94pseudunguis SPINELLI & GROGAN (Parabezzia) 84Psychophaena PHILIPPI 40, 70-71Pterobosca MACFIE 20, 56pulchripennis MACFIE (Culicoides) 73pulchripennis MACFIE (Sphaeromias) 76pulchripes KIEFFER (Bezzia) 31, 94pulicaris (LINNAEUS) (Culicoides) 132pullata (WIRTH) (Mallochohelea) 91pumila MACFIE (Dasyhelea) 61punctipennis (WILLISTON) (Bezzia) 93-94punctipes WIRTH (Stilobezzia) 87punctiradialis SPINELLI & WIRTH (Alluaudomyia) 77punctivenosa WIRTH & GROGAN (Alluaudomyia) 77punctulata LANE (Stilobezzia) 87puracensis WIRTH & DOW (Forcipomyia) 57puracensis WIRTH & LEE (Culicoides) 17, 63pusilla (LUTZ) (Dasyhelea) 27, 61pusilloides WIRTH & BLANTON (Culicoides) 63pusillus LUTZ (Culicoides) 22, 33, 63pygmaea (WILLISTON) (Dasyhelea) 61pyrsonota MACFIE (Dasyhelea) 61quartibrunneus BORKENT & PICADO (Atrichopogon) 46quasicornuta SAUNDERS (Forcipomyia) 26, 52quasidentica FELIPPE-BAUER & QUINTELAS

(Downeshelea) 81quasiingrami MACFIE (Forcipomyia) 49quasiparaensis CLASTRIER (Culicoides) 67quatei SPINELLI & GROGAN (Borkenthelea) 78quatei WIRTH (Forcipomyia) 50


quechua MARINO & SPINELLI (Forcipomyia) 151rabelloi LANE (Stilobezzia) 30, 87raccurti SPINELLI & GROGAN (Parabezzia) 84rachoui TAVARES & SOUZA (Culicoides) 67raleighi MACFIE (Forcipomyia) 25, 50rangeli ORTÍZ & MIRSA (Culicoides) 72raposoensis CLASTRIER & WIRTH (Forcipomyia) 55raposoensis SPINELLI & WIRTH (Bezzia) 94raposoensis WIRTH & BARRETO (Culicoides) 64raposoi DE MEILLON & WIRTH (Forcipomyia) 55rava INGRAM & MACFIE (Stilobezzia) 86recifei BARBOSA (Culicoides) 69recifensis BARBOSA (Culicoides) 74reconditus CAMPBELL & PELHAM-CLINTON

(Culicoides) 135redactus BORKENT & PICADO (Atrichopogon) 46remigatus EWEN (Atrichopogon) 25, 46remigera UTMAR & WIRTH (Forcipomyia) 48reticulatus LUTZ (Culicoides) 29, 74rettenmeyerorum CLASTRIER & WIRTH (Forcipomyia) 55reynoldsi INGRAM & MACFIE (Dasyhelea) 61Rhynchoforcipomyia WIRTH & DOW 40, 48, 56Rhynchohelea WIRTH & BLANTON 84, 101, 103, 107,

134, 137-138, 152ricardoi RONDEROS & SPINELLI (Leptoconops) 44richardsi MACFIE (Echinohelea) 81ringueleti SPINELLI (Brachypogon) 79riojana SPINELLI & MARINO (Forcipomyia) 58rioplatensis MARINO & SPINELLI (Forcipomyia) 26, 51rodriguezi ORTÍZ (Culicoides) 72roldani SPINELLI & WIRTH (Bezzia) 31, 94ronderosae SPINELLI & BORKENT (Culicoides) 65ronderosi GROGAN & WIRTH (Amerohelea) 92rondoniensis SPINELLI & DIPPOLITO (Groganhelea)

90, 114, 120, 150roraimensis FELIPPE-BAUER & SPINELLI (Monohelea) 83rostratus WIRTH & BLANTON (Culicoides) 25, 63roubaudi CLASTRIER & DELÉCOLLE (Forcipomyia) 58rozeboomi BARBOSA (Culicoides) 68-69rubriceps KIEFFER (Clinohelea) 88rufa KIEFFER (Heteromyia) 89ruizi FORATTINI (Culicoides) 69rusticus MACFIE (Atrichopogon) 47sabalitensis UTMAR & WIRTH (Forcipomyia) 48

saintjusti TAVARES & RUIZ (Culicoides) 68Saliohelea WIRTH & RATANAWORABHAN 48, 57saltaensis SPINELLI & WIRTH (Culicoides) 65saltanensis LANE & DURET (Clinohelea) 88saltensis (CAVALIERI) (Forcipomyia) 52sanctaeclarae MACFIE (Atrichopogon) 47sanctaeclarae WIRTH (Forcipomyia) 57sanctibernardini KIEFFER (Stilobezzia) 86sanctilaurentii KIEFFER (Atrichopogon) 47sanmartini WIRTH & BARRETO (Culicoides) 67santaemarthae (KIEFFER) (Dasyhelea) 61santanderi BROWNE (Culicoides) 73saundersi EWEN (Atrichopogon) 25, 47, 132sayhuequei MARINO & SPINELLI (Forcipomyia) 58scabra (COQUILLETT) (Palpomyia) 96Schizoforcipomyia CHAN & LEROUX 57Schizonyxhelea CLASTRIER 30, 32, 41, 85, 101, 107,

138, 152schmitzi SPINELLI & GROGAN (Brachypogon) 79schmitzorum DIPPOLITO & SPINELLI (Bezzia) 94schnacki SPINELLI & GROGAN (Clastrieromyia) 95, 114schnacki SPINELLI (Alluaudomyia) 29, 77schwarzii (COQUILLETT) (Nilobezzia) 30, 92scissurae MACFIE (Dasyhelea) 61scopus ROOT & HOFFMAN (Culicoides) 65scorzai ORTÍZ (Culicoides) 75scutata LANE & FORATTINI (Stilobezzia) 87seminole WIRTH (Forcipomyia) 26, 52-53sentior (INGRAM & MACFIE) (Austrosphaeromias) 90sequax (WILLISTON) (Atrichopogon) 47setifer (LUTZ) (Culicoides) 64setiger MACFIE (Stenoxenus) 99setigera SAUNDERS (Forcipomyia) 25, 49setigera SPINELLI & WIRTH (Bezzia) 94setosa WIRTH (Macrurohelea) 82setosicrus (KIEFFER) (Forcipomyia) 49setosilateralis BORKENT & PICADO (Atrichopogon) 47seudoobfuscatus SPINELLI (Atrichopogon) 47sexpunctata SPINELLI & WIRTH (Alluaudomyia) 77sexvittata WIRTH (Forcipomyia) 51shannoni (INGRAM & MACFIE) (Forcipomyia) 58shannoni (WIRTH & BLANTON) (Austrohelea) 77,

113, 148shannoni INGRAM & MACFIE (Dasyhelea) 61


silvicola MACFIE (Stilobezzia) 87similans LANE & FORATTINI (Stilobezzia) 87similis SPINELLI & GROGAN (Macrurohelea) 82similis SPINELLI & MARINO (Atrichopogon) 47similis SPINELLI (Amerohelea) 92simplex LANE & FORATTINI (Stilobezzia) 87sinclairi BORKENT (Dasyhelea) 61sinuosa DOW & WIRTH (Forcipomyia) 58smarti MACFIE (Echinohelea) 81smithi (LEWIS) (Mallochohelea) 114snowi LANE (Bezzia) 31, 94soibelzoni MARINO & SPINELLI (Forcipomyia) 57somuncurensis MARINO & SPINELLI (Forcipomyia) 51sonorensis WIRTH & RATANAWORABHAN

(Paryphoconus) 98sordidipes (MACFIE) (Amerohelea) 30, 92soriai WIRTH & WAUGH (Dasyhelea) 61soriai WIRTH (Forcipomyia) 51spangleri DE MEILLON & WIRTH (Forcipomyia) 55spangleri WIRTH (Parabezzia) 84spathicerca WIRTH (Dasyhelea) 61spatulifera SAUNDERS (Forcipomyia) 25, 49spatuliformis SPINELLI & GROGAN (Brachypogon) 79spatuligera MACFIE (Forcipomyia) 51Sphaerohelea SPINELLI & FELIPPE-BAUER 41, 92,

102-103, 122-123, 134, 143-144, 152spilmani DE MEILLON & WIRTH (Forcipomyia) 55spinellii GROGAN & WIRTH (Amerohelea) 82spinellii WIRTH & GROGAN (Alluaudomyia) 77spinellii WIRTH & GROGAN (Diaphanobezzia) 80spinifemur (LANE) (Palpomyia) 96spinifemur WIRTH & BLANTON (Lanehelea) 91spinosa LUTZ (Palpomyia) 96spinosa SAUNDERS (Forcipomyia) 26, 59spinosus BORKENT & PICADO (Atrichopogon) 47spurius WIRTH & BLANTON (Culicoides) 67squamithorax CLASTRIER (Forcipomyia) 53squamitibia LUTZ (Forcipomyia) 49, 51squamosa LUTZ (Forcipomyia) 55steineri SPINELLI & WIRTH (Paryphoconus) 98Stenoxenus COQUILLETT 32, 99, 102, 122-123, 129,

143-144, 146, 152stigmalis (COQUILLETT) (Brachypogon) 111stigmalis WIRTH (Culicoides) 75

Stilobezzia KIEFFER 17, 24, 32, 40, 84-86, 101-103, 115-117, 126, 134, 140-142, 149, 152

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


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

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


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

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

Documents

Neotropical Ceratopogonidae Diptera Insecta AquaticBiodiversity in Latin America