Ceratopogonidae (Dipetara) Neotropicales (Borkent & Spinelli, 2007) PDF

Aquatic Biodiversity in Latin America ABLA
Biodiversidad Acutica en Amrica Latina Vol. 4
FOR MANY YEARS,
Neotropical Ceratopogonidae (Diptera: Insecta)

STUDENTS FROM NUMEROUS COUNTRIES HAVE COMPLAINED
ABOUT THE LACK OF TOOLS TO IDENTIFY AQUATIC INVERTEBRATES FROM
LATIN 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 COVER
NEOTROPICAL REPRESENTATIVES IN DUE DETAIL. THE FEW THAT DO, HOWEVER,
Art Borkent & Gustavo R. Spinelli

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 SCIENTISTS
FROM 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 AQUATIC
BIODIVERSITY. THE SERIES WILL BE AN IMMINENT TOOL FOR ANY BIOLOGICAL
LIBRARY.
Volume 4
This book brings together, for the first time, all known information on the
51 genera and 1095 described Neotropical species of biting midges. An
overview of the group includes a discussion of bionomics, adaptations of the
various life stages, habitats, their ecological roles (including disease transmis-
sion), collecting and preparation techniques, zoogeography and classification. A
key to adults (including males) provides the first key to all the genera of this
region. Limited keys to eggs and larvae are presented. Keys are written in both
English and Spanish. A table of all previous descriptions of immatures and a
catalog of all Neotropical species is included.
This book will be of interest to systematists, ecologists, medical entomolo-
gists and students wishing to undertake innovative research in a remarkably
interesting and instructive group of insects.
ABLA Vol. 4 Art Borkent & Gustavo R. Spinelli: Neotropical Ceratopogonidae 1
Aquatic Biodiversity in Latin America
Biodiversidad Acutica en Amrica Latina
Volume 4
Volumen 4
2 ABLA Vol. 4 Art Borkent & Gustavo R. Spinelli: Ceratopogonidae Neotropicales
BIODIVERSIDAD ACUTICA EN AMRICA LATINA
Volumen 4
CERATOPOGONIDAE
NEOTROPICALES
(DIPTERA: INSECTA)
por
Lector Cientfico a los Editores:

William L. Grogan
Editores de la Serie:
Joachim Adis, Jorge R. Arias,
Guillermo Rueda-Delgado & Karl Matthias Wantzen
SofiaMoscow
2007
AQUATIC BIODIVERSITY IN LATIN AMERICA
Volume 4
NEOTROPICAL
CERATOPOGONIDAE
(DIPTERA: INSECTA)
by
Scientific Reader to the Editors:

William L. Grogan
Series Editors:
Joachim Adis, Jorge R. Arias,
SofiaMoscow
2007

Series Editors/Editores de la Serie:

Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen
VOLUME 4. VOLUMEN 4.
NEOTROPICAL CERATOPOGONIDAE CERATOPOGONIDAE NEOTROPICALES
(DIPTERA: INSECTA) (DIPTERA: INSECTA)
by por
Art Borkent & Gustavo R. Spinelli Art Borkent & Gustavo R. Spinelli
Scientific Reader to the Editors: Lector Cientfico a los Editores:
William L. Grogan William L. Grogan
Front cover:
Female Forcipomyia (Microhelea) sp. feeding on phasmid antenna (Chlorosphasma sp.)
in Costa Rica; Amazon River near Manaus
(photos: P. Naskrecki, WJ. Junk; design: Zheko Aleksiev & Elke Bustorf).
Aquatic Biodiversity in Latin America Vol. 4

ISSN: 1312-7276
First published 2007
ISBN-13: 978-954-642-301-6
The book should be cited as follows:

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

Habitus of female Culicoides variipennis COQUILLETT

(Nearctic Region; from McALPINE et al., 1981).
Series Editors: Joachim Adis, Jorge R. Arias,

Biodiversity is a key word in science and global management schemes; however very few
people are able to identify the species and their ecology that make up biodiversity. For
many years, researchers and students from numerous countries complain about the lack of
tools to identify aquatic invertebrates from Latin America. Keys found in accepted
entomological textbooks are mostly highly limited, superficial and rarely cover Neotropical
biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology
is scattered throughout the literature in many single publications.
An international team of editors have combined their efforts with Pensoft Publishers
to launch a new major series on the Aquatic Biodiversity of Latin America (ABLA). Their
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and status of the taxa (written in English) is combined with illustrated identification keys to
families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists,
hydrobiologists, biogeographers, conservationists and students interested in aquatic
biodiversity. The series will be an essential tool for any biological library.
Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher.

Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642-
258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and
photos (15 plates in color); publication date: March 2006.
Volume 2: Ephemeroptera of South America by Eduardo Domnguez, 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-
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Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarn & Cecilia L.

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

R. Spinelli.
Pensoft Publishers, Sofia-Moscow, ISBN-13: 978-954-642-301-6, 165x240 mm, 198 pp.,
including 26 plates of figures in line drawings and photos; publication date: June 2007.
Ceratopogonidae are found in virtually every habitat with even a small amount of sustained
moisture, are notorious as severe pests of man and domestic animals, and are one of the
most common fly families in traps. Yet they are by far the most poorly understood of all
the biting flies. In part, this is due to their remarkably diversity, both in numbers of species
and in their various adaptations. Larvae range from being detritivores and scavengers to
predators and adult females of various species feed on a wide array of vertebrates and
invertebrates. Some females draw blood from conspecific males while they are mating.
Adults are also important pollinators of a wide array of plants.
This book brings together, for the first time, all known information on the 51 genera
and 1095 described Neotropical species of biting midges. An overview of the group includes
a discussion of bionomics, adaptations of the various life stages, habitats, their ecological
roles (including disease transmission), collecting and preparation techniques, zoogeography
and classification. A key to adults (including males) provides the first key to all the genera
of this region. Limited keys to eggs and larvae are presented. Keys are written in both
English and Spanish. A table of all previous descriptions of immatures and a catalog of all
Neotropical species is included.
This book will be of interest to systematists, ecologists, medical entomologists and
students wishing to undertake innovative research in a remarkably interesting and instructive
group of insects.
ADDRESSES
Authors:
DR. ART BORKENT M.Sc. GUILLERMO RUEDA-DELGADO
691-8th Avenue SE Grupo de Investigacin en Cuencas y
Salmon Arm, British Columbia V1E 2C2 Humedales Tropicales GICHT
Canada UNIMAGADALENA
e-mail: aborkent@sunlite.ca Universidad de Bogot Jorge Tadeo Lozano
Laboratorio de Limnologa
DR. GUSTAVO R. SPINELLI Carrera 4 No. 22-61
Divison Entomologia Bogot, D.C., Colombia
Museo de La Plata e-mail: guillermo.ruedadelgado@utadeo.edu.co
Paseo Del Bosque s/n
1900 La Plata, Argentina DR. KARL MATTHIAS WANTZEN
e-mail: spinelli@fcnym.unlp.edu.ar Institute of Limnology
University of Konstanz
Series Editors: Postfach M 659
PROF. DR. JOACHIM ADIS 78457 Konstanz, Germany
Max-Planck-Institute for Evolutionary e-mail: matthias.wantzen@uni-konstanz.de
Biology
Tropical Ecology Working Group Scientific Reader to the Editors:
Postfach 165 DR. WILLIAM L. GROGAN
24302 Pln, Germany Department of Biological Sciences
e-mail: adis@evolbio.mpg.de Salisbury University
Salisbury, Maryland 21801-6837
DR. JORGE R. ARIAS e-mail: wlgrogan@salisbury.edu
5870 Colfax Avenue
Alexandria, Virginia 22311, USA
e-mail: jaria2@fairfaxcounty.gov
FOREWORD

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

relationships among taxa including references to the fossil record and a preliminary
discussion of the zoogeography of selected species. A synopsis of the medical importance
of ceratopogonids in Latin America is also included as are useful syntheses of information
about Neotropical ceratopogonids (e.g., Table 1 includes references for all species whose
immature stages have been described). The color photographs in the bionomics chapter of
biting midges feeding on other insect species are particularly striking. Another useful feature
found in the Catalog of the Ceratopogonidae of the Neotropical Region is the designation of
species in the genera Culicoides and Leptoconops which are known to bite humans. Borkent and
Spinelli acknowledge more than once that many more ceratopogonid species will be described
from Latin America following expeditions to poorly collected areas. Their foundational work
will most certainly facilitate future taxonomic work. The authors have provided an invaluable
resource for scientists who investigate aquatic biodiversity in the Neotropics.
STEVE MURPHREE
Editor, Ceratopogonidae Information Exchange
Belmont University
Nashville, Tennessee, USA
PREFACE
The Ceratopogonidae are commonly known as biting midges, no-see-ums or punkies, in

Spanish as manta blanca, polvorines, or jejenes (which also may refer to Simuliidae), and in
Portuguese as maruim or mosquito plvora. They have a bad reputation as being nasty
biters that pester humans and domestic animals and, in some instances, transmit harmful
diseases. Because of their small size, the females of some species can pass through screen
and mesh that keeps other biting pests outside and these can make life insufferable. Biting
midges may occur in such huge numbers that in some areas people are driven indoors (or
complain loudly and then suffer numerous itchy bites!).
Few people, however, realize that this group of flies provides important services in a
wide array of ecosystems. Some species are important pollinators of such plants as cacao
(without them we wouldnt enjoy chocolate!) and rubber trees, and the larvae of many are
important predators of other organisms in semiaquatic and aquatic habitats. The adults of
most biting midges actually suck blood from other insects and may even be important
vectors of viruses that kill caterpillars.
The biting midges are a diverse group in both numbers of species and in their habits
and there are some intriguing stories associated with the way they obtain their food, mate
and the kinds of different habitats they occupy. This contribution provides the basis for
understanding the family in the Neotropical Region, allows for their identification to the
generic level, gives a synopsis of each genus, details some aspects of their behaviour and
ecology, discusses various collecting and preparation techniques, and provides a catalog of
the Neotropical species.
The first author thanks his wife and dearest friend Annette BORKENT for providing
moral and financial support throughout the writing of this contribution and for her steadfast
faith in his work. Pablo DELLAP and Maria M. RONDEROS provide substantial advice and
assistance in preparing the plates and we express our gratitude to them. A number of the
figures were skillfully produced by Ana BRENES (INBio) and we appreciate permission
from the Instituto Nacional de Biodiversidad in Costa Rica to reproduce these here. Nelida
CALIGARIS redrew some of the illustrations of previously published figures. The Series
Editors and the Scientific Reader are thanked for valuable suggestions, improving the content
and the final shape of the book. Many of the figures reproduced here are from Chapter 28
of Volume 1 of the Manual of Nearctic Diptera (MCALPINE et al. (eds), Agriculture Canada
Monograph 27, Agriculture & Agri-Food Canada, 1981) and we are grateful for permission
to do so from the Minister of Public Works and Government Services Canada (2006).
Annia PICADO (INBio) has slide mounted tens of thousands of Ceratopogonidae from
Costa Rica with tremendous skill and this has helped our understanding of Neotropical
biting midges immeasurably. Heron HUERTA provided a careful review of this work and we
thank him for his detailed critique.
We gratefully acknowledge permission to use photographs from several colleagues

who are noted in the figure captions.
THE AUTHORS
Table of Contents
1 B References to species of Neotropical Ceratopogonidae

IONOMICS 15
described as eggs, larvae and/or pupae (Table 1) 25

Number of named species with described immatures for
genera of Neotropical Ceratopogonidae (Table 2) 32
Organisms transmitted by Neotropical
Ceratopogonidae (Table 3) 33
2C APTURE AND STUDY OF CERATOPOGONIDAE 34
3R EARING CERATOPOGONIDAE 38
4P HYLOGENETIC RELATIONSHIPS AND FOSSIL

RECORD OF CERATOPOGONIDAE 39
5Z OOGEOGRAPHY 40
6 C Catalog of the Ceratopogonidae of the Neotropical

LASSIFICATION 42
Region (Table 4) 43
7I DENTIFICATION
Numbers of named valid species in each genus of
100
Neotropical Ceratopogonidae (Table 5) 101

Diagnosis of the Ceratopogonidae 102
Key to the genera of adult Ceratopogonidae in the

Neotropical Region 103
Key to the genera of eggs of Ceratopogonidae in the
Key to the genera of larvae of Ceratopogonidae in the
Clave para los gneros de adultos de Ceratopogonidae
de la Regin Neotropical 134
Clave para los huevos de gneros de Ceratopogonidae
Clave para las larvas de gneros de Ceratopogonidae
8 SNYNOPSIS OF THE
EOTROPICAL
CERATOPOGONIDAE OF THE
REGION 148
9R EFERENCES 154
10 S UBJECT INDEX 182

1
BIONOMICS
Ceratopogonidae are one of the common of fly families in the Neotropical Region (BROWN,
2005). Adults may be found from the coast (some are intertidal) to the highest elevations. We
have records of Forcipomyia MEIGEN and Dasyhelea KIEFFER from 3700 to 4000 meters in
northwestern Argentina and Forcipomyia and Atrichopogon KIEFFER from 3819 meters in Costa
Rica (which is that countrys highest point). They are present in nearly all habitats where there
is even a little long-lasting moisture. Adults of a few species are vicious biters of humans and
domestic animals and some act as vectors of various diseases or parasites (BORKENT, 2004).
There is little information on the specific life cycle of Ceratopogonidae in the Neotro-
pical Region. Further to this, our survey of the literature (Tables 1, 2) indicates that fewer
than 15 percent of the species named from this area are described in even one immature
stage. Nineteen genera are entirely unknown. Some descriptions, especially those that are
older, are likely based on misidentified species and these immatures need to be restudied.
Aware that many species remain to be described even as adults, there is a rich field available
for investigation. Undoubtedly, many of the larvae of Ceratopogonidae play an important
ecological role that is nearly entirely uninvestigated. Rearings of larvae will certainly provide
important information regarding the structure of tropical ecosystems but clearly, the first
step is to have these described and to have keys made available. At the present time there are
no reliable keys to the larvae or pupae to even the generic level although these stages are
presently being studied at a worldwide level (BORKENT, in prep.).
All members of the family have an egg, four larval instars, a pupal and an adult stage.
Eggs may be laid on moist substrate or, in the case of fully aquatic species, in water. The
fact that adult ceratopogonids, as a group, are present throughout the year in tropical areas
suggests that many species have multiple life cycles, without the third or fourth instar
experiencing a diapause as do nearly all species in northern temperate and arctic regions. In
some tropical areas, it appears that adult Ceratopogonidae may be somewhat more diverse
during the rainy season. It is unknown how tropical Ceratopogonidae respond in areas with
a pronounced dry season. The life cycle of Ceratopogonidae in the southern temperate
zone of South America is entirely unstudied but the presence of both numbers of species
and adults in late spring in this area indicates that many species likely overwinter in the late
larval stage, as they do in the northern temperate region.
Larvae are important detritivores or predators in semiaquatic and aquatic systems of
all sizes, from tiny tree holes and the water held by tropical flowers, to large rivers and lakes,
where they may be an important source of food for other invertebrates and fish. In tropical
areas of Latin America, the immatures of Ceratopogonidae form an important compo-
nent of arboreal aquatic systems (such as in bromeliads and tree holes). The most diverse
group, both in terms of numbers of taxa and structurally, are those in the subfamily 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/or
taxon, feeding on microorganisms or lashing out at prey. Some prey are swallowed whole
(e.g. nematodes) but in the case of larger prey, which are mostly other insect larvae, the
biting midge larvae of some taxa burrow through the cuticle of the host and eat their
insides. It seems certain that in many habitats, biting midge larvae play an important role in
the biological control of other insects, but this has not been studied in much detail.
The immatures of each of the four subfamilies differ substantially.
The Leptoconopinae includes just one genus, Leptoconops SKUSE in the Neotropical
Region. The group is an ancient one and fossil Leptoconops are known from Lebanese am-
ber, 121 million years old. Their larvae live in wet sand (especially on marine beaches) or in
cracked alkaline, clay soils. The sluggish larvae burrow through wet sand or loose clay soil
and feed on microorganisms by raking their heads over the substrate.
Larvae of Forcipomyiinae are aquatic, semiaquatic or terrestrial. When they are in
aquatic habitats, they are generally restricted to lentic (standing) waters although a few are
known from streams and small rivers and are associated with mosses and algae. They retain
the primitive anterior and posterior prolegs and therefore can walk on surfaces beneath or
above water. As they move, they graze the substrate surface for microorganisms, looking
somewhat like cattle eating grass. Some species feed on decaying vegetation and a few
species may be found associated with manure. The larvae of most species have specialized
setae along the length of their bodies that secrete a sticky fluid that repels ants. When the
larvae pupate, most retain the larval exoskeleton on the posterior end of their bodies.
Larvae of Dasyheleinae are aquatic and semiaquatic and tend to be sluggish. They
have posterior hooks on their abdomens and move slowly over the substrate in which they
are found. Most species live in small water bodies such as those in epiphytes, tree holes,
broken or damaged bamboo, rotten banana stems, fruit husks and small rock pools on the
margins of streams and rivers. A few species are common in mangrove swamps and salt
marshes. Larvae may also be found in wet vegetation. One species in Costa Rica is a leaf-
miner of the floating leaves of Salvinia and another is found in the wet refuse piles of some
ants (BORKENT, pers. obs.). Larvae feed on detritus, algae, and, in a few species, on carrion
(dead insects). There are some species whose adults are associated with rocky areas in the
upper tidal zone of coastal Costa Rica but the immatures of these are not known.
Larvae of the Ceratopogoninae, the most species-rich group of biting midges, are prog-
nathous, lack prolegs and hooks (at least in instars 2-4), and their abdominal segments are not
secondarily divided. Larvae of most species swim in a characteristic, very rapid, serpentine
motion through the water column. When on or in substrate they assume a slower motion,
working their way through wet substrate, eating microorganisms or lashing out at larger prey,
looking very much like small snakes. Some smaller invertebrates are swallowed whole but in
the case of larger prey, which are mostly other insect larvae, the biting midge larva penetrates
the cuticle of the prey and devours the internal contents. The earlier phylogenetic lineages
tend to feed on microorganisms and/or smaller invertebrates, often ingesting these whole.
The more derived lineages, especially in the tribes Heteromyiini, Sphaeromiini, and Palpomyiini,
feed on larger invertebrates such as chironomid larvae and sometimes burrow into the body
of their host. In many habitats, biting midge larvae probably exert an important biological
control on other insects. Larvae of the species-rich genus Culicoides LATREILLE may be found
in decaying vegetation, fruits, and manure, epiphytes, tree holes, wet mud, mangrove swamps
and a large array of other aquatic habitats. In streams and rivers, Ceratopogoninae larvae may
be an important source of food for other invertebrates and fish. In general, the most 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, and
larger lotic and lentic habitats such as rivers and lakes.
Pupae are generally sluggish and move their abdomens slowly in more or less circular
movements until they find an appropriate resting position in the midst of the substrate or
at the water surface where their paired respiratory organs can obtain air. The pupae of
some Sphaeromiini move above the water line and have specialized membranous pads on
the venter of the abdomens which they use to stick to the dry substrate. BORKENT & CRAIG
(2001) and CAZORLA & MARINO (2004) described unusual pupae of Stilobezzia KIEFFER from
Costa Rica and Peru respectively, which use their respiratory organs to obtain oxygen by
piercing the submerged leaves of aquatic plants.
Adult females of Leptoconops, Culicoides and Forcipomyia (only in the subgenus Lasiohelea
KIEFFER) are haematophagous on vertebrates, needing the blood to develop their eggs.
Female Leptoconops and Culicoides (Fig. 1B, frontispiece) may occur in large numbers in 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 to
their 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 marine
beaches and adult females are daytime biters, generally on the lower legs of humans. The
adults of some species (including L. bequaerti (KIEFFER) in Central America) have an 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 Neotropical
Region, from Mexico to southern Chile (Puerto Natales, at almost 52 S), from the coast to
high elevations (at least 3320 m in the Colombian Andes (WIRTH & LEE, 1967)). Although
the females of a few species may bite during the day and may occur in huge numbers, most
are crepuscular or nocturnal feeders.
Of the 266 species of Culicoides known from the Neotropical Region, 70 (26%) have
been recorded feeding on humans (Table 4) but of these only the following occur in 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 coastal
Brazil, C. insinuatus ORTZ & LEON, C. puracensis WIRTH & LEE at high elevations in the Andes
of Colombia, C. pseudodiabolicus FOX and C. paraensis (GOELDI) in the Amazon basin, C. lahillei
(ICHES) and C. debilipalpis LUTZ in Paraguay and northeastern Argentina, C. caridei (BRTHES) in
Uruguay and the province of Buenos Aires, Argentina, and C. patagoniensis RONDEROS & SPINELLI
in Argentinean and Chilean Patagonia. Further pestiferous species will likely be discovered in
the largely unknown Andes. Females of some other species feed on domestic animals. For
example, females of C. debilipalpis feed on both humans and horses and C. arubae FOX &
HOFFMAN, C. foxi ORTZ and C. barbosai WIRTH & BLANTON have been collected from mules or
horses. Culicoides insignis LUTZ is typically associated with cattle. Culicoides phlebotomus, a well
known species in the Neotropics biting humans and dogs on marine beaches, is also known to
suck blood from leatherback turtles that are on the beach laying their eggs (BORKENT, 1996).
The hosts of the remaining species of Culicoides are largely unknown.
The biting habits of female adult Forcipomyiinae, including only two genera, Forcipomyia
and Atrichopogon, are remarkably diverse. In general, female adults act as ectoparasites of large
insects, with some sucking blood from their hosts for extended periods of time. In particular,
those Forcipomyia in the ixodoides species group of the subgenus Microhelea KIEFFER attack
phasmids (Fig. 2B) and katydids (Figs. 2A, 4A) and when replete and developing their eggs,
look like small ticks (CLASTRIER & WIRTH, 1995). One female of this species group we exam-
ined in Costa Rica had developed 2778 eggs. Other species of Forcipomyia suck hemolymph
Culicoides furens
Det. G.R. Spinelli
2 km N. Montezeuma,
Playa Grande, Costa Rica,
21-XI-2000, A. Borkent
Biting humans
CD5099
A
B
Figure 1. A: Dissected adult Ceratopogonidae on a microscope slide. B: Female Culicoides sp., biting
human (taken by S.A. Marshall in South Carolina, USA).
B
Figure 2. Female Forcipomyia (Microhelea) spp., feeding. A: On katydid abdomen (Orophus sp.) (taken by M.
Zumbado in Costa Rica). B: On phasmid antenna (Chlorophasma sp.) (taken by P. Naskrecki in Costa Rica).
from caterpillars (Fig. 3B), sawfly larvae (WIRTH, 1972), spiders, crane flies, the wing veins of
dragonflies and damselflies (Fig. 3A), lacewings and butterflies (WIRTH, 1956a; WIRTH &
MESSERSMITH, 1971; MARINO & VON ELLENRIEDER, 1999). The feeding habits of female Atri-
chopogon are more poorly known but those of at least one subgenus feed on the blood of adult
Meloidae beetles (BORKENT & ROCHA-FILHO, 2006). Species in the subgenus Forcipomyia (Lasio-
helea) are the only Forcipomyiinae that are vertebrate feeders. Their hosts are poorly known
and females have been recorded feeding only on frogs and lizards in the New World (SPINELLI
et al., 2002). The males and females of many adult Forcipomyia and Atrichopogon feed on nectar
to fuel their flight and are often common on flowers, particularly those that are small and
white or green. One unusual example of consuming sugars is Forcipomyia adults from the
backs of pentastomid bugs in Costa Rica (Fig. 4B).
B
Figure 3. A: Female Forcipomyia (Pterobosca) sp. feeding on damselfly wing (Argia hinei) (taken by D.
Huntington in New Mexico, USA). B: Female Forcipomyia (Microhelea) sp. feeding on caterpillar
(Automeris zugana) (taken by D.H. Janzen in Costa Rica).
B
Figure 4. A: Female Forcipomyia (Microhelea) spp., feeding on katydid (Pristonotus latistylus) (taken by
P. Naskrecki in Costa Rica). B: Forcipomyia sp. females likely feeding on surface sugars on a
pentatomid (taken by M. Moraga in Costa Rica).
Adult females of Dasyhelea have reduced mouthparts and do not suck blood from
either vertebrates or invertebrates. Some species are common at flowers where they, like
many other biting midges, obtain nectar from flowers.
Adult females in the subfamily Ceratopogoninae, other than those of Culicoides, are
predators and feed on other flying insects that are approximately the same size as 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 injects
an 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 the
body of the male and sucks out its contents (DOWNES, 1978). Such females may later be
found with the dried male genitalia still attached to their own abdomen, after the bulk of
the dried male has broken off. The males of these species contribute not only their sperm
and body tissues but also an effective mating plug to the female with which they mate.
BORKENT (2004) provided a list of all diseases and parasites known to be transmitted
by biting midges worldwide. The females of Leptoconops, Forcipomyia (Lasiohelea) and Culi-
coides are the only genera which include species which bite vertebrates (from frogs to birds
to mammals) in the Neotropical Region. Worldwide, Ceratopogonidae are known to trans-
mit 66 viruses, 15 species of protozoa and 26 species of filarial nematodes. It is highly
likely that many more remain undetected. This strongly suggests that Ceratopogonidae will
be found to transmit many more organisms in the Neotropical Region than the six viruses
and eight nematodes currently known (Table 3). For example, it would be quite remarkable
if there were no protozoan parasites of birds in the entire Neotropical Region when at
least nine species of protozoans are known from North America. The adults of those
biting midges that suck hemolymph from large insects (some Atrichopogon and Forcipomyia)
suggest the possibility that they may be involved in transmitting viruses from one insect
host to another, particularly among caterpillars.
Bluetongue is a viral disease of cattle, sheep, goats, and wild ruminants. It is particu-
larly damaging in sheep, where up to half of infected animals may die. In cattle and goats
the disease primarily affects the animals reproductive capabilities. Although it was known
that the virus occurred in Central America (MO et al., 1994) and antibodies were detected in
cattle in the late 1980s in southern Brazil (CUNHA, 1990), there were no published data of
animal illness in the Neotropics until recently. The Office International des Epizooties,
World Health Organization (OIE, 2001) reported the presence of the disease in the state
of Paran, Brazil, in which eight sheep and one goat showed classical symptoms, including
the death of four of these animals, and 70 cattle showed susceptibility as well. More re-
cently, the virus was isolated and characterized from asymptomatic bovines from north-
eastern Argentina (GORCH et al., 2002). Bluetongue occurs virtually worldwide and has
different, phylogenetically unrelated vectors in different regions of the globe, although all
are in the genus Culicoides. In the Neotropical Region the primary vectors are likely C.
insignis, C. pusillus LUTZ (WIRTH & DYCE, 1985; SENZ & GREINER, 1994; GORCH et al., 2002)
and possibly C. filarifer HOFFMAN (SENZ & GREINER, 1994).
There are other diseases of humans and domestic animals that occur elsewhere in the
Neotropical Region. One of the more significant is Oropouche fever, a viral infection pro-
B
Figure 5. A: Female Palpomyia sp. (taken by S.A. Marshall in Ontario, Canada). B: Female Sphaeromias
sp. (taken by K. van der Krieke in the Netherlands).
ducing severe flu-like symptoms, mostly in Brazil. It has been responsible for at least 27
epidemic outbreaks in humans, including one in the state of Par, Brazil in 1978-1980 which
afflicted at least 130,000 people. The disease is primarily transmitted by species of Culicoides
(and particularly by C. paraensis) but has also been isolated (at low levels) from some mosqui-
toes. Outbreaks occur primarily in urban areas, where populations of C. paraensis may occur at
great densities (HOCH et al., 1990; MELLOR et al., 2000). MERCER et al. (2003) reported decaying
platano (plantain: Musa parasidiaca) stems, stumps, flowers, fruits and debris beneath platano
trees, as well as soil beneath a fruiting mamey (Pouteria sapota) tree and organic-rich mud along
a lake shoreline, as larval substrates of C. paraensis in Iquitos, Peru, a city which has suffered
several outbreaks of Oropouche fever. LEDUC & PINHEIRO (1989) provide an excellent 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 thought
of in negative terms. The adults of many genera, primarily Forcipomyia and Dasyhelea, are
important pollinators of some commercial trees such as cacao (Theobroma cacao) and rubber
(Hevea brasiliensis). Species of these ceratopogonid genera, and those of Atrichopogon, Culi-
coides, and Stilobezzia, are likely to be pollinators for a number of other plants in tropical
America (possibly including mango (Mangifera indica) and a number of palms). Further
information about pollination by ceratopogonids may be found in a series of papers by
WARMKE (1951, 1952), SAUNDERS (1959), SAUNDERS & Bowman (1956), WINDER (1977a, b),
WIRTH (1956c) and YOUNG (1982, 1983, 1986a, b). Although it is unpleasant for many, the
vicious attacks on humans and domestic animals by the females of some species of 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 parasites
and 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 stages
of Dasyhelea necrophila SPINELLI & RODRGUEZ 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, larvae

and/or pupae. E refers to egg; L refers to larva and P to pupa. Theses are cited only
when their results have not been published. Duplicated figures or repeated descriptions by
subsequent authors are not listed here. Some descriptions, especially those that are older,
may be based on incorrectly identified species.
Taxon Reference Stages Described

Leptoconops bequaerti (as Leptoconops sp.) PAINTER, 1927 L P
MAYER, 1934 P
LINLEY, 1965a E
LINLEY, 1968 L
Atrichopogon bifidus EWEN & SAUNDERS, 1958 L P

A. caribbeanus EWEN & SAUNDERS, 1958 L P
A. 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 P
A. fusculus or rostratus
(as Kempia haesitans) LENZ, 1934 L P
A. incultus EWEN & SAUNDERS, 1958 L P
A. longicornis EWEN & SAUNDERS, 1958 L P
A. obscurus EWEN & SAUNDERS, 1958 L P
A. remigatus EWEN & SAUNDERS, 1958 L P
A. saundersi EWEN & SAUNDERS, 1958 L P
A. tuberculatus EWEN & SAUNDERS, 1958 L P
Forcipomyia (Caloforcipomyia) caerulea SAUNDERS, 1957 L P

F. (C.) glauca (as monilis) KRIVOSHEINA, 1968 L P
F. (C.) varicolor SAUNDERS, 1957 L P
Forcipomyia (Euprojoannisia) blantoni SORIA & BYSTRAK, 1975 L P

BYSTRAK & WIRTH, 1978 L P
F. (E.) bromeliae SAUNDERS, 1957 L P
F. (E.) calcarata WIRTH, 1952a L P
F. (E.) falcifera SAUNDERS, 1957 L P
F. (E.) longispina SAUNDERS, 1957 L P
F. (E.) mortuifolii SAUNDERS, 1957 L P
BYSTRAK & WIRTH, 1978 L P
F. (E.) setigera SAUNDERS, 1957 L P
F. (E.) spatulifera SAUNDERS, 1957 L P
F. (Forcipomyia) argenteola LANE, 1947c L P

F. (F.) genualis (as raleighi) SAUNDERS, 1957 L P
(as raleighi) GUTSEVICH & GLUKHOVA, 1970 L P
F. (F.) harpegonata WIRTH & SORIA, 1975 L P
Table 1. Continued.

F. (F.) pictoni WIRTH, 1976 L P
F. (F.) rioplatensis SPINELLI et al., 2005a L P
F. (Lasiohelea) attenuata SAUNDERS, 1964 L P

F. (L.) cornuta SAUNDERS, 1964 L P
KRIVOSHEINA, 1968 L
F. (L.) intermedia SAUNDERS, 1964 L P
F. (L.) quasicornuta SAUNDERS, 1964 L P
F. (L.) stylifer LANE, 1947c L P
F. (Lepidohelea) brasiliensis WIRTH & SPINELLI, 1993a P

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

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

WIRTH & SORIA, 1980 L P
F. (W.) lesliei (as bicolor) SAUNDERS, 1957 L P
F. (W.) spinosa SAUNDERS, 1957 L P
F. (W.) terrestris SAUNDERS, 1964 L P
Table 1. Continued.

F. (W.) tuberculata SAUNDERS, 1957 L P
F. sp. (as an Apelma sp.) SAUNDERS, 1925 L P
F. sp. (as species A) SAUNDERS, 1957 L
F. sp. (as species A) SAUNDERS, 1964 L
F. sp. (as species B) SAUNDERS, 1964 L
Dasyhelea atlantis WAUGH & WIRTH, 1976 P

BORKENT, 1991 P
D. bahamensis RONDEROS et al., 2003 L P
D. calvescens WILLIAMS, 1944 E L P
D. cincta (as penthesileae) SPINELLI, 1983b P
D. correntina RONDEROS et al., 2004a P
D. filibranchia (as a Ceratopogon) LUTZ, 1914 P
D. flavicauda SPINELLI & RONDEROS, 1987 P
D. lacustris SPINELLI & RONDEROS, 1987 P
D. mediomunda MINAYA, 1978 P
D. mutabilis THOMSEN, 1937 L P
WIRTH, 1952a P
WAUGH & WIRTH, 1976 P
D. necrophila SPINELLI & RODRGUEZ, 1999 L P
RONDEROS et al., 2003 L P
RONDEROS et al., 2006 E L
D. paracincta BORKENT, 1991 L P
D. paulistana FORATTINI & RABELLO, 1957 P
D. pusilla FORATTINI & RABELLO, 1957 P
Culicoides annettae SPINELLI & BORKENT, 2004a P

C. annuliductus VITALE et al., 1981 P
C. arubae (as sp. 1) FOX, 1942 P
FORATTINI, 1957 P
MURPHREE & MULLEN, 1991 L
C. bambusicola LUTZ, 1913 L
(as sp. 2) FOX, 1942 P
LANE, 1947c L P
BARBOSA, 1952, 1953 L P
FORATTINI, 1957 P
RONDEROS & SPINELLI, 2000 L P
RONDEROS et al., 2000 L P
C. barbosai LINLEY, 1965b P
LINLEY & DAVIES, 1971 E
BLANTON & WIRTH, 1979 P
C. bayano VITALE et al., 1981 P
C. bermudensis JAMNBACK, 1965 L P
Table 1. Continued.

BLANTON & WIRTH, 1979 L P
C. borinqueni LINLEY, 1965b P
C. cancer HOGUE & WIRTH, 1968 L P
C. charruus SPINELLI et al., 1993 P
C. chaverrii SPINELLI & BORKENT, 2004a L P
C. cochisensis MURPHREE & MULLEN, 1991 L
C. crepuscularis THOMSEN, 1937 P
FOX, 1942 P
JAMNBACK, 1965 L P
C. debilipalpis BARBOSA, 1952, 1953 P
FORATTINI, 1957 P
C. dicrourus WIRTH & SORIA, 1981 L P
C. edeni (as possibly haematopotus) JONES, 1961 P
C. fililductus VITALE et al., 1981 P
C. floridensis LINLEY, 1970 P
C. furens PAINTER, 1927 L P
(as dovei) DOVE et al., 1932 E L P
FOX, 1942 P
WIRTH, 1952d L P
FORATTINI, 1957 L P
JONES, 1961 P
LINLEY & KETTLE, 1964 L P
JAMNBACK, 1965 L P
LINLEY & DAVIES, 1971 E
HRIBAR & MULLEN, 1991 L
C. guyanensis FORATTINI & RABELLO, 1956 L P
FORATTINI, 1957 L P
C. haematopotus THOMSEN, 1937 P
JONES, 1961 P
JAMNBACK, 1965 L P
ATCHLEY & WIRTH, 1979 L P
C. hoffmani LINLEY & KETTLE, 1964 L P
C. hylas (as sp. 3) FOX, 1942 P
FORATTINI, 1957 P
Table 1. Continued.

C. insignis FORATTINI et al., 1956 L P
FORATTINI, 1957 L P
LINLEY, 1965b P
SPINELLI et al., 1993 P
C. lahillei LAMBERSON et al., 1992 P
C. luteovenus WIRTH, 1952a L P
C. macieli WIRTH & SORIA, 1981 L P
C. maruim LUTZ, 1913 E P
BARBOSA, 1952, 1953 P
FORATTINI, 1957 P
SPINELLI et al., 1993 P
C. melleus WIRTH, 1952d L P
JAMNBACK et al., 1958 L P
JONES, 1961 P
JAMNBACK, 1965 L P
C. neopulicaris JONES, 1961 P
C. oklahomensis MURPHREE & MULLEN, 1991 L
LAMBERSON et al., 1992 P
C. paraensis MURPHREE & MULLEN, 1991 L
LAMBERSON et al., 1992 P
C. phlebotomus PAINTER, 1927 L P
FOX, 1942 P
FORATTINI, 1957 P
C. reticulatus LUTZ, 1913 L
FORATTINI et al., 1960 L P
C. venezuelensis RONDEROS & SPINELLI, 1989 P
DAZ et al., 2005 E L P
C. sp. PAINTER, 1927 L P
Alluaudomyia bella (as splendida) THOMSEN, 1937 L P

WIRTH & GROGAN, 1981 L P
A. biestroi SPINELLI, 1988 P
A. caribbeana SPINELLI & WIRTH, 1984c P
A. distispinulosa SPINELLI, 1997b P
A. schnacki SPINELLI, 1983b L P
Baeodasymyia christopheri BORKENT & CRAIG, 1999 E L P

B. michaeli BORKENT & CRAIG, 1999 L P
Echinohelea lanei WIRTH, 1994b P

Table 1. Continued.

Parabezzia alexanderi GROGAN & WIRTH, 1977 P
WIRTH & GROGAN, 1981 P
SPINELLI & GROGAN, 1987 P
P. balseiroi SPINELLI & GROGAN, 1987 P
Schizonyxhelea forattinii BORKENT, 2000b L P
Stilobezzia antennalis THOMSEN, 1937 L P

WIRTH & GROGAN, 1981 L P
S. bicolor FORATTINI & RABELLO, 1956 P
S. chaconi LANE & FORATTINI, 1961 P
S. coquilletti LANE & FORATTINI, 1961 P
S. dorsofasciata (as a Palpomyia) LUTZ, 1914 E
S. dubitans LANE et al., 1955 P
S. fiebrigi FORATTINI & RABELLO, 1956 L P
CAZORLA et al., 2006 P
S. glauca LANE et al., 1955 P
S. panamensis FORATTINI et al., 1960 P
S. rabelloi BORKENT & CRAIG, 2001 P
S. wygodzinskyi LANE et al., 1955 P
Clinohelea horacioi SPINELLI & DURET, 1993 P

C. nigripes SPINELLI & DURET, 1993 P
Heteromyia clavata WIRTH & GROGAN, 1977 P

H. wokei WIRTH & GROGAN, 1977 P
Mallochohelea termophila SPINELLI, 1984 P
Neobezzia amnicola MAYER, 1959 P
Nilobezzia schwarzii WIRTH, 1962 P

PALCHICK, 1981 P
Amerohelea sordidipes LANE et al., 1955 P
Bezzia bivittata HRIBAR & MULLEN, 1991 L

B. blantoni SPINELLI & WIRTH, 1989b P
B. brevicornis SPINELLI, 1983c P
SPINELLI & WIRTH, 1989b P
B. bromeliae SPINELLI & WIRTH, 1989b P
B. glabra THOMSEN, 1937 E L P
Table 1. Continued.

WIRTH, 1952a L P
CHAN & LEROUX, 1967 E
PALCHICK, 1981 P
WIRTH, 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 L
HRIBAR, 1993 L
B. pulchripes MAYER, 1959 P
SPINELLI & WIRTH, 1989b P
B. roldani SPINELLI & WIRTH, 1981, 1989b L P
B. snowi SPINELLI & WIRTH, 1991 P
Clastrieromyia dycei SPINELLI & GROGAN, 1986 P
Pachyhelea pachymera SPINELLI, 1983c P
Palpomyia guarani RONDEROS et al., 2004b L P

P. lacustris LANE et al., 1955 P
P. subaspera (as essigi) WIRTH, 1952a P
GROGAN & WIRTH, 1979c P
P. wirthi LANE et al., 1955 P
Paryphoconus angustipennis MAYER, 1952 E

P. flavidus (as lanei) MAYER, 1959 P
P. mayeri MAYER, 1959 P
P. oliveirai WIRTH & RATANAWORABHAN, 1972b P
Stenoxenus johnsoni WIRTH & RATANAWORABHAN, 1972b E

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

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

known for this area and are in the following genera: Leptoconops, Forcipomyia or Culicoides.
Taxon Vector Vertebrate Hosts Location

VIRUSES
Bunyavirus
Ananindeua C. paraensis Birds, marsupials, Brazil
rodents, primates
Oropouche C. paraensis Sloths, humans; antibodies Trinidad, Panama,
in other mammals Brazil, Peru
Utinga Ceratopogonidae Sloths; antibodies in South and Central
other mammals America
Orbivirus
Bluetongue C. insignis,C. filarifer, Wild and domestic New World, Southern
C. pusillus ruminants Europe, Africa, Asia,
Australia
Vesiculovirus
Vesicular stomatitis C. spp. Horses, cattle, pigs, humans New World
New Jersey
Unclassified Arboviruses
Buritirana C. spp. Unknown Brazil
FILARIOIDEA
Dipetalonema caudispina C. hollensis (in lab) Monkeys (Cebidae) South America
Dipetalonema gracile C. hollensis (in lab) Monkeys 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 South
L. bequaerti, C. furens America
C. phlebotomus
Dipetalonema perstans C. spp. Humans, other primates Central and South
America, Africa
Icosiella neglecta F. spp. Frogs South America,
Europe, Asia, Africa
Onchocerca cervicalis C. spp. Equidae Worldwide
Onchocerca gutturosa C. spp. Bovidae Worldwide
2
CAPTURE AND STUDY OF
CERATOPOGONIDAE
Adult biting midges may be collected by any one of the following techniques, with varying
results.
1. Light trap. Although many specialized light traps exist that work very well, a good
sample may even be obtained at a lit window pane when there is good surrounding habitat.
A simple and inexpensive technique is to place a funnel, approximately 20 cm in diameter
above a jar of alcohol, close to a source of light. Another productive method is to lay a UV
light on metal mesh (1 cm square, to keep out larger insects) which is lain over a shallow
pan with soapy water; the water must be screened with fine mesh in the morning and the
specimens preserved in alcohol. Ultraviolet light works well to attract adult midges but
regular florescent or incandescent light may also bring in large numbers. An advantage of
this technique is that it collects both males and females of a given species so that they can
be associated.
2. CO2 trap. These traps collect females of those species that feed on vertebrate hosts
(generally only those of species of Culicoides). A serious limitation of this technique is that
males are not collected and therefore cannot be studied.
3. Sweeping vegetation, especially in areas with wet habitats, and aspirating the adults
from the net. This provides a good method to sample diverse habitats and often results in
species which are not discovered otherwise. This method also results in samples of both
males and females.
4. Malaise traps. These work very well to sample males and females of many species.
Larvae and pupae are obviously restricted to specific habitats and may be sampled
directly from those aquatic, semiaquatic, or moist terrestrial habitats where they are found.
The variety of habitats in which biting midge larvae and pupae may be found is described
above. Sampling methods include the following:
1. Careful examination of wet substrate with a large magnifying glass or under a dis-
secting microscope.
2. Samples of substrate from aquatic habitats (e.g. mud, silt, detritus, leaves) placed in a
pan with additional water. Most Ceratopogoninae larvae swim with a rapid serpentine motion
and these may be removed with an eyedropper. Pupae almost always float at the water sur-
face. Detritus removed from the margins of ponds, streams and lakes and placed in a white
bucket allows for manual removal of pupae as they float. Some workers add salt or sugar to
float larvae and pupae from sample substrate and this is the only practical way to collect those
few species in which the larvae do not swim or the pupae do not float at the surface.
3. Berlese funnels are a good method for extracting larvae from semiaquatic or merely
moist material (e.g. mosses growing on trees in cloud forest).
Once specimens are preserved in 70% alcohol (or some other preservative), most
need to be placed on microscope slides (Fig. 1A) to be identified beyond the generic level.
Methods vary but the following gives excellent results for slide mounting adult specimens
stored in ethyl alcohol and which are not too old (3-4 years or, if stored under dark and
very cold conditions (below 0 C), over 10 years). This method works particularly well for
material to be used for taxonomic studies. A more rapid method is given further below.
Prepare for slide mounting by putting the following solutions into a series of four
stender dishes: 15% acetic acid, 2-propanol, 2-propanol layered over clove oil, clove oil
(each dish about 3/4 full). Each specimen needs to go through this sequence of solutions.
Preparation of five specimens at a time is convenient and manageable, so that 20 dishes are
in use at a time. Placing all the dishes on a thin board so that they can be moved around as
a group on a table top makes it easier to look at specimens while they are in the solutions
and still put all the dishes out of harms way while they are soaking.
A series of slides needs to be ready, by cleaning these well with a cloth or paper towel
and placing a label on each slide. These slides should be placed in a slide tray. In making the
following preparations, round, 10 mm in diameter coverslips work very well.
1. Place the specimen into a dish of ethyl alcohol under the microscope. Remove the
wings by grabbing the very base of the wing with fine forceps. Place both wings in the dish
with 15% acetic acid.
2. For most specimens, separate head and abdomen from thorax. For very small spec-
imens, just tear the membrane between the head and thorax and between the abdomen and
thorax so that all the parts remain attached to each other. Place parts, or torn specimen,
into a three dram vial filled about 1/4 to 1/3 full with 8% KOH. Prepare a total of five
specimens in this way. Each vial needs to be numbered with a grease pencil so that it can be
associated with the wings that are in the acetic acid.
3. Place vials of KOH into a beaker partially filled with water. Place beaker on hot
plate and heat water to boiling point. Specimens will clear after 2-5 minutes, depending on
the size of the specimens. It is important that specimens of a similar size are cooked at
the same time so that they all clear at the same rate.
4. The following step requires some speed because specimens are damaged by re-
maining in hot KOH for too long. Remove the specimen from the vial into a dish with
some 8% KOH in it and put under the microscope to see if it is properly cleared. The
muscles should have dissolved and freely flow out of the specimen if squeezed very, very
gently with the forceps. If the specimens are over-cleared they will make for a poor 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 size
of 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 that
there 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 well
ventilated area and the 2-propanol will evaporate from the mixture.
7. Move specimen to pure clove oil and leave for at least 30 minutes (or longer if
necessary).
8. Place small drop of Canada Balsam on the microscope slide; place the wings in the
drop and cover with a clean coverslip. If Canada Balsam has already hardened too much to
move the wings around easily, add a small drop of xylene, which will make the Canada
Balsam more fluid. Use xylene for the following procedures as well; it is always needed
because Canada Balsam dries faster than the time it takes for a good slide to be made.
9. Place three small drops of Canada Balsam on the microscope slide and put the
head, thorax and abdomen into each. If the head, thorax and abdomen are still partially
attached (small specimens), put specimen into the drop which will contain just the thorax
and remove the head and abdomen; put the head and abdomen into their proper places.
10. Remove the right legs and place all these under a separate coverslip.
11. Arrange the thorax so that the left side is facing upward. Orient the head so that
the front of the head is oriented upward and the antennae are splayed outward. Do not add
a coverslip.
12. Details of the genitalia are often very important in species identification, so that
the following step is a very important one. Arrange the abdomen so that the ventral part of
the genitalia is facing upward. In some males, the genitalia may be only partially turned in
relation to the rest of the abdomen and in this case, sometimes the genitalia will need to be
removed from the abdomen (by tearing with the forceps and a pin) so that the separated
genitalia lies with the ventral side pointed up. The rest of the abdomen should be straight,
with the tergites dorsal, if possible. Do not add a coverslip.
13. Put the slide into the drying oven at 40-50 C. After 1-2 days, the Canada Balsam
with the head, abdomen and thorax will have dried somewhat. Add some more Canada
Balsam to each of these and very gently put a coverslip over each, so they are not crushed.
The partially dried Canada Balsam should keep these parts from being damaged. While the
slide is drying, some of the Canada Balsam may leave a small air bubble under the coverslip.
If so, add a tiny bit of Canada Balsam. Dont worry if the added Canada Balsam traps a
bubble under the coverslip; the bubble nearly always moves out by itself (after some time).
14. Put the slide back into the drying oven. Check for air bubbles for the next several
days and add Canada Balsam as needed. Remove the slide after 1-2 weeks and put into a
slide box. Keep the slide box upright, so the slides are still horizontal.
Larvae and pupae may be treated with the same solutions and should be mounted
dorsal side up. Larvae require no dissection but the operculum of the pupa as well as one
of the respiratory organs should be removed and placed under a separate coverslip (leaving
the Canada Balsam to dry for a day or two and then adding the coverslips). If the larva or
pupa is represented by an exuvia, the specimens obviously do not require clearing and may
be placed directly into 2-propanol and processed from there.
A simpler method for mounting specimens, which is particularly useful for mass mount-
ing material which is likely to include previously described species, is as follows: remove
specimens from 70% ethanol (many can be done as one lot), very briefly blot excess 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 one
or 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 structures
difficult to see clearly. Workers using this method should use a fume hood as both xylene
and phenols are dangerous to human health.
3
REARING CERATOPOGONIDAE
The immatures may be reared using one or more methods. Field collected female adults of
some species may be induced to lay eggs by keeping them in a vial with a loose-fitting lid or
a cotton ball, or in a small Petri dish with wet mud (as described by BORKENT & CRAIG,
2004). In such instances, food for hatching larvae will be somewhat uncertain and, 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 collected
from nature will often molt to the pupal stage if given some of the substrate in which they
were discovered, along with additions of either a rich microbial soup or small prey items,
such as nematodes (depending on the taxon collected). It is always an advantage to collect
mature larvae of either the third or fourth instar to enhance the likelihood of maturation.
Pupae may either be left in a Petri dish or other small container with some of the substrate
in which they were living or with a piece of moist to wet filter paper. Emerging adults
should be kept alive for at least 12-24 hours at room temperature before preservation to
ensure complete hardening of the cuticle and maturation of pigmentation patterns.
4
PHYLOGENETIC RELATIONSHIPS AND
FOSSIL RECORD OF CERATOPOGONIDAE
The phylogenetic and fossil record of the Ceratopogonidae is one of the best understood
among insects and provides a good example of combining cladistic analysis of extant taxa
with interpretation of a detailed fossil record (BORKENT, 2000a; BORKENT & CRAIG, 2004).
The biting midges are an exemplary group for studying evolutionary relationships
because of three outstanding features. First, they are diverse in numbers of species. Gen-
erally, when a group is species-rich, there are intermediate species or groups present which
provide good clues on the direction of evolutionary change of a given structure. Second,
the biting midges have three very different stages (larva, pupa, adult) each of which have
diversified structurally. Unlike many other groups, the immatures of biting midges have
evolved in many different morphological directions. Therefore, it is possible to compare
conclusions about relationships based on one stage against the conclusions based on a
different stage. Third, the biting midges have an incredible fossil record in amber that has
been investigated in some detail. We now have 11 major deposits ranging from 17 to 121
million years of age, with thousands of specimens studied. The fossil biting midges are
largely in excellent condition and most taxonomic features can be studied in detail. These
fossils provide an excellent gauge as to the accuracy of our interpretation of evolutionary
relationships based on living species.
General conclusions show that the Leptoconopinae are the sister group of all remain-
ing Ceratopogonidae, the Forcipomyiinae and Dasyheleinae are sister groups, and that the
preceding 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 relationships
between most of the remaining genera of the Ceratopogoninae.
5
ZOOGEOGRAPHY
Our understanding of the zoogeography of Ceratopogonidae is rather limited. This is for
two important reasons. One is that the phylogenetic relationships between many taxa are
unknown and it is therefore impossible to interpret the significance of their distributions in
light of those relationships. Secondly, in the Neotropical Region, large areas have not been
sampled, including much of the Amazon Basin and nearly all of the Andes. Without doubt,
there remain a great number of species to be discovered.
Nevertheless, there are some broad geographic patterns which are apparent. Most
notable is the distinctive fauna of Patagonia. The following genera are endemic to the area,
with most of these present in forested habitats: Borkenthelea SPINELLI & GROGAN (three
forest species), Diaphanobezzia INGRAM & MACFIE (four species, three from forest, one from
steppe), Isthmohelea INGRAM & MACFIE (monotypic, from forest), Notiohelea GROGAN & WIRTH
(two forest species), Physohelea GROGAN & WIRTH (two forest species), Austrosphaeromias
SPINELLI (two forest species). In addition, the following are restricted within the 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; six
others from Australia or New Zealand), Macrurohelea INGRAM & MACFIE (13 species (two
undescribed) from Patagonian forest, except one species that extends northward to arid
zones of Chile and Argentina, one undescribed species in the Somuncura Plateau in the
Patagonian steppe; three described and two unnamed from Australia), and Paradasyhelea
MACFIE (three Patagonian species, two from forest and one from steppe; seven others from
New Caledonia, Australia, New Zealand; one Nearctic species). The latter three genera
may have southern hemisphere connections that reflect patterns of continental drift but
their phylogenetic relationships await analysis to test this possibility.
Study of other ceratopogonids in Patagonia indicates that most species are endemic
in this region including species of Forcipomyia, Dasyhelea, and Stilobezzia (all of which are
members of the subgenus Acanthohelea KIEFFER and all restricted to Nothofagus forest).
In addition to those endemic Patagonian genera discussed above, several genera or
subgenera of Ceratopogonidae are restricted or nearly restricted to the Neotropical Re-
gion: Forcipomyia (Rhynchoforcipomyia WIRTH & DOW) (seven species), Culicoides (Anilomyia
VARGAS) (17 species), C. (Cotocripus BRTHES) (five species), C. (Macfiella FOX) (two species),
C. (Mataemyia VARGAS) (15 species), C. (Psychophaena PHILIPPI) (two species), Baeodasymyia
CLASTRIER & RACCURT (five species), Baeohelea WIRTH & BLANTON (one species), Bahiahelea
WIRTH (one species), Cacaohelea WIRTH & GROGAN (two species), Leptohelea WIRTH & BLAN-
TON (one species), Parastilobezzia WIRTH & BLANTON (one species), Echinohelea MACFIE (Echi-
noideshelea WIRTH) (one species), Fittkauhelea WIRTH & BLANTON (one species), Schizonyxhelea
CLASTRIER (two species), Clinohelea KIEFFER (Ceratobezzia KIEFFER) (one species), Groganhelea
SPINELLI & DIPPOLITO (one species), Lanehelea WIRTH & BLANTON (two species), Neobezzia
WIRTH & RATANAWORABHAN (eight species), Sphaerohelea SPINELLI & FELIPPE-BAUER (one
species), Clastrieromyia SPINELLI & GROGAN (four species), Pachyhelea WIRTH (one of the two
species extends north into Texas), Amerohelea GROGAN & WIRTH (one of the 11 known
species extends north into Texas), Paryphoconus ENDERLEIN (one of 40 species is Nearctic).
Several of these groups are clearly monophyletic and this indicates that they have speciated
within the Neotropical Region (i.e. Baeodasymyia + Baeohelea, Clastrieromyia, Paryphoconus).
A few of these genera, each with only 1-2 species, are endemic to Central America
and northern South America: Cacaohelea (Costa Rica), Leptohelea (Costa Rica, Colombia),
Parastilobezzia (Costa Rica, Colombia).
In Central America, the genus Ceratoculicoides WIRTH & RATANAWORABHAN is present in
tropical Mexico (one described species) and Costa Rica (one undescribed species) and this
is likely a southern extension of an otherwise Holarctic genus.
Within Costa Rica we recognize a pattern, common in many groups of organisms,
that lowland species tend to be broadly distributed in the Neotropical Region and those
that occur higher than 1500 meters in elevation tend to be more restricted in distribution
and are often endemic to Costa Rica or Costa Rica and Panama. This general pattern will
almost certainly be true in other high elevation areas in the Neotropics once they are inves-
tigated more thoroughly.
6
CLASSIFICATION
Ceratopogonidae belong to the superfamily Chironomoidea and are the sister group of the
Chironomidae. Worldwide, extant Ceratopogonidae are divided into four well established
subfamilies, based on cladistic relationships (BORKENT, 2000a; BORKENT & CRAIG, 2004):
Leptoconopinae (2 genera), Forcipomyiinae (2 genera), Dasyheleinae (1 genus) and Cer-
atopogoninae (104 genera). All subfamilies are known from the Neotropical Region (Table
4). REMM (1975) proposed recognizing the Forcipomyiinae and Dasyheleinae as tribes in
the subfamily Forcipomyiinae and, as sister groups, this is conceivable but has not yet been
widely accepted.
The Ceratopogoninae, which make up the bulk of the family both generically and at
the species level, is divided into 6 tribes: Culicoidini (3 genera), Ceratopogonini (57 gen-
era), Heteromyiini (8 genera), Sphaeromiini (28 genera), Palpomyiini (6 genera) and the
Stenoxenini (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, the
Ceratopogonini are paraphyletic in relation to the remaining tribes and that the Palpomyi-
ini and Stenoxenini form a monophyletic group. The relationships among most of the
genera 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 is
clear that many more remain unnamed. It is certainly reasonable to suggest that at least
15,000 species exist on our planet. Within the Neotropical Region, the Andes are virtually
uncollected, in spite of the fact that Ceratopogonidae are abundant and diverse there
(BORKENT, pers. obs.). Many areas of Brazil, Peru, northern Chile and northwestern Argen-
tina are also particularly poorly known. Limited portions of the tropical rain forests of
Central America and Amazonia were extensively collected during the 1960s and 1970s,
especially by F.S. BLANTON and E. FITTKAU, respectively, providing the basis for numerous
taxonomic papers mainly by W.W. WIRTH and his coworkers. Nevertheless, recent collec-
tions of biting midges in Costa Rica and Peruvian Amazonia reveal the presence of many
more 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 most
taxonomic attention, has many undescribed species and its subgeneric classification is in
great need of analysis.
Table 4. Catalog of the Ceratopogonidae of the Neotropical Region. The names of those
species of Leptoconops and Culicoides which are known to bite humans are underlined.
SUBFAMILY LEPTOCONOPINAE NO, 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 NO, 1905: 114. Type species: Mycterotypus bezzii NO, designation by
CARTER, 1921: 3.
PROTERSESTHES KIEFFER, 1921c: 107. Type species: Tersesthes brasiliensis LUTZ, by original
designation.
REFERENCES: RONDEROS, 1988 (key to females from Argentina); RONDEROS & SPINELLI,
1992 (key to females); WIRTH & ATCHLEY, 1973 (key to subgenera of females, other
than Proleptoconops; key to Nearctic females).
Subgenus BRACHYCONOPS WIRTH & ATCHLEY

BRACHYCONOPS WIRTH & ATCHLEY, 1973: 15 (as subgenus of Leptoconops). Type species:
Leptoconops californiensis WIRTH & ATCHLEY, by original designation.
patagoniensis RONDEROS, 1990b: 423 (female; Argentina). Distr.- Southern Argentina.
Subgenus HOLOCONOPS KIEFFER

HOLOCONOPS KIEFFER, 1918: 135. Type species: Leptoconops kerteszi KIEFFER, by original
designation.
MICROCONOPS KIEFFER, 1921c: 108. Type species: Microconops vexans KIEFFER, by original
designation.
REFERENCE: CLASTRIER & WIRTH, 1978 (key to Nearctic males and females).
bassoi RONDEROS & SPINELLI, 1993: 305 (male; Chile). Distr.- Northern Chile.
bequaerti (KIEFFER), 1925: 405 (Holoconops, as becquaerti; female; Honduras). Distr.- USA
(Texas) to Panama and Trinidad.
hondurensis HOFFMAN, 1926: 135. Honduras.
knowltoni CLASTRIER & WIRTH, 1978: 26 (male, female; USA, California). Distr.- USA
(western) south to Mexico (Sinaloa), Brazil (Santa Catarina).
Subgenus LEPTOCONOPS SKUSE

brasiliensis (LUTZ), 1913: 66 (Tersesthes; female; Brazil). Distr.- Argentina (Salta), Brazil
(Amazonas), Uruguay.
casali CAVALIERI & CHIOSSONE, 1966: 45 (female; Argentina). Distr.- Argentina (Salta, San
Luis), Uruguay.
chilensis FORATTINI, 1958: 38 (female; Chile). Distr.- Bolivia, Chile, Argentina (Jujuy, Salta,
La Pampa).
panamensis RONDEROS & SPINELLI, 1993: 308 (male; Panama). Distr.- Panama.
petrocchiae SHANNON & DEL PONTE, 1927: 734 (female; Argentina). Distr.- Northwestern
Argentina.
ricardoi RONDEROS & SPINELLI, 1992: 43 (female; Argentina). Distr.- Southwestern Argentina.
venezuelensis ORTZ, 1952a: 165 (female; Venezuela). Distr.- Venezuela.
Subgenus MEGACONOPS WIRTH & ATCHLEY

MEGACONOPS WIRTH & ATCHLEY, 1973: 18 (as subgenus of Leptoconops). Type species:
Leptoconops floridensis WIRTH, by original designation.
floridensis WIRTH, 1951a: 282 (female; USA, Florida). Distr.- USA (Florida), Jamaica, Colombia.
SUBFAMILY FORCIPOMYIINAE LENZ, 1934: 96

Genus ATRICHOPOGON KIEFFER
ATRICHOPOGON KIEFFER, 1906a: 53 (as subgenus of Ceratopogon). Type species:
Ceratopogon exilis COQUILLETT (= Ceratopogon levis COQUILLETT), designation by COQUILLETT,
1910: 512.
REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE, 1931
(key males and females from Patagonia and southern Chile); MACFIE, 1939b (key to
males and females from the Neotropical Region).
Subgenus ATRICHOPOGON KIEFFER

adamsoni MACFIE, 1937a: 4 (female; Trinidad). Distr.- Mexico (Chiapas), Trinidad.
albinensis INGRAM & MACFIE, 1931: 228 (female; Argentina). Distr.- Argentina (Buenos Aires).
altivolans MACFIE, 1949: 113 (female; Mexico). Distr.- Mexico (Chiapas).
aridus SPINELLI & MARINO, in SPINELLI et al., 2006: 304 (male, female; Argentina). Distr.-
Argentina (Mendoza, Rio Negro).
asuturus BORKENT & PICADO, 2004: 29 (male; Costa Rica). Distr.- Costa Rica.
auricoma KIEFFER, 1917b: 300 (male, female; Colombia). Distr.- Colombia.
balseiroi SPINELLI, 1982: 206 (male, female; Argentina). Distr.- Argentina (Buenos Aires).
barbatus BORKENT & PICADO, 2004: 23 (male; Costa Rica). Distr.- Costa Rica.
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 (Crdoba, 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 (Maranho, Santa Catarina).
gamboai BORKENT & PICADO, 2004: 26 (male; Costa Rica). Distr.- Costa Rica.
glaber MACFIE, 1935a: 50 (female; Brazil). Distr.- Costa Rica, Trinidad, Brazil (Par,
Maranho).
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 (Maranho), 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 (ORTZ), 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 DELCOLLE & 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.-
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 (Maranho).
utricularis MACFIE, 1953: 102 (female; Costa Rica). Distr.- Costa Rica.
warmkei WIRTH, 1956c: 243 (male, female; Puerto Rico). Distr.- USA (Florida, Puerto Rico).
woodfordi MACFIE, 1938: 162 (female; Trinidad). Distr.- Trinidad.
yolancae BORKENT & PICADO, 2004: 24 (male; Costa Rica). Distr.- Costa Rica.
Subgenus LOPHOMYIDIUM CORDERO

LOPHOMYIDIUM CORDERO, 1929: 94. Type species: Lophomyidium uruguayense CORDERO,
by original designation.
REFERENCE: WIRTH 1994a (key to Nearctic males and females).
archboldi WIRTH, 1994a: 27 (male, female; USA, Florida). Distr.- USA (California to Florida)
to Argentina.
fusculus (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, New Jersey). Distr.- Northern
Eurasia, North America, Central America and Caribbean to Bolivia and Brazil (Par).
uruguayensis (CORDERO), 1929: 95 (Lophomyidium; male; Uruguay). Distr.- Paraguay, eastern
Argentina, Uruguay.
Genus FORCIPOMYIA MEIGEN

FORCIPOMYIA MEIGEN, 1818: 73, 75. Type species: Tipula bipunctata Linnaeus, designation
by WESTWOOD, 1840: 126. Generic name first published in synonymy with Ceratopogon
but available under ICZN Code Article 11(e).
TETRAPHORA PHILIPPI, 1865: 630. Type species: Tetraphora fusca PHILIPPI, by monotypy.
REFERENCES: DEBENHAM, 1987 (key to males and females to subgenera from Australasian
Region which includes all Neotropical subgenera other than Rhynchoforcipomyia and
Saliohelea); INGRAM & MACFIE, 1931 (key to males and females from Patagonia and
southern Chile); WIRTH & RATANAWORABHAN, 1978 (key to females to subgenera; their
genus A near Lepidohelea = Pedilohelea).
Subgenus BLANTONIA WIRTH & DOW

BLANTONIA WIRTH & DOW, 1971: 289 (as subgenus of Forcipomyia). Type species:
Forcipomyia caribbea WIRTH & DOW, by original designation.
caribbea WIRTH & DOW, 1971: 291 (male, female; Jamaica). Distr.- USA (Florida) to Trinidad,
Mexico (Yucatn), 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 (Par, 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
(Par, So 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, Par).
nigrescens MACFIE, 1939b: 156 (male, female; Brazil). Distr.- Panama, Brazil (So Paulo,
Rio de Janeiro, Santa Catarina).
remigera UTMAR & WIRTH, 1976: 121 (male, female; Colombia). Distr.- Panama, Colombia,
French Guiana, Brazil (Par).
sabalitensis UTMAR & WIRTH, 1976: 127 (male, female; Costa Rica). Distr.- Honduras to
Colombia, Brazil (Paran, Santa Catarina), northeastern Argentina.
varicolor SAUNDERS, 1957: 683 (larva, pupa, male; Brazil). Distr.- Brazil (Rio de Janeiro).
Subgenus EUPROJOANNISIA BRTHES

EUPROJOANNISIA BRTHES, 1914: 155. Type species: Euprojoannisia platensis BRTHES,
REFERENCES: BYSTRAK & WIRTH, 1978 (key to Nearctic larvae, pupae, males, females);
HUERTA, 1996 (key to males and females from Mexico).
blantoni SORIA & BYSTRAK, 1975: 3 (larva, pupa, male, female; Brazil). Distr.- Eastern USA
to Ecuador, Brazil (Bahia).
bromeliae SAUNDERS, 1957: 665 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro).
calcarata (COQUILLETT), 1905: 64 (Ceratopogon; male; Mexico). Distr.- Eastern USA, Mexico
(Tabasco, Quintana Roo).
dowi BYSTRAK & WIRTH, 1978: 21 (male, female; USA, Florida). Distr.- USA (Florida),
Mexico (Quintana Roo).
esteparia MARINO & SPINELLI, 2001b: 12 (male, female; Argentina). Distr.- Southern Argentina.
falcifera SAUNDERS, 1959: 39 (larva, pupa, male, female; Trinidad). Distr.- Venezuela, Trinidad.
galliarii MARINO & SPINELLI, 1999a: 5 (female; Argentina). Distr.-. Argentina (Misiones).
longispina SAUNDERS, 1957: 669 (larva, pupa, male, female; Brazil). Distr.- Brazil (Rio de Janeiro).
lota (WILLISTON), 1896: 282 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.
mortuifolii SAUNDERS, 1959: 35 (larva, pupa, male, female; Trinidad). Distr.- USA (Florida)
to St. Lucia.
navaiae BYSTRAK & WIRTH, 1978: 31 (male, female; USA, Florida). Distr.- USA (Florida),
Mexico (Yucatn).
platensis (BRTHES), 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, So
Paulo), Venezuela.
unica BYSTRAK & WIRTH, 1978: 44 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas.
Subgenus FORCIPOMYIA MEIGEN

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

argenteola MACFIE, 1939b: 146 (female; Brazil). Distr.- Brazil (Rio de Janeiro, Santa Catarina).
calatheae WIRTH, 1982a: 573 (larva, pupa, male, female; Dominica). Distr.- Colombia,
Dominica, Brazil (Bahia, Santa Catarina). .
marini SPINELLI & DIPPOLITO, 1995: 155 (male, female; Venezuela). Distr.- Venezuela.
quatei WIRTH, 1952a: 142 (male, female; USA, California). Distr.- USA (California to Florida)
to Brazil (Bahia).
uramaensis SPINELLI & DIPPOLITO, 1995: 156 (female; Venezuela). Distr.- Venezuela.
youngi WIRTH, 1982a: 579 (larva, pupa, male, female; Panama). Distr.- Costa Rica, Panama,
Ecuador.
fusca Species Group

caliginosella WIRTH, 1974: 5. New name for caliginosa. Distr.- Southwestern Argentina.
caliginosa INGRAM & MACFIE, 1931: 167 (preoccupied by Forcipomyia caliginosa (INGRAM &
MACFIE), 1924). Argentina.
fusca (PHILIPPI), 1865: 630 (Tetraphora; female; Chile). Distr.- Central and southern Chile
south to Deceit Island, southwestern Argentina south to Tierra del Fuego.
patagonica INGRAM & MACFIE, 1931: 165. Argentina.
piroskyi CAVALIERI, 1961b: 172 (female; Argentina). Distr.- Argentina (Tierra del Fuego).
wygodzinskyi CAVALIERI, 1961a: 17 (female; Argentina). Distr.- Argentina (Tierra del Fuego).
delpontei CAVALIERI, 1961b: 169. Argentina.
genualis Species Group

genualis (LOEW), 1866: 128 (Ceratopogon; male; Cuba). Distr.- USA (Louisiana to Florida),
Bahamas, Mexico (Yucatn, Chiapas) to Colombia, Galpagos Islands, Cayman Islands,
Cuba to Argentina (Buenos Aires province); Sao Tom.
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, Par, 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,
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 (Santa
Catarina).
poulaineae INGRAM & MACFIE, 1931: 227 (male; Argentina). Distr.- Brazil (Bahia, Santa
Catarina), Argentina (Buenos Aires).
somuncurensis MARINO & SPINELLI, 2001a: 116 (male, female; Argentina). Distr.- Argentina
(Rio Negro).
townsendi KNAB, 1915: 111 (male, female; Peru). Distr.- Peru.
utae KNAB, 1915: 109 (female; Peru). Distr.- Peru.
yamana MARINO & SPINELLI, 2001a: 118 (female; Chile). Distr.- Chile (Deceit Island).
zonogaster INGRAM & MACFIE, 1931: 168 (male; Chile). Distr.- Southern Chile.
squamitibia Species Group

catarinensis MARINO & SPINELLI, 2002: 309 (male, female; Brazil). Distr.- Mexico, Costa
Rica, Colombia, Brazil.
chilensis (PHILIPPI), 1865: 601 (Ceratopogon; male; Chile). Distr.- Central and southern Chile,
pinamarensis SPINELLI, 1983a: 121 (male, female; Argentina). Distr.- Costa Rica, Panama,
Venezuela, Brazil (Par, 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 Fernndez Islands). Distr.- Juan Fernndez
Islands.
Unplaced to Species Group

flava (WILLISTON), 1896: 280 (Ceratopogon; male; St. Vincent). Distr.- St. Vincent.
pictoni MACFIE, 1938: 161 (male; Trinidad). Distr.- USA (Louisiana to Florida) to Brazil
(Amazonas, Par, Bahia, So 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 & IBAEZ-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 (So Paulo).
guarani RONDEROS & SPINELLI, 1999: 153 (female; Paraguay). Distr.- Paraguay.
insigniforceps MACFIE, 1939b: 165 (male; Brazil). Distr.- Brazil (Santa Catarina).
intermedia SAUNDERS, 1964: 468 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica.
multidentata RONDEROS & SPINELLI, 1999: 153 (female; Brazil). Brazil (Bahia, Itaguipe,
Fazenda Almirante).
quasicornuta SAUNDERS, 1964: 467 (larva, pupa, male, female; Costa Rica). Distr.- Costa Rica.
saltensis (CAVALIERI), 1962: 360 (Lasiohelea; female; Argentina). Distr.- Northwestern Argentina.
stylifer (LUTZ), 1913: 63 (Centrorhynchus; female; Brazil). Distr.- Belize to Ecuador, Trinidad,
Venezuela to northeastern Argentina.
Subgenus LEPIDOHELEA KIEFFER

LEPIDOHELEA KIEFFER, 1917b: 364. Type species: Ceratopogon chrysolophus KIEFFER, by
original designation.
REFERENCES: WIRTH, 1991c (key to species groups; key to males and females of bicolor
group); WIRTH & SPINELLI, 1992a (males and females of seminole subgroup); WIRTH &
SPINELLI, 1993a (key to males and some females of annulatipes group).
abercrombyi MACFIE, 1938: 161 (male, female; Trinidad). Distr.- Trinidad.

annulatipes MACFIE, 1939b: 154 (male; Brazil). Distr.- Colombia, Brazil (Par, 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 previous
type designation).
PHASMIDOHELEA MAYER, 1937: 233. Type species: Phasmidohelea crudelis MAYER
(= Ceratopogon fuliginosus MEIGEN), by original designation.
REFERENCES: CLASTRIER & WIRTH, 1995 (key to females in ixodoides group); HUERTA, 1996
(key to males and females from Mexico); WIRTH, 1972 (key to females in fuliginosa group).
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 (Par).
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 (Par).
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 (Par).
donskoffi CLASTRIER & WIRTH, 1995: 125 (female; French Guiana). Distr.- French Guiana.
dunklei CLASTRIER & WIRTH, 1995: 130 (female; Peru). Distr.- Peru.
eriophora (WILLISTON), 1896: 279 (Ceratopogon; male; St. Vincent). Distr.- USA (Florida),
Caribbean, Belize to Brazil (Bahia).
felippebauerae CLASTRIER & WIRTH, 1995: 122 (female; Peru). Distr.- Peru.
grandcolasi CLASTRIER & WIRTH, 1995: 112 (female; French Guiana). Distr.- French Guiana.
franklini CLASTRIER & WIRTH, 1995: 141 (female; Costa Rica). Distr.- Costa Rica, Panama.
fuliginosa (MEIGEN), 1818: 86 (Ceratopogon; Germany). Distr.- Widespread in the Palaearctic,
Oriental, Afrotropical, Nearctic and Neotropical regions. In the Neotropics south to
Argentina and Uruguay.
crudelis KNAB, 1914: 66 (preoccupied by Forcipomyia crudelis (KARSCH), 1886). Mexico.
tropica (KIEFFER), 1917b: 297 (Ceratopogon). Costa Rica.
galapagensis (COQUILLETT), 1901b: 372 (Ceratopogon; male; Galpagos Islands). Distr.-
Galpagos 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 (Par).
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, Par).
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 (Par).
valleensis CLASTRIER & WIRTH, 1995: 145 (female; Colombia). Distr.- Colombia.
vernoni CLASTRIER & WIRTH, 1995: 138 (female; Colombia). Distr.- Colombia.
wagneri (SGUY), 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 (Par).
archboldi DE MEILLON & WIRTH, 1980: 13 (male, female; Dominica). Distr.- Jamaica, Puerto
Rico, Brazil (Par).
raposoi DE MEILLON & WIRTH, 1980: 21 (male; Colombia). Distr.- Colombia.
spangleri DE MEILLON & WIRTH, 1980: 21 (male, female; Guatemala). Distr.- Mexico
(Yucatn), Guatemala, Panama, Colombia, Ecuador.
spilmani DE MEILLON & WIRTH, 1980: 22 (male, female; Dominica). Distr.- Panama,
Colombia, Dominica, Trinidad, Brazil (Santa Catarina).
Subgenus PHYTOHELEA REMM

PHYTOHELEA REMM, 1971: 189 (as subgenus of Forcipomyia). Type species: Ceratopogon
bromelicola LUTZ, by original designation.
REFERENCE: DE MEILLON & WIRTH, 1979 (key to females).

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 & DELCOLLE, 1994: 51 (male, female; French Guiana). Distr.- French Guiana.
oligarthra SAUNDERS, 1957: 698 (larva, pupa, male, female; Puerto Rico). Distr.- USA (Florida,
Puerto Rico), Cuba, Dominica, Mexico, Guyana, also from Singapore and Micronesia.
Subgenus PTEROBOSCA MACFIE

PTEROBOSCA MACFIE, 1932: 266. Unavailable name; proposed after 1930 without type
species designation.
PTEROBOSCA MACFIE, 1940f: 16. Type species: Ceratopogon aeschnosuga DE MEIJERE, by
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 (Par).
messersmithi WIRTH & DOW, 1972: 864 (male, female; El Salvador). Distr.- Mexico
(Veracruz), Central America to Colombia, Jamaica.
puracensis WIRTH & DOW, 1972: 867 (female; Colombia). Distr.- Colombia.
zeteki WIRTH & DOW, 1972: 870 (female; Panama). Distr.- Panama, Colombia.
Subgenus SALIOHELEA WIRTH & RATANAWORABHAN

SALIOHELEA WIRTH & RATANAWORABHAN, 1978: 494 (as subgenus of Forcipomyia). Type
species: Forcipomyia leei WIRTH & RATANAWORABHAN, by original designation.
leei WIRTH & RATANAWORABHAN, 1978: 498 (male, female; Colombia). Distr.- Eastern USA
and Caribbean to southern Brazil (Amazonas, Par, Bahia, Rio de Janeiro).
Subgenus SCHIZOFORCIPOMYIA CHAN & LEROUX

SCHIZOFORCIPOMYIA CHAN & LEROUX, 1971: 271 (as subgenus of Forcipomyia). Type
species: Forcipomyia petersoni CHAN & LEROUX ( = Forcipomyia borbonica CLASTRIER), by
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 (Yucatn).
sanctaeclarae WIRTH, 1952b: 90 (male, female; Juan Fernndez Islands). Distr.- Juan
Fernndez Islands, central Chile, northwestern Argentina.
soibelzoni MARINO & SPINELLI, 2001c: 14 (female; Argentina). Distr.- Argentina (Rio Negro).
tenuiforceps MACFIE, 1939b: 167 (male; Brazil). Distr.- USA (California, New Mexico) to
Brazil (Santa Catarina).
Subgenus THYRIDOMYIA SAUNDERS

THYRIDOMYIA SAUNDERS, 1925: 268. Type species: Thyridomyia palustris SAUNDERS
(= Ceratopogon monilicornis COQUILLETT), by original designation.
REFERENCE: WIRTH, 1970 (key to males and females).
calchaqui SPINELLI & MARINO, 1997: 188 (female; Argentina). Distr.- Northwestern
Argentina.
colombiae WIRTH, 1970: 435 (male, female; Colombia). Distr.- Mexico (Veracruz), Costa
Rica, Panama, Trinidad, Colombia.
jamaicensis WIRTH, 1970: 436 (male, female; Jamaica). Distr.- Jamaica, Puerto Rico.
jipajapae WIRTH, 1970: 437 (male, female; Panama). Distr.- Costa Rica to Ecuador, Trinidad.
nana (MACFIE), 1939b: 171 (Lasiohelea; female; Brazil). Distr.- Mexico (Veracruz), Jamaica
through Caribbean to southern Brazil (Santa Catarina).
nodosa SAUNDERS, 1959: 43 (larva, pupa, male, female; Costa Rica). Distr.- USA (Arizona,
Kansas, Mississippi), Mexico (Baja California, Sonora, Sinaloa, San Luis Potosi), Costa
Rica, Colombia.
riojana SPINELLI & MARINO, 1997: 188 (male; Argentina). Distr.- Western and southern Argentina.
sinuosa DOW & WIRTH, 1972: 189 (male, female; Mexico). Distr.- Mexico (Sinaloa) (possibly
Iowa in USA).
univesicula MACFIE, 1939b: 170 (female; Brazil). Distr.- Brazil (Santa Catarina).
Subgenus TRICHOHELEA GOETGHEBUER

APELMA KIEFFER, 1919a: 64 (preoccupied by BILLBERG, 1820). Type species: Apelma auronitens
KIEFFER (= Trichohelea tonnoiri GOETGHEBUER), designation by MACFIE, 1940f: 16.
TRICHOHELEA GOETGHEBUER, 1920: 39. Type species: Trichohelea tonnoiri GOETGHEBUER,
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 (So Paulo).
roubaudi CLASTRIER & DELCOLLE, 1997: 379 (female; French Guiana). Distr.- French Guiana.
sayhuequei MARINO & SPINELLI, 2004b: 2257 (female; Argentina). Distr.- Southwestern
Argentina, southern Chile.
tehuelche MARINO & SPINELLI, 2004b: 2260 (male, female; Argentina). Distr.- Southwestern
Argentina.
trinidadensis SAUNDERS, 1964: 476 (pupa, female; Trinidad). Distr.- Trinidad.
Subgenus WARMKEA SAUNDERS

WARMKEA SAUNDERS, 1957: 671 (as subgenus of Forcipomyia). Type species: Forcipomyia
bicolor SAUNDERS (= Forcipomyia lesliei WIRTH), by original designation.
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 (Par).
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 (Par, Maranho, 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 (Par, 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, Par), Argentina
(Buenos Aires).
SUBFAMILY DASYHELEINAE LENZ, 1934: 96

Genus DASYHELEA KIEFFER
DASYHELEA KIEFFER, 1911c: 5. Type species: Dasyhelea halophila KIEFFER, by monotypy.
PSEUDOCULICOIDES M ALLOCH , 1915a: 309. Type species: Ceratopogon mutabilis
COQUILLETT, by original designation.
REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM & MACFIE,
1931 (key males and females from Patagonia and southern Chile); SPINELLI & WIRTH,
1984d (key to males and females of cincta group).
aegealitis SPINELLI & WIRTH, 1984d: 589 (male, female; Jamaica). Distr.- USA (Florida), Jamaica.
albopicta INGRAM & MACFIE, 1931: 187 (male, female; Argentina). Distr.- Southern Chile,
ancora (COQUILLETT), 1902: 87 (Ceratopogon; female; USA, Florida). Distr.- USA (California,
Arizona, New Mexico, Connecticut to Florida), Galpagos 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, Galpagos 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 (Yucatn), 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, Galpagos Islands, Brazil (Bahia).
cacaoi WIRTH & WAUGH, 1976: 231 (male, female; Trinidad). Distr.- Panama, Trinidad.
calvescens MACFIE, 1938: 157 (male, female; USA, Hawaii). Distr.- Mexico (Baja California,
Sonora), Panama, Hawaii.
caribbeana SPINELLI & WIRTH, 1984d: 596 (female; Jamaica). Distr.- Mexico (Quintana
Roo, Chiapas), Panama, Jamaica, Haiti.
chilensis INGRAM & MACFIE, 1931: 186 (female; Chile). Distr.- Southern Chile
cincta (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Florida). Distr.- USA (California
to New York to Florida) to Brazil and Argentina.
penthesileae MACFIE, 1935a: 55. Brazil.
columbiana (KIEFFER), 1917b: 304 (Culicoides; female; Colombia). Distr.- Colombia.
correntina RONDEROS & DAZ, 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 DELCOLLE & RIEB, 1994: 272 (male, female; Guadeloupe). Distr.-
Guadeloupe.
hippolytae MACFIE, 1935a: 54 (female; Brazil). Distr.- Brazil (Maranho).
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 DELCOLLE & 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), Galpagos
Islands.
necrophila SPINELLI & RODRGUEZ, 1999: 59 (male, female, pupa, larva; Argentina). Distr.-
Mexico (Coahuila), Argentina (Buenos Aires).
paracincta WIRTH, 1969: 576 (male, female; Galpagos Islands). Distr.- Galpagos 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 (So 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 (So 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; Galpagos Islands). Distr.- Galpagos Islands.
soriai WIRTH & WAUGH, 1976: 229 (male, female; Brazil). Distr.- Brazil (Bahia).
spathicerca WIRTH, 1969: 577 (male, female; Galpagos Islands). Distr.- Galpagos Islands.
suarezi SPINELLI & RONDEROS, 1987: 11 (male, female; Argentina). Distr.- Argentina (Buenos
Aires).
thalestris MACFIE, 1935a: 55 (female; Brazil). Distr.- Trinidad, Guyana, Brazil (Maranho).
villosipes (KIEFFER), 1917b: 305 (Culicoides; male; Paraguay). Distr.- Paraguay.
viridans FORATTINI & RABELLO, 1957: 247 (male; Brazil). Distr.- Brazil (Rio de Janeiro).
williamsi WIRTH & WAUGH, 1976: 228 (male, female; Trinidad). Distr.- Panama, Dominica,
Trinidad, Brazil (Bahia).
winderi WIRTH & WAUGH, 1976: 233 (male, female; Brazil). Distr.- Brazil (Bahia).
SUBFAMILY CERATOPOGONINAE NEWMAN, 1834: 388

TRIBE CULICOIDINI KIEFFER, 1911c: 1, 1911b: 319
Genus CULICOIDES LATREILLE
CULICOIDES L ATREILLE , 1809: 251. Type species: Culicoides punctatus L ATREILLE
(= Ceratopogon punctatus MEIGEN), by monotypy.
PADROSIA RAFINESQUE, 1815: 130 (unnecessary replacement name for Culicoides LATREILLE).
Type species: Culicoides punctatus LATREILLE (= Ceratopogon punctatus MEIGEN), automatic.
REFERENCES: AITKEN et al., 1975 (key to females from Trinidad and Tobago); BLANTON
& WIRTH, 1979 (key to larvae, males and females from Florida (USA)); FORATTINI,
1957 (key to females of all species); HUERTA, 1996 (key to females from Mexico);
ORTZ & LEN, 1955 (key to females from Ecuador); RONDEROS, 1988 (key to females
from Argentina); RONDEROS & SPINELLI, 1998 (key to males and females from the
Yacyret Dam, between Paraguay and Argentina); SPINELLI, RONDEROS, DAZ & 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 and
some males from Panama); WIRTH & BLANTON 1973 (key to females and some males
from Amazon Basin); WIRTH & BLANTON, 1974b (key to females and some males
from the Caribbean); WIRTH, DYCE & SPINELLI, 1988 (some meristic character states
and photographs of wings of all species).
There is no reliable key to the subgenera and species groups of Neotropical Culicoides. For
species outside of local areas for which revisions are available (as listed above), the
easiest guide to species identification is that of WIRTH, DYCE & SPINELLI (1988). From
there readers may refer to keys to subgenera or species groups referred to below.
Subgenus AMOSSOVIA GLUKHOVA

AMOSSOVIA GLUKHOVA, 1989: 226 (as subgenus of Culicoides). Type species: Ceratopogon
dendrophilus Amosova, by original designation.
REFERENCE: WIRTH & BLANTON, 1967 (key to males and females)
oklahomensis KHALAF, 1952: 355 (as subspecies of villosipennis; male; USA, Oklahoma).
Distr.- USA (California to Mississippi) to Guatemala.
Subgenus ANILOMYIA VARGAS

ANILOMYIA VARGAS, 1960: 37 (as subgenus of Culicoides). Type species: Culicoides covagarciai
ORTZ, 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 ORTZ, 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 obsoletus
MEIGEN, by original designation.
REFERENCES: RODRGUEZ & 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
suarezi RODRGUEZ & WIRTH, 1986: 313 (female; Colombia). Distr.- Colombia.
Subgenus BELTRANMYIA VARGAS

BELTRANMYIA VARGAS, 1953a: 34 (as subgenus of Culicoides). Type species: Culicoides
crepuscularis MALLOCH, by original designation.
REFERENCE: BLANTON & WIRTH, 1979 (males and females included in key to species
from Florida).
bermudensis WILLIAMS, 1956: 298 (female; Bermuda). Distr.- USA (New York to Florida
and Texas), Bermuda.
crepuscularis MALLOCH, 1915a: 303 (male, female; USA, Illinois). Distr.- Canada (British
Columbia to Nova Scotia) to Costa Rica.
knowltoni BECK, 1956: 136 (male, female; USA, Florida). Distr.- Bahamas, USA (Florida).
Subgenus COTOCRIPUS BRTHES

COTOCRIPUS BRTHES, 1912: 451. Type species: Cotocripus caridei BRTHES, by monotypy.
bambusicola LUTZ, 1913: 62 (female; Brazil). Distr.- Colombia, Venezuela, Brazil (Bahia,
Esprito Santo, Rio de Janeiro, So Paulo), Argentina (Misiones, Buenos Aires).
bahiensis BARBOSA, 1947: 11. Brazil.
caridei (BRTHES), 1912: 451 (Cotocripus; female; Argentina). Distr.- Argentina (Buenos Aires
province), southeastern Brazil, Uruguay.
setifer (LUTZ), 1913: 64 (Centrorhynchus). Brazil.
irwini SPINELLI & WIRTH, 1984a: 180 (female; Chile). Distr.- Chile, southwestern Argentina.
patagoniensis RONDEROS & SPINELLI, 1997: 34 (male, female; Argentina). Distr.- Southern
Chile, southern Argentina.
raposoensis WIRTH & BARRETO, 1978: 561 (male, female; Colombia). Distr.- Colombia.
Subgenus CULICOIDES LATREILLE

elutus MACFIE, 1948: 75 (female; Mexico). Distr.- Mexico (Chiapas, Oaxaca) to Panama.
hondurensis SPINELLI & BORKENT, 2004a: 369 (male, female; Honduras). Distr.- El Salvador,
Honduras.
luteovenus ROOT & HOFFMAN, 1937: 156 (male, female; Mexico). Distr.- USA (California,
Utah, Washington) to Panama.
neopulicaris WIRTH, 1955: 355 (male, female; USA,Texas). Distr.- USA (Texas and
Louisiana) to Costa Rica.
Subgenus DIPHAOMYIA VARGAS

DIPHAOMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides baueri
HOFFMAN, by original designation.
blantoni VARGAS & WIRTH, 1955: 33 (male, female; Mexico). Distr.- USA (Texas), Mexico
(Sinaloa, Tamaulipas, Morelos).
edeni WIRTH & BLANTON, 1974a: 23 (male, female; notes, syn.; USA, Florida). Distr.-
Southeastern USA to Bahamas.
evansi WIRTH & BLANTON, 1959: 342 (male, female; Panama). Distr.- Honduras, Costa
Rica, Panama.
freitasi WIRTH & BLANTON, 1973: 434 (female; Brazil). Distr.- Brazil (Par).
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 (Par).
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 ORTZ, 1953a: 801 (female; Venezuela). Distr.- Panama, Venezuela, Trinidad.
ronderosae SPINELLI & BORKENT, 2004a: 377 (female; Costa Rica). Distr.- Costa Rica.
tarapaca SPINELLI & WIRTH, 1984a: 184 (female; Chile). Distr.- Northern Chile.
Subgenus DRYMODESMYIA VARGAS

DRYMODESMYIA VARGAS, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides
copiosus ROOT & HOFFMAN, by original designation.
REFERENCE: WIRTH & HUBERT, 1960: 646 (key to females and some males; as copiosus group).
borinqueni FOX & HOFFMAN, 1944: 110 (male, female; Puerto Rico). Distr.- Jamaica to St. Croix.
bredini WIRTH & BLANTON, 1970b: 41 (male, female; Dominica). Distr.- Dominica.
chacoensis SPINELLI & WIRTH, 1984a: 174 (male, female; Argentina). Distr.- Bolivia, Paraguay,
Argentina, Uruguay.
haitiensis DELCOLLE, 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 (Yucatn), Central America and Caribbean to Colombia and Venezuela.
loughnani EDWARDS, 1922: 165 (female; Jamaica). Distr.- USA (Texas to Georgia), Mexico
(Yucatn), 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 (Par).
poikilonotus MACFIE, 1948: 82 (female; Mexico). Distr.- Mexico (Chiapas), Central America,
Venezuela and Trinidad to Brazil (Bahia).
cacozelus MACFIE, 1948: 85. Mexico.
hertigi WIRTH & BLANTON, 1953b: 229. Panama.
saltaensis SPINELLI & WIRTH, 1984a: 183 (female; Argentina). Distr.- Western Argentina.
uruguayensis RONDEROS, 1990a: 117 (female; Uruguay). Distr.- Uruguay.
wirthomyia VARGAS, 1953b: 227 (male; Mexico). Distr.- Mexico (Guerrero).
Subgenus GLAPHIROMYIA VARGAS

GLAPHIROMYIA VARGAS, 1960: 41 (as subgenus of Culicoides). Type species: Culicoides
scopus ROOT & HOFFMAN, by original designation.
parascopus WIRTH & BLANTON, 1978: 238 (male, female; Mexico). Distr.- Mexico (Michoacan).
scopus ROOT & HOFFMAN, 1937: 170 (male, female; Mexico). Distr.- Mexico (DF), Costa
Rica, Panama.
Subgenus HAEMATOMYIDIUM GOELDI

HAEMATOMYIDIUM GOELDI, 1905: 137. Type species: Haematomyidium paraense GOELDI,
REFERENCES: VITALE, WIRTH & AITKEN, 1981 (key to females of debilipalpis group);
WIRTH & FELIPPE-BAUER, 1989 (key to paraensis group).
annuliductus WIRTH, in VITALE et al, 1981: 150 (pupa, male, female; Panama). Distr.- Costa
Rica, Panama.
austroparaensis SPINELLI, in SPINELLI et al., 2005b: 141 (male, female; Argentina). Distr.-
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,
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, Yucatn), 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 ORTZ, 1951c: 586 (female; Venezuela). Distr.- El Salvador to Panama, Colombia,
Venezuela, Trinidad, Brazil (Par), northeastern Argentina.
glabrior MACFIE, 1940d: 27 (as variety of debilipalpis; female; Guyana). Distr.- Honduras to
Ecuador, Guyana, Surinam, Trinidad, Brazil (Par).
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, southeastern
Brazil.
imitator ORTZ, 1953b: 807 (male, female; Venezuela). Distr.- Guatemala to Panama, Venezuela.
insinuatus ORTZ & LEN, 1955: 577 (female; Ecuador). Distr.- Colombia, Ecuador, Peru,
Trinidad, Guyana, Brazil (Amazonas, Par, (?) So 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 ORTZ & LEN, 1955: 576 (female; Ecuador). Distr.- Ecuador, Venezuela, Brazil
(Par, So 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, Rondnia).
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
(Par, Santa Catarina).
youngi WIRTH & BARRETO, 1978: 562 (male, female; Colombia). Distr.- Colombia.
Subgenus HOFFMANIA FOX

HOFFMANIA FOX, 1948: 21 (as subgenus of Culicoides). Type species: Culicoides inamollae
FOX & HOFFMAN (= Culicoides insignis LUTZ), by original designation.
REFERENCES: SPINELLI, GREINER & WIRTH, 1993 (key to species groups; key to males
and females of guttatus group); WIRTH & BLANTON, 1968a (key to males and females in
hylas group).
aitkeni WIRTH & BLANTON, 1968a: 214 (female; Trinidad). Distr.- Trinidad, Brazil (Amazonas).
annettae SPINELLI & BORKENT, 2004a: 365 (male, female; Costa Rica). Distr.- Costa Rica.
batesi WIRTH & BLANTON, 1973: 426 (male, female; Brazil). Distr.- Guatemala, Colombia,
Ecuador, Bolivia, Brazil (Par).
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 (Par).
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,
Par, So 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
(Esprito 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 (Par, Pernambuco, Mato Grosso, Esprito Santo, So Paulo, Rio de
Janeiro, Santa Catarina).
foxi ORTZ, 1950c: 461 (male; Puerto Rico). Distr.- Mexico (Chiapas) to Bolivia, Puerto
Rico to northeastern Argentina.
franklini SPINELLI, in SPINELLI et al., 1993: 45 (male, female; Panama). Distr.- Mexico
(Guerrero) to Bolivia, Brazil (Par).
fusipalpis WIRTH & BLANTON, 1973: 435 (male, female; Brazil). Distr.- El Salvador to Ecuador,
Bolivia, French Guiana, Guyana, Brazil (Amazonas, Par, Bahia, Rio de Janeiro).
guttatus (COQUILLETT), 1904b: 35 (Ceratopogon; female; Brazil). Distr.- Paraguay, northeastern
Argentina, southeastern Brazil.
heliconiae FOX & HOFFMAN, 1944: 108 (male, female; Venezuela). Distr.- Belize to Ecuador,
Venezuela, Grenada, Trinidad and Tobago.
rozeboomi BARBOSA, 1947: 26. Trinidad.
hylas MACFIE, 1940d: 26 (female; Guyana). Distr.- Mexico (Veracruz) to Peru, Brazil
(Amazonas).
ignacioi FORATTINI, 1957: 215 (male, female; Brazil). Distr.- Brazil (Minas Gerais, So 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 (Yucatn, Chiapas), Central America and Caribbean to central Argentina.
inamollae FOX & HOFFMAN, 1944: 110. Puerto Rico.
painteri FOX, 1946: 257. Honduras.
lutzi COSTA L IMA, 1937a: 419 (female; Brazil). Distr.- Colombia to French Guiana,
northeastern Argentina, Brazil (Roraima, Amazonas, Par, Mato Grosso, Gois, So
Paulo, 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 ORTZ, 1950b: 455 (male, female; Venezuela). Distr.- Mexico (Oaxaca, Tabasco)
to northern Brazil (Par, Rondnia).
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, Par).
paramaruim WIRTH & BLANTON, 1973: 443 (male, female; Brazil). Distr.- Brazil (Par).
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, Gois).
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,
Par, 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 ORTZ & LEN, 1955: 574 (as variety of rozeboomi). Ecuador.
xanifer WIRTH & BLANTON, 1968a: 210 (male, female; Panama). Distr.- Honduras to Panama.
Subgenus MACFIELLA FOX

MACFIELLA FOX, 1955: 217 (as subgenus of Culicoides). Type species: Ceratopogon phlebotomus
WILLISTON, by original designation.
phlebotomus (WILLISTON), 1896: 281 (Ceratopogon; female; St. Vincent). Distr.- Coastal;
Mexico (Sinaloa) to Ecuador, Jamaica to Brazil (Maranho, Cear, Pernambuco, Gois).
amazonius MACFIE, 1935a: 52. Brazil.
willistoni WIRTH & BLANTON, 1953c: 116 (male, female; Panama). Distr.- Mexico (Sonora),
Honduras, Panama.
Subgenus MATAEMYIA VARGAS

MATAEMYIA VARGAS, 1960: 43 (as subgenus of Culicoides). Type species: Culicoides mojingaensis
WIRTH & BLANTON, by original designation.
REFERENCE: WIRTH & SORIA, 1981 (key males and females to some species, as discrepans
group).
albuquerquei WIRTH & BLANTON, 1973: 424 (male, female; Brazil). Distr.- Brazil (Par).
avilaensis ORTZ & 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 ORTZ, 1951a: 445 (male; Venezuela). Distr.- Ecuador, Venezuela, Bolivia, Brazil
(Amazonas, Par).
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 ORTZ & 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 (Par).
Subgenus OECACTA POEY

OECACTA POEY, 1853: 238. Type species: Oecacta furens POEY, by monotypy.
alahialinus BARBOSA, 1952: 11 (1953: 12) (female; Ecuador). Distr.- Costa Rica, Panama,
Colombia, Ecuador.
barbosai WIRTH & BLANTON, 1956a: 161 (male, female; Panama). Distr.- USA (Florida) to
Ecuador.
cancer HOGUE & WIRTH, 1968: 2 (larva, pupa, male, female; Costa Rica). Distr.- Mexico
(Sinaloa), El Salvador, Costa Rica.
furens (POEY), 1853: 238 (Oecacta; female; Cuba). Distr.- USA (Massachusetts to Florida
and Texas), Mexico (Campeche, Sinaloa, Yucatn, Veracruz) and Caribbean to Ecuador
and coastal Brazil.
maculithorax (WILLISTON), 1896: 277 (Ceratopogon). St. Vincent.
birabeni CAVALIERI, 1966: 59. Venezuela.
gorgasi WIRTH & BLANTON, 1953b: 232 (male, female; Panama). Distr.- Costa Rica to
Colombia.
Subgenus PSYCHOPHAENA PHILIPPI

PSYCHOPHAENA PHILIPPI, 1865: 628. Type species: Psychophaena pictipennis PHILIPPI (=
Culicoides venezuelensis ORTZ & MIRSA), by monotypy.
lacustris RONDEROS, 1990a: 116 (male, female; Argentina). Distr.- Southern Argentina.
venezuelensis ORTZ & MIRSA, 1950: 137 (Venezuela). Distr.- Costa Rica to Chile and
Central Argentina.
pictipennis (PHILIPPI), 1865: 628 (Psychophaena) (preoccupied by Culicoides
pictipennis (STAEGER), 1839). Chile.
ortizi FOX, 1952: 366. Venezuela.
Subgenus unplaced, acotylus Species Group

acotylus LUTZ, 1913: 69 (female; Brazil). Distr.- Mexico (DF), Honduras, Panama, Venezuela,
Trinidad, Surinam, Brazil (Mato Grosso, Par).
panamericanus FOX, 1947: 90. Mexico.
atripalpis WIRTH & BLANTON, 1973: 425 (female; Brazil). Distr.- Brazil (Par).
carsiomelas WIRTH & BLANTON, 1955a: 100 (male, female; Panama). Distr.- Panama,
Colombia, Brazil (Par).
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, Par).
camposi ORTZ & LEN, 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 ORTZ, 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, Par).
estevezae RONDEROS & SPINELLI, 1994: 47 (female; Argentina). Distr.- Northern Argentina.
guerrai WIRTH & BLANTON, 1971: 41 (male, female; Trinidad. Distr.- Trinidad, Brazil (Par, Bahia).
rodriguezi ORTZ, 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 ORTZ, 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 ORTZ & 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 ORTZ & LEN, 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 ORTZ & MIRSA, 1952a: 267. Venezuela.
fernandezi ORTZ, 1954: 223 (female; Venezuela). Distr.- Venezuela.
fluvialis MACFIE, 1940d: 25 (female; Guyana). Distr.- Honduras to Colombia, Venezuela,
Trinidad, Guyana, Brazil (Amazonas, Par).
leopoldoi ORTZ, 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,
yaracuyensis ORTZ, 1959: 364 (as yaracuyanus; female; Venezuela). Distr.- Venezuela.
Subgenus unplaced, leoni Species Group

benarrochi ORTZ & MIRSA, 1952b: 126 (as benarrochei; female; Venezuela). Distr.- Brazil
(Rio de Janeiro), Venezuela, Trinidad.
fieldi WIRTH & BLANTON, 1956c: 50 (male, female; Honduras). Distr.- Honduras, Costa
Rica, Panama.
gabaldoni ORTZ, 1954: 221 (female; Venezuela). Distr.- Mexico (Tabasco) to Ecuador,
Venezuela, Trinidad, Brazil (Bahia), Paraguay, northeastern Argentina.
glabellus WIRTH & BLANTON, 1956c: 47 (male, female; Panama). Distr.- Honduras to
Ecuador, Trinidad, Brazil (Bahia, Par).
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 (Gois, So 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 (Par).
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 (Par, Mato
Grosso, So Paulo, Rio de Janeiro, Santa Catarina) to northeastern Argentina.
lobatoi FELIPPE-BAUER, in FELIPPE-BAUER & QUINTELAS, 1994: 25 (female; Brazil). Distr.-
lopesi BARRETTO, 1944: 102 (male; Brazil). Distr.- Panama, Surinam, Brazil (So 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 (Par).
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
(Par, Mato Grosso).
kintzi WIRTH & BLANTON, 1953a: 72. Panama.
cylindricornis WIRTH & BLANTON, 1973: 430 (female; Brazil). Distr.- Brazil (Par).
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 ORTZ, 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 (Par).
forattinii ORTZ, 1961: 211 (female; Venezuela). Distr.- Venezuela.
goeldii WIRTH & BLANTON, 1973: 437 (male, female; Brazil). Distr.- Brazil (Par).
guamai WIRTH & BLANTON, 1973: 438 (male, female; Brazil). Distr.- Brazil (Par).
guyanensis FLOCH & ABONNENC, 1942a: 4 (male, female; French Guiana). Distr.- Panama,
Venezuela to French Guiana, Trinidad and Tobago, Brazil (Par, Pernambuco, So Paulo).
recifensis BARBOSA, 1943: 263. Brazil.
stubalensis FOX, 1946: 254. Trinidad and Tobago.
kuscheli WIRTH & BLANTON, 1978: 236 (female; Chile). Distr.- Northern Chile.
lanei ORTZ, 1950a: 431 (male; Panama). Distr.- Mexico (Veracruz), Honduras, Costa Rica,
Panama, Venezuela, Trinidad, Brazil (Par).
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, Par).
pifanoi ORTZ, 1951c: 588 (male, female; Venezuela). Distr.- Belize to Colombia, Venezuela,
Trinidad, Brazil (Par, Bahia), Paraguay.
tricoloratus WIRTH & BLANTON, 1953b: 233. Panama.
reticulatus LUTZ, 1913: 49 (male, female; Brazil). Distr.- Honduras to Colombia, Brazil
(Pernambuco, Bahia, So 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 ORTZ, 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 ORTZ, 1956: 93. Venezuela.
stigmalis WIRTH, 1952c: 245 (female; Guatemala). Distr.- Mexico (Oaxaca), Guatemala,
Costa Rica, Panama.
Subgenus unplaced, stonei Species Group

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

arubae FOX & HOFFMAN, 1944: 109 (male, female; Aruba). Distr.- USA (Texas), Mexico
(Yucatn), Aruba and Grenada, to Colombia and Venezuela.
floridensis BECK, 1951: 135 (male, female; USA, Florida). Distr.- USA (Florida), Bahamas, Bermuda.
malariologiensis PERRUOLO, 1990: 28 (female; Venezuela). Distr.- Venezuela.
pancensis BROWNE, 1980: 537 (male, female; Colombia). Distr.- Colombia.
polystictus KIEFFER, 1921b: 181 (female; Paraguay). Distr.- Paraguay.
propinquus MACFIE, 1948: 81 (male; Mexico). Distr.- Mexico (Chiapas).
trilineatus FOX, 1946: 250 (female; Virgin Islands). Distr.- Guatemala to Panama, Puerto
Rico, Virgin Islands, Dominica, Barbados, Paraguay.
unetensis PERRUOLO, 2001: 35 (female; Venezuela). Distr.- Venezuela.
wokei FOX, 1947: 90 (female; Panama). Distr.- Costa Rica, Panama.
Genus PARADASYHELEA MACFIE

PARADASYHELEA MACFIE, 1940f: 17. Type species: Dasyhelea brevipalpis INGRAM &
MACFIE, by original designation.
REFERENCES: SPINELLI, 1987 (description of female); SPINELLI & GROGAN, 2003 (key to
males and females).
brevipalpis (INGRAM & MACFIE), 1931: 178 (Dasyhelea; male; Argentina). Distr.- Southwestern
Argentina south to Tierra del Fuego.
ingrami SPINELLI & GROGAN, 2003: 572 (male, female; Argentina). Distr.- Southtern Argentina.
macfiei SPINELLI & GROGAN, 2003: 574 (male, female; Argentina). Distr.- Southwestern
TRIBE CERATOPOGONINI
Genus ALLOHELEA KIEFFER
ALLOHELEA KIEFFER, 1917b: 364. Type species: Sphaeromias pulchripennis KIEFFER, by original
designation.
neotropica WIRTH, 1991a: 503 (male, female; Jamaica). Distr.- Belize, Panama, Colombia, Jamaica.
Genus ALLUAUDOMYIA KIEFFER

ALLUAUDOMYIA KIEFFER, 1913: 12. Type species: Alluaudomyia imparunguis KIEFFER, by
monotypy.
NEOCERATOPOGON MALLOCH, 1915b: 310. Type species: Ceratopogon bellus COQUILLETT,
ISOECACTA GARRETT, 1925: 9. Type species: Isoecacta poeyi GARRETT (= Ceratopogon bellus
COQUILLETT), by original designation.
REFERENCES: HUERTA, 1996 (key to males and females from Mexico); SPINELLI & WIRTH,
1984c (key to males and females).
amazonica SPINELLI & WIRTH, 1984c: 676 (female, male; Brazil). Distr.- Brazil (Amazonas),
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
(Yucatn, 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, Par).
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, Par).
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 (Par),
Paraguay, northeastern Argentina.
punctiradialis SPINELLI & WIRTH, 1984c: 692 (preoccupied by Alluaudomyia punctiradialis
CHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Brazil.
schnacki SPINELLI, 1983b: 403 (larva, pupa, male, female; Argentina). Distr.- Eastern
Argentina, Uruguay.
sexpunctata SPINELLI & WIRTH, 1984c: 695 (female; Colombia). Distr.- Colombia.
spinellii WIRTH & GROGAN, 1988: 21. New name for tripunctata. Distr.- Colombia.
tripunctata SPINELLI & WIRTH , 1984c: 697 (preoccupied by Alluaudomyia tripunctata
CHAUDHURI, DAS GUPTA & CHAUDHURI, 1972). Colombia.
tenuiannulata SPINELLI & WIRTH, 1984c: 696 (male, female; Guatemala). Distr.- Guatemala,
Costa Rica.
youngi SPINELLI & WIRTH, 1984c: 699 (male, female; Colombia). Distr.- Panama, Colombia.
Genus AUSTROHELEA WIRTH & GROGAN

AUSTROHELEA WIRTH & GROGAN, 1988: 22. Type species: Monohelea shannoni WIRTH &
BLANTON, by original designation.
REFERENCE: WIRTH & GROGAN, 1988 (description of male).
shannoni (WIRTH & BLANTON), 1972b: 175 (Monohelea; female; Argentina). Distr.- Southern
Chile, southwestern Argentina south to Tierra del Fuego.
Genus BAEODASYMYIA CLASTRIER & RACCURT

BAEODASYMYIA CLASTRIER & RACCURT, 1979b: 100. Type species: Baeodasymyia modesta
CLASTRIER & RACCURT, by monotypy.
REFERENCE: BORKENT & CRAIG, 1999 (keys to pupae, males, females).
christopheri BORKENT, in BORKENT & CRAIG, 1999: 15 (larva, pupa, male, female; Costa
Rica). Distr.- Costa Rica.
gustavoi BORKENT, in BORKENT & CRAIG, 1999: 20 (female; Paraguay). Distr.- Paraguay,
Argentina (Misiones).
lydiae BORKENT, in BORKENT & CRAIG, 1999: 19 (male, female; Costa Rica). Distr.- Costa Rica.
michaeli BORKENT, in BORKENT & CRAIG, 1999: 13 (larva, pupa, male, female; Costa Rica).
Distr.- Costa Rica, Panama.
modesta CLASTRIER & RACCURT, 1979b: 100 (male, female; Haiti). Distr.- Haiti.
Genus BAEOHELEA WIRTH & BLANTON

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

BAHIAHELEA WIRTH, 1992: 276. Type species: Bahiahelea brasiliensis WIRTH, by original
designation.
brasiliensis WIRTH, 1992: 280 (male, female; Brazil). Distr.- Brazil (Bahia).
Genus BORKENTHELEA SPINELLI & GROGAN

BORKENTHELEA SPINELLI & GROGAN, 1993: 321. Type species: Borkenthelea nothofagus
SPINELLI & GROGAN, by original designation.
REFERENCE: SPINELLI & GROGAN, 2001 (key to males and females species).
harii SPINELLI & GROGAN, 2001: 148 (male, female; Argentina). Distr. - Southwestern
nerudai SPINELLI & GROGAN, 2001: 151 (male, female; Chile). Distr. - Southern Chile.
nothofagus SPINELLI & GROGAN, 1993: 323 (male, female; Argentina). Distr.- Southwestern
quatei SPINELLI & GROGAN, 2001: 153 (male, female; Argentina). Distr.- Argentina (Chubut).
Genus BRACHYPOGON KIEFFER

BRACHYPOGON KIEFFER, 1899: 69. Type species: Ceratopogon vitiosus WINNERTZ, by
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 (Rondnia).
bifidus SPINELLI & GROGAN, 1998: 66 (male, female; Dominica). Distr.- El Salvador, Dominica.
bimaculatus SPINELLI & GROGAN, 1998: 67 (male, female; Colombia). Distr.- Mexico
(Yucatn), 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
(Yucatn, 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 (Par).
pseudoparaensis SPINELLI & GROGAN, 1998: 64 (male; Brazil). Distr.- Brazil (Par).
ringueleti SPINELLI, 1990: 748 (male, female; Argentina). Distr.- Southern Argentina.
schmitzi SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondnia).
spatuliformis SPINELLI & GROGAN, 1998: 65 (male; Brazil). Distr.- Brazil (Rondnia).
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
(Yucatn), 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, southern
Argentina.
cuacuahuitlus HUERTA & BORKENT, 2005: 115 (male; Mexico). Distr.- Mexico (Jalisco).
hugoi SPINELLI & GROGAN, 1994: 2 (male, female; Colombia). Distr.- Colombia.
mapuche SPINELLI, 1990: 750 (male, female; Argentina). Distr.- Southwestern Argentina.
misionensis SPINELLI, 1990: 752 (male; Argentina). Distr.- Northeastern Argentina,
southeastern Brazil.
pallidipennis SPINELLI & GROGAN, 1994: 6 (male, female; Honduras). Distr.- Honduras.
wirthi SPINELLI, 1990: 753 (female; Argentina). Distr.- Northwestern Argentina.
Genus CACAOHELEA WIRTH & GROGAN

CACAOHELEA WIRTH & GROGAN, 1988: 33. Type species: Cacaohelea youngi WIRTH &
GROGAN, by original designation.
youngi WIRTH & GROGAN, 1988: 34 (female; Costa Rica). Distr.- Costa Rica.
Genus CERATOCULICOIDES WIRTH & RATANAWORABHAN

CERATOCULICOIDES WIRTH & RATANAWORABHAN, 1971a: 170. Type species: Helea
longipennis WIRTH, by original designation.
aliciae HUERTA & BORKENT, 2005: 112 (male, female; Mexico). Distr.- Mexico (Jalisco).
Genus DIAPHANOBEZZIA INGRAM & MACFIE

DIAPHANOBEZZIA INGRAM & MACFIE, 1931: 223. Type species: Diaphanobezzia pellucida
INGRAM & MACFIE, by original designation.
araucaria SPINELLI, 1996: 77 (female; Argentina). Distr.- Southwestern Argentina.

patagonica SPINELLI & GROGAN, 1990: 127 (female; Argentina). Distr.- Southwestern
Argentina.
pellucida INGRAM & MACFIE, 1931: 223 (male; Argentina). Distr.- Southwestern Argentina.
spinellii WIRTH & GROGAN, 1988: 48 (male, female; Argentina). Distr.- Southwestern
Argentina.
Genus DOWNESHELEA WIRTH & GROGAN

DOWNESHELEA WIRTH & GROGAN, 1988: 50. Type species: Monohelea stonei WIRTH, by
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 (Par).
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 (Par), Guyana.
multilineata (LUTZ), 1914: 93 (Palpomyia; male, female; Brazil). Distr.- USA (Florida, Puerto
Rico), Mexico (Yucatn), 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, by
REFERENCE: WIRTH, 1994b (key to males and females).
Subgenus ECHINOHELEA MACFIE

blantoni WIRTH, 1994b: 232 (male, female; Panama). Distr.- Mexico (Yucatn, 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 (Par, Rondnia).
leei WIRTH, 1994b: 237 (male; Colombia). Distr.- Colombia.
macfiei LANE, 1948: 228 (female; Brazil). Distr.- Brazil (So Paulo).
neotropica WIRTH, 1994b: 237 (male, female; Colombia). Distr.- Panama to Ecuador, Brazil
(Amazonas, Par, Rio de Janeiro).
ornatipennis MACFIE, 1940c: 188 (male, female; Guyana). Distr.- Panama, Trinidad, Guyana,
Brazil (Par).
panamensis WIRTH, 1994b: 239 (male, female; Panama). Distr.- Panama to Ecuador.
richardsi MACFIE, 1940c: 189 (female; Guyana). Distr.- Guyana, Brazil (Par, Rondnia,
So 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 (Par).
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 &
micronyx WIRTH & BLANTON, 1970a: 12 (female; Colombia). Distr.- Costa Rica, Colombia.
Genus MACRUROHELEA INGRAM & MACFIE

MACRUROHELEA INGRAM & MACFIE, 1931: 203. Type species: Macrurohelea caudata
INGRAM & MACFIE, by original designation.
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
wirthi SPINELLI & GROGAN, 1984: 965 (female; Argentina). Distr.- Southwestern Argentina.
yamana SPINELLI & GROGAN, 1999: 710 (male, female; Argentina). Distr.- Argentina (Tierra
del Fuego).
Genus MONOHELEA KIEFFER

MONOHELEA KIEFFER, 1917b: 294. Type species: Monohelea hieroglyphica KIEFFER, by
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 (Gois,
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 (Yucatn, Tamaulipas), Guatemala, Panama.
maya FELIPPE-BAUER, HUERTA & IBAEZ-BERNAL, 2000a: 815 (male, female; Mexico). Distr.-
Mexico (Yucatn).
mayeri ORTZ, 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,
roraimensis FELIPPE-BAUER & SPINELLI, 1991: 202 (male; Brazil). Distr.- Brazil (Roraima).
urracaisi LANE & WIRTH, 1964: 231 (male, female; Panama). Distr.- Panama, Guyana.
uruguayensis FELIPPE-BAUER & SPINELLI, 1998: 63 (male; Uruguay). Distr.- Uruguay.
Genus NANNOHELEA GROGAN & WIRTH

NANNOHELEA GROGAN & WIRTH, 1980a: 374. Type species: Ceratopogon bourioni
CLASTRIER, by original designation.
clastrieri GROGAN & WIRTH, 1980a: 381 (male; Colombia). Distr.- Colombia.
Genus NOTIOHELEA GROGAN & WIRTH

NOTIOHELEA GROGAN & WIRTH, 1979b: 283. Type species: Notiohelea chilensis GROGAN
& WIRTH, by original designation.
chilensis GROGAN & WIRTH, 1979b: 284 (female; Chile). Distr.- Southern Chile.
pilosa SPINELLI & GROGAN, 1990: 133 (female; Argentina). Distr.- Southwestern Argentina.
Genus PARABEZZIA MALLOCH

PARABEZZIA MALLOCH, 1915a: 358. Type species: Parabezzia petiolata MALLOCH, by original
designation.
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 Ros).
blantoni WIRTH, 1965b: 220 (female; Panama). Distr.- Panama.
brasiliensis SPINELLI & GROGAN, 1987: 34 (female; Brazil). Distr.- Brazil (Mato Grosso).
brunnea WIRTH, 1965b: 220 (female; Panama). Distr.- Belize, Panama.
caribbeana CLASTRIER & RACCURT, 1979a: 173 (female; Haiti). Distr.- Haiti.
cayoensis SPINELLI & GROGAN, 1987: 36 (female; Belize). Distr.- Belize.
clastrieri SPINELLI & GROGAN, 1987: 23 (female; El Salvador). Distr.- Belize, El Salvador.
costalis WIRTH, 1965b: 221 (male, female; Panama). Distr.- Panama.
fuscipennis WIRTH, 1965b: 223 (female; Panama). Distr.- Panama.
haitiensis CLASTRIER & RACCURT, 1979a: 170 (female; Haiti). Distr.- Haiti.
hondurensis SPINELLI & GROGAN, 1987: 25 (male, female; Honduras). Distr.- Honduras.
inaequalis SPINELLI & GROGAN, 1987: 38 (male, female; Brazil). Distr.- Brazil (Amazonas).
jamaicensis WIRTH, 1965b: 224 (female; Jamaica). Distr.- Jamaica.
neunguis GROGAN & WIRTH, 1977: 65 (female; USA, Virgina). Distr.- USA (Virginia,
Maryland), El Salvador.
pallida SPINELLI & GROGAN, 1987: 40 (female; Mexico). Distr.- Mexico (Oaxaca).
panamensis WIRTH, 1965b: 225 (male, female; Panama). Distr.- Panama.
pseudunguis SPINELLI & GROGAN, 1987: 30 (male, female; Panama). Distr.- Panama.
raccurti SPINELLI & GROGAN, 1987: 42 (male, female; El Salvador). Distr.- El Salvador,
northern Argentina.
spangleri WIRTH, 1965b: 227 (female; Puerto Rico). Distr.- Puerto Rico.
uncinata (JOHANNSEN), 1943: 761 (Stilobezzia; female; USA, Alabama). Distr.- USA (Alabama,
Maryland, Virginia, Arkansas), Jamaica.
unguis WIRTH, 1965b: 228 (male, female; USA, Arizona). Distr.- USA (Arizona, Texas) to Colombia.
wirthi CLASTRIER & RACCURT, 1979a: 169 (female; Haiti). Distr.- Haiti.
Genus PARASTILOBEZZIA WIRTH & BLANTON

PARASTILOBEZZIA WIRTH & BLANTON, 1970a: 10. Type species: Parastilobezzia leei WIRTH
& BLANTON, by original designation.
leei WIRTH & BLANTON, 1970a: 10 (male, female; Colombia). Distr.- Costa Rica, Colombia.
Genus RHYNCHOHELEA WIRTH & BLANTON

RHYNCHOHELEA WIRTH & BLANTON, 1970c: 96. Type species: Rhynchohelea monilicornis
WIRTH & BLANTON, by original designation.
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,
forattinii WIRTH & GROGAN, 1988: 81 (male, female; Brazil). Distr.- Costa Rica, Brazil
(Santa Catarina).
guyana CLASTRIER, 1984: 2 (female; French Guiana). Distr.- Mexico (Yucatn), French
Guiana, Brazil (Mato Grosso).
Genus STILOBEZZIA KIEFFER

STILOBEZZIA KIEFFER, 1911a: 118. Type species: Stilobezzia festiva KIEFFER, by original
designation. HARTOMYIA MALLOCH, 1915a: 339. Type species: Ceratopogon pictus
COQUILLETT (= Stilobezzia coquilletti KIEFFER), by original designation.
REFERENCES: HUERTA, 1996 (key to males and females from Mexico); INGRAM &MACFIE,
1931 (key males and females from Patagonia and southern Chile); LANE & FORATTINI,
1961 (key to males and females from Neotropical Region north of Patagonia and
southern Chile); WIRTH & GROGAN, 1988 (key to subgenera, as part of generic key).
Subgenus ACANTHOHELEA KIEFFER

ACANTHOHELEA KIEFFER, 1917a: 198. Type species: Acanthohelea pruinosa KIEFFER, by
monotypy.
amazonica CLASTRIER, 1991: 306 (male, female; French Guiana). Distr.- French Guiana, Peru.
atrichopogon LANE & FORATTINI, 1956: 208 (male, female; Panama). Distr.- Panama.
bicinctipes INGRAM & MACFIE, 1931: 195 (female; Chile). Distr.- Southern Chile.
edwardsi INGRAM & MACFIE, 1931: 198 (male, female; Argentina). Distr.- Southwestern Argentina.
furva INGRAM & MACFIE, 1931: 200 (female; Argentina). Distr.- Southern Chile, southwestern
Argentina.
guianae (MACFIE), 1940d: 28 (Acanthohelea; male, female; Guyana). Distr.- Guyana.
hirsuta INGRAM & MACFIE, 1931: 201 (male, female; Argentina). Distr.- Southwestern Argentina.
maia LANE & FORATTINI, 1958: 204 (male, female; Panama). Distr.- Panama, Colombia.
manaosensis LANE & FORATTINI, 1958: 205 (male, female; Brazil). Distr.- Brazil (Amazonas).
nigerrima INGRAM & MACFIE, 1931: 196 (female; Argentina). Distr.- Southwestern Argentina.
ornata LANE & FORATTINI, 1958: 206 (female; Panama). Distr.- Panama.
ornaticrus INGRAM & MACFIE, 1931: 194 (female; Chile). Distr.- Southern Chile.
patagonica INGRAM & MACFIE, 1931: 196 (male, female; Argentina). Distr.- Southwestern
Argentina.
rava INGRAM & MACFIE, 1931: 203 (male, female; Chile). Distr.- Southern Chile.
sanctibernardini KIEFFER, 1917b: 308 (female; Paraguay). Distr.- Paraguay.
succinea INGRAM & MACFIE, 1931: 200 (male, female; Argentina). Distr.- Chile, southwestern
Argentina south to Tierra del Fuego.
tibialis LANE & FORATTINI, 1956: 209 (male, female; Panama). Distr.- Panama.
varia INGRAM & M ACFIE , 1931: 191 (male, female; Chile). Distr.- Southern Chile,
Subgenus EUKRAIOHELEA INGRAM & MACFIE

EUKRAIOHELEA INGRAM & MACFIE, 1921: 347. Type species: Eukraiohelea africana INGRAM
& MACFIE, designation by MACFIE, 1940f: 22.
REFERENCE: WIRTH & SPINELLI, 1992b (key males and females).
amnigena (MACFIE), 1935a: 56 (Eukraiohelea; female; Brazil). Distr.- USA (Florida, South
Carolina), (?) Jamaica, (?) Puerto Rico, Brazil (Maranho), 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 (Yucatn), 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 (Yucatn), Panama, Peru.
bicolor LANE, 1947a: 208 (male; Brazil). Distr.- Panama, Brazil (So Paulo), Argentina
(Buenos Aires).
bimaculata LANE & FORATTINI, 1956: 211 (male, female; Panama). Distr.-Panama, Trinidad,
bispinosa KIEFFER, 1917b: 310 (female; Paraguay). Distr.- Brazil (So 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 (Gois, So Paulo).
coquilletti KIEFFER, 1917b: 308. (New name for pictus COQUILLETT, female; USA, Virginia).
Distr.- USA (Virgina, Illinois, Maryland), Mexico (Yucatn, Chiapas) to Panama,
Trinidad, Brazil (So Paulo).
picta (COQUILLETT), 1905: 60 (Ceratopogon) (preoccupied by Sphaeromias pictus (MEIGEN),

1818). USA (Virginia).
coracina KIEFFER, 1917b: 311 (male; Paraguay). Distr.- Brazil (So 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 (So Paulo).
dureti LANE & FORATTINI, 1958: 210 (male; Brazil). Distr.- Brazil (So Paulo).
esmeralda LANE & FORATTINI, 1958: 211 (male, female; Panama). Distr.- Panama.
femoralis LANE & FORATTINI, 1956: 212 (male; Panama). Distr.- Mexico (Yucatn), Panama.
fiebrigi KIEFFER, 1917b: 309 (female; Paraguay). Distr.- Brazil (So 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 (Yucatn,
Tamaulipas), Panama, Brazil (Minas Gerais).
punctipes WIRTH, 1953a: 79. USA (Florida).
macfiei LANE, 1947a: 213 (male, female; Brazil). Distr.- Brazil (So Paulo, Rio de Janeiro).
maculata LANE, 1947a: 207 (male; Brazil). Distr.- Mexico (Yucatn), Panama, Venezuela,
merceri CAZORLA & SPINELLI, in CAZORLA et al., 2005: 290 (male, female, pupa; Peru). Distr.- Peru.
modesta LANE, 1947a: 206 (female; Brazil). Distr.- Brazil (So 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
(So Paulo).
panamensis LANE & FORATTINI, 1958: 218 (male, female; Panama). Distr.- Panama, Puerto
Rico, Brazil (So Paulo, Rio de Janeiro).
paulistensis LANE, 1947a: 200 (male, female; Brazil). Distr.- Panama, USA (Puerto Rico),
Brazil (So Paulo, Rio de Janeiro).
punctulata LANE, 1947a: 204 (female; Brazil). Distr.- Mexico (Tabasco), Panama, Per,
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
(Yucatn), 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 (Esprito Santo).
venezuelensis ORTZ, 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 (So Paulo).
TRIBE HETEROMYIINI WIRTH, 1962: 275

Genus CLINOHELEA KIEFFER
CLINOHELEA KIEFFER, 1917b: 295. Type species: Ceratopogon variegatus WINNERTZ
(= Ceratopogon unimaculata MACQUART), by original designation.
Subgenus CERATOBEZZIA KIEFFER

CERATOBEZZIA KIEFFER, 1917b: 326. Type species: Ceratobezzia fallax KIEFFER, by
fallax KIEFFER, 1917b: 326 (female; Paraguay). Distr.- El Salvador to Colombia, Venezuela,
Brazil (Amazonas, Par, 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 (Par, So Paulo), Paraguay,
argentina LANE & DURET, 1954: 248 (female; Argentina). Distr.- Northeastern Argentina.
damascenoi LANE & DURET, 1954: 250 (female; Brazil). Distr.- Peru, Bolivia, French Guiana,
Brazil (Par), Paraguay.
horacioi LANE, 1944: 257 (male, female; Brazil). Distr.- Mexico (Yucatn, 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 (Gois, Minas Gerais,
So Paulo, Rio de Janeiro), Paraguay.
nigripes MACFIE, 1939b: 205 (female; Brazil). Distr.- Costa Rica to Colombia, Brazil (So
Paulo, Santa Catarina), northeastern Argentina.
rubriceps KIEFFER, 1917b: 318 (female; Paraguay). Distr.- Paraguay.
saltanensis LANE & DURET, 1954: 252 (female; Argentina). Distr.- Brazil (Rondnia, 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 ARRIBLZAGA), 1893: 227 (Pachyleptus; female; Argentina). Distr.-

Northeastern Argentina.
bejaranoi DURET & LANE, 1955: 36 (male, female; Argentina). Distr.- Northeastern
Argentina.
castanea LANE, 1946a: 214 (female; Brazil). Distr.- Brazil (So 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
(Rondnia), 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 (So Paulo, Rio de Janeiro,
Paran), Paraguay.
fasciatus (WALKER), 1856: 426 (Pachyleptus) (preoccupied by Heteromyia fasciata SAY, 1825).
South America.
orellana (ROBACK), 1957: 1 (Palpomyia; female; Peru). Distr.- Peru.
wokei WIRTH & GROGAN, 1977: 181 (pupa, male, female; Nicaragua). Distr.- Nicaragua.
Genus PELLUCIDOMYIA MACFIE

PELLUCIDOMYIA MACFIE, 1939a: 99. Type species: Pellucidomyia ugandae MACFIE, by
MACFIEHELEA LANE, 1946a: 208. Type species: Macfiehelea oliveirai LANE, by original
designation.
REFERENCE: WIRTH, 1960 (World key to males and females).
blantoni (LANE), 1956b: 435 (Macfiehelea; female; Panama). Distr.- Panama.

lanei WIRTH & RATANAWORABHAN, 1971b: 446 (female; Colombia). Distr.- Colombia, Jamaica.
oliveirai (LANE), 1946a: 209 (Macfiehelea; female; Brazil). Distr.- Brazil (Mato Grosso, Minas Gerais).
wirthi (LANE), 1956b: 437 (Macfiehelea; female; Panama). Distr.- USA (Texas) to Colombia.
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 (Rondnia), Uruguay.
Genus JOHANNSENOMYIA MALLOCH

JOHANNSENOMYIA MALLOCH, 1915a: 332. Type species: Johannsenomyia halteralis
MALLOCH (= Ceratopogon argentata LOEW), designation by WIRTH, 1952a: 211.
DICROHELEA KIEFFER, 1917b: 363. Type species: Palpomyia filicornis KIEFFER, designation
by MACFIE, 1940f: 26.
blantoni (LANE & WIRTH ), 1961: 81 (Dicrohelea; female; Panama). Distr.- Panama.
lanei WIRTH, 1965a: 4 (female; Brazil). Distr.- Brazil (Santa Catarina).
Genus LANEHELEA WIRTH & BLANTON

LANEHELEA WIRTH & BLANTON, 1972a: 433. Type species: Lanehelea leei WIRTH and
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,
aenipes (MACFIE), 1940c: 193 (Bezzia; female; Guyana). Distr.- Guyana

limitrofe SPINELLI & FELIPPE-BAUER, 1990a: 87 (female; Argentina). Distr.- Northeastern
Argentina, Uruguay.
nemoralis (MACFIE), 1940c: 190 (Johannsenomyia; male; Guyana). Distr.- Guyana.
nigripes (MACFIE), 1939b: 216 (Johannsenomyia; female; Brazil). Distr.- Brazil (Santa Catarina).
pullata (WIRTH), 1952a: 213 (Johannsenomyia; female; USA, California). Distr.- USA (California,
Arizona, Texas) to Brazil (Amap).
bicellii (LANE), 1961b: 449 (Nilobezzia). Brazil.
termophila (SPINELLI), 1984: 197 (Neobezzia; male, female; Argentina). Distr.- Brazil
(Rondnia), northern Argentina.
Genus NEOBEZZIA WIRTH & RATANAWORABHAN

NEOBEZZIA WIRTH & RATANAWORABHAN, 1972a: 477. Type species: Neobezzia clavipes
WIRTH & RATANAWORABHAN, by original designation.
REFERENCE: WIRTH & RATANAWORABHAN, 1972a (key to females).
albitarsis WIRTH & RATANAWORABHAN, 1972a: 478 (female; Brazil). Distr.- Brazil (Par).
amnicola (MACFIE), 1940d: 30 (Bezzia; female; Guyana). Distr.- Belize to Brazil (Amazonas,
Rondnia, Mato Grosso, So Paulo).
blantoni WIRTH & RATANAWORABHAN, 1972a: 482 (female; Panama). Distr.- Nicaragua to
Colombia, Brazil (Amazonas, Par, Rondnia), 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 (Rondnia, Bahia).
paraensis (LANE), 1958: 33 (Bezzia; female; Brazil). Distr.- Brazil (Par).
schwarzii (COQUILLETT), 1901a: 605 (Ceratopogon; female; USA, Texas). Distr.- USA (South
Carolina to Texas and Florida), Mexico (Yucatn), Bahamas, Cuba, Panama, Brazil
(Rio de Janeiro), northeastern Argentina.
banksi (GERRY), 1933: 94 (Bezzia). Cuba.
brasiliensis (LANE), 1958: 28 (Bezzia). Brazil.
Genus SPHAEROHELEA SPINELLI & FELIPPE-BAUER

SPHAEROHELEA SPINELLI & FELIPPE-BAUER, 1990b: 195. Type species: Sphaerohelea biestroi
SPINELLI & FELIPPE-BAUER, by original designation.
biestroi SPINELLI & FELIPPE-BAUER, 1990b: 197 (female; Argentina). Distr.- Northeastern
Argentina.
TRIBE PALPOMYIINI ENDERLEIN, 1936: 49

Genus AMEROHELEA GROGAN & WIRTH
AMEROHELEA GROGAN & WIRTH, 1981: 1280. Type species: Amerohelea galindoi GROGAN
& WIRTH, by original designation.
REFERENCES: GROGAN & WIRTH, 1981 (key to females); HUERTA, 1996 (key to females
from Mexico).
dalcyi GROGAN & WIRTH, 1981: 1289 (female; Brazil). Distr.- Brazil (Amazonas).
fasciata GROGAN & WIRTH, 1981: 1283 (male, female; Belize). Distr.- Mexico (Sonora,
Morelos, Tabasco, Chiapas) to Colombia.
frontispina (DOW & TURNER), 1976: 138 (Bezzia; female; USA, Texas). Distr.- USA
(California to Texas) south to Colombia and Venezuela.
galindoi GROGAN & WIRTH , 1981: 1294 (male, female; Colombia). Distr.- Panama,
Colombia,Venezuela, northern Argentina.
nelsoni GROGAN & WIRTH, 1981: 1290 (female; Brazil). Distr.- Brazil (Amazonas).
pseudofasciata GROGAN & WIRTH, 1981: 1286 (female; Brazil). Distr.- Southeastern Brazil,
Argentina (Misiones, Ro 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 (Par,
Minas Gerais, So 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 and
females of nobilis and punctipennis groups); SPINELLI & WIRTH, 1989b (key to females
and some males of glabra and brevicornis groups); SPINELLI & WIRTH, 1990 (key to males
and females of gibbera group); SPINELLI & WIRTH, 1991 (key to males and females of
dentifemur and venustula groups).
aitkeni SPINELLI & WIRTH, 1991: 5 (female; Brazil). Distr.- Brazil (Par).
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 Ros, Buenos Aires, Ro 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, Rondnia).
dentifemur SPINELLI & WIRTH, 1991: 3 (male, female; Colombia). Distr.- Belize, Colombia,
Trinidad, Brazil (Par).
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, Par, Brasilia,
D.F., So 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, Rondnia).
megatheca SPINELLI & WIRTH, 1990: 22 (female; Colombia). Distr.- Colombia, Brazil
(Rondnia).
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 (Par).
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 (Yucatn, Tabasco, Quintana
Roo), El Salvador, Honduras, Jamaica, Dominica, Brazil (Amazonas), Argentina
(Buenos Aires province).
raposoensis SPINELLI & WIRTH, 1991: 4 (male, female; Colombia). Distr.- Mexico (Yucatn),
Colombia.
roldani SPINELLI & WIRTH, 1981: 187 (larva, pupa, male, female; Argentina). Distr.- Eastern
Argentina (Entre Rios, Buenos Aires).
schmitzorum DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 54 (female; Brazil). Distr.-
Brazil (Rondnia).
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 (Par).
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 schnacki
SPINELLI & GROGAN, by original designation.
REFERENCE: SPINELLI & GROGAN, 1986 (key to females).
dycei SPINELLI & GROGAN, 1986: 456 (pupa, male, female; Uruguay). Distr.- Uruguay, eastern
and central Argentina.
kremeri SPINELLI & GROGAN, 1985: 332 (female; Brazil). Distr.- Brazil (Amazonas).
schnacki SPINELLI & GROGAN, 1985: 331 (female; Ecuador). Distr.- Ecuador, northern
Argentina.
uruguayensis SPINELLI & GROGAN, 1986: 458 (female; Uruguay). Distr.- Northeastern and
central Argentina, Uruguay.
Genus PACHYHELEA WIRTH

PACHYHELEA W IRTH , 1959a: 50. Type species: Ceratopogon magnus C OQUILLETT
(= Ceratopogon pachymerus WILLISTON), by original designation.
albidiventris (KIEFFER), 1917b: 316 (Sphaeromias; female; Colombia). Distr.- Colombia.

pachymera (WILLISTON), 1900: 224 (Ceratopogon; female; Mexico). Distr.- USA (Texas) to
central Argentina.
latifemoris (INGRAM & MACFIE), 1931: 231 (Johannsenomyia). Argentina.
Genus PALPOMYIA MEIGEN

PALPOMYIA MEIGEN, 1818: 82. Type species: Ceratopogon flavipes MEIGEN, by monotypy.
Generic name first published in synonymy with Ceratopogon but available under ICZN
Code Article 11(e).
REFERENCES: INGRAM & MACFIE, 1931 (key males and females from Patagonia and
southern Chile); LANE, 1960 (key to females).
aculeata INGRAM & MACFIE, 1931: 215 (female; Chile). Distr.- Southern Chile.
almeidai (LANE), 1946b: 219 (Sphaeromias; female; Brazil). Distr.- Brazil (So 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 (So Paulo).
castanea MACFIE, 1939b: 211 (female; Brazil). Distr.- Brazil (So Paulo, Santa Catarina).
catarinensis LANE, 1960: 385 (female; Brazil). Distr.- Brazil (Santa Catarina).
chilensis INGRAM & MACFIE, 1931: 218 (female; Chile). Distr.- Southern Chile, ? Brazil (So
Paulo).
columbiana KIEFFER, 1917b: 323 (female; Colombia). Distr.- Colombia.
conifera MACFIE, 1939b: 214 (male; Brazil). Distr.- Brazil (Brasilia, D.F., Gois, Santa Catarina).
coroicoensis WIRTH, 1974: 54. New name for boliviensis. Distr.- Bolivia.
boliviensis (KIEFFER), 1917b: 331 (Jenkinsia; female) (preoccupied by Palpomyia boliviensis
KIEFFER, 1917b). Bolivia.
crassicrus KIEFFER, 1917b: 321 (female; Paraguay). Distr.- Brazil (So Paulo), Paraguay.
guarani LANE, 1946b: 221 (pupa, male, female; Brazil). Distr.- Brazil (So Paulo, Esprito
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 (So Paulo).
lacustris LANE, FORATTINI & RABELLO, 1955: 83 (pupa, male; Brazil). Distr.- Brazil (So 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 (So Paulo).
nigroscutellata LANE, 1947b: 444 (female; Brazil). Distr.- Brazil (So Paulo, Rio de Janeiro).
oliveirai LANE, 1947b: 443 (female; Brazil). Distr.- Brazil (Rondnia, Minas Gerais).
pampana LANE, 1960: 386 (female; Brazil). Distr.- Brazil (Rio Grande do Sul).
paraensis LANE, 1960: 387 (female; Brazil). Distr.- Brazil (Par).
patagonica INGRAM & MACFIE, 1931: 222 (female; Argentina). Distr.- Southwestern
Argentina.
paulistensis LANE, 1947b: 445 (female; Brazil). Distr.- Panama, Brazil (So Paulo).
pseudolacustris DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 55 (male, female; Brazil).
Distr.- Brazil (Rondnia).
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
(So 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,
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 (Rondnia, Santa Catarina).
wirthi LANE, FORATTINI & RABELLO, 1955: 82 (pupa, female; Brazil). Distr.- Brazil (So 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
(Rondnia).
TRIBE STENOXENINI COQUILLETT, 1899: 61

Genus PARYPHOCONUS ENDERLEIN
PARYPHOCONUS ENDERLEIN , 1912: 57. Type species: Paryphoconus angustipennis
ENDERLEIN, by original designation.
REFERENCES: HUERTA, 1996 (key to females from Mexico); SPINELLI & WIRTH, 1984b
(key to females).
aemulus MACFIE, 1940b: 180 (female; Guyana). Distr.- Peru, Guyana, Brazil (Amazonas,
Mato Grosso).
amapaensis LANE, 1961b: 450 (female; Brazil). Distr.- Brazil (Amap).
angustipennis ENDERLEIN, 1912: 57 (female; Brazil). Distr.- Colombia, Peru, Venezuela,
Brazil (Gois, So 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 (Gois, So 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 (Par).
fuscus LANE, 1946a: 206 (female; Brazil). Distr.- Brazil (Gois).
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 (Par, Santa Catarina).
guianae MACFIE, 1940b: 180 (female; Guyana). Distr.- Ecuador, Guyana, Brazil (Par, Mato
Grosso, So Paulo).
kiefferi LANE, 1956a: 302 (female; Brazil). Distr.- Brazil (Gois).
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 (Par, Mato
Grosso, So Paulo).
maya SPINELLI & WIRTH, 1984b: 899 (female; Belize). Distr.- Mexico (San Luis Potosi), Belize.
mayeri WIRTH, 1959b: 236 (female; Brazil). Distr.- Brazil (Amazonas, Rondnia).
misionensis SPINELLI, 1998: 52 (female; Argentina). Distr.- Northeastern Argentina.
neotropicalis (LANE), 1948: 229 (Macropeza; female; Brazil). Distr.- Brazil (So Paulo).
nigripes MACFIE, 1939c: 8 (female; Argentina). Distr.- Guyana, Brazil (Par, Rondnia),
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, Par).
paranaensis SPINELLI & WIRTH, 1984b: 902 (female; Argentina). Distr.- Northeastern Argentina.
paulistensis LANE, 1961c: 457 (female; Brazil). Distr.- Brazil (So 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,
Par, Rondnia).
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 (Par).
wirthi LANE, 1961c: 458 (female; Brazil). Distr.- Brazil (Brasilia, D.F.).
wygodzinskyi LANE, 1946a: 208 (female; Brazil). Distr.- Brazil (Rio de Janeiro).
Genus STENOXENUS COQUILLETT

STENOXENUS COQUILLETT, 1899: 61. Type species: Stenoxenus johnsoni COQUILLETT, by
monotypy.
REFERENCE: WIRTH & RATANAWORABHAN, 1972b (key to females).
aductus DIPPOLITO & SPINELLI, in DIPPOLITO et al., 1995: 56 (female; Brazil). Distr.- Brazil
(Rondnia).
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),
carrerai LANE, 1956a: 305 (as carreirai; female; Brazil). Distr.- Brazil (Gois).
dimorphus KIEFFER, 1909: 47 (male, female; Peru). Distr.- Peru.
excentricus LANE, 1961c: 453 (female; Brazil). Distr.- Brazil (Par).
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 (Gois).
paraensis LANE, 1956a: 306 (female; Brazil). Distr.- Brazil (Par).
pseudofulvus SPINELLI, 1998: 54 (female; Argentina). Distr.- Northeastern Argentina,
Uruguay.
setiger MACFIE, 1939b: 138 (female; Brazil). Distr.- Jamaica, Dominican Republic, Brazil
(Santa Catarina), northeastern Argentina.
Unplaced genus of Forcipomyiinae

DIDYMOPHLEPS WEYENBERGH , 1883: 108. Type species: Didymophleps hortorum
WEYENBERGH, by monotypy.
Unplaced species of Forcipomyiinae

Didymophleps hortorum WEYENBERGH, 1883: 110. Argentina.
Unplaced species of Ceratopogonidae

Ceratopogon claripennis LYNCH ARRIBLZAGA, 1893: 229. Argentina.
7
IDENTIFICATION
BORKENT & WIRTH (1997) listed the living and extinct species of biting midges for the World
and an updated version is available on the web (http://www.inhs.uiuc.edu/cee/FLYTREE/)
CeratopogonidaeCatalog.pdf). BORKENT & SPINELLI (2000) provide a catalog of the
ceratopogonids south of the USA which included those species from the Nearctic portion
of Mexico. Additional information was given concerning which life stages were originally
described and the distribution of each of the 1066 mentioned species. References to all
pertinent taxonomic literature was provided for each genus, including available keys. Here,
we provide an updated catalog of those species restricted to the Neotropical Region, as
defined by GRIFFITHS (1980) (Table 4). The number of species in each genus is given in
Table 5. WIRTH & GROGAN (1988) provided a revision of the tribe Ceratopogonini, the
most diverse tribe at the generic level. This revision included a key to the genera and
subgenera, and a diagnosis, distribution and list of included species for each genus.
The best work for identifying Neotropical Culicoides is a wing atlas published by WIRTH
et al. (1988), which contains wing photographs and a summary of the numerical characters
of the majority of the species presently known for the region. This work may be used
together with the revision of the genus for Panama by WIRTH & BLANTON (1959), and the
keys for the Amazon Basin by WIRTH & BLANTON (1973), for the species occurring south
of the Amazon Basin by SPINELLI & WIRTH (1986b), and for Argentina (SPINELLI et al.,
2005b). Description of further species may be found in SPINELLI & BORKENT (2004a) for
Costa Rica, BRICKLE & HAGAN (1999) for Belize, FELIPPE-BAUER & QUINTELAS (1994) and
FELIPPE-BAUER et al. (2000a, b) for Brazil, FELIPPE-BAUER et al. (2003, 2005) for Peru and
PERRUOLO (1990, 2001) for Venezuela.
Adult ceratopogonids are easily collected with a variety of methods and are best
preserved in 70% alcohol. These specimens can be identified to genus while in fluid but
generally need to be placed on microscope slides to be identified to species. Specimens that
are not slide mounted should be pinned, as old alcohol material is impossible to slide
mount and study properly. For more information, see chapter Capture and study of
Ceratopogonidae.
Table 5. Numbers of named valid species in each genus of Neotropical Ceratopogonidae.
Taxon Number of species

Subfamily Leptoconopinae NO
Leptoconops SKUSE 12
Subfamily Forcipomyiinae LENZ
Atrichopogon KIEFFER 100
Forcipomyia MEIGEN 211
Subfamily Dasyheleinae LENZ
Dasyhelea KIEFFER 58
Subfamily Ceratopogoninae NEWMAN
Tribe Culicoidini KIEFFER
Culicoides LATREILLE 266
Paradasyhelea MACFIE 3
Tribe Ceratopogonini
Allohelea KIEFFER 1
Alluaudomyia KIEFFER 19
Austrohelea WIRTH & GROGAN 1
Baeodasymyia CLASTRIER & RACCURT 5
Baeohelea WIRTH & BLANTON 1
Bahiahelea WIRTH 1
Borkenthelea SPINELLI & GROGAN 4
Brachypogon KIEFFER 25
Cacaohelea WIRTH & GROGAN 1
Ceratoculicoides WIRTH & RATANAWORABHAN 1
Diaphanobezzia INGRAM & MACFIE 4
Downeshelea WIRTH & GROGAN 18
Echinohelea MACFIE 11
Fittkauhelea WIRTH & BLANTON 1
Isthmohelea INGRAM & MACFIE 1
Leptohelea WIRTH & BLANTON 1
Macrurohelea INGRAM & MACFIE 12
Monohelea KIEFFER 18
Nannohelea GROGAN & WIRTH 1
Notiohelea GROGAN & WIRTH 2
Parabezzia MALLOCH 24
Parastilobezzia WIRTH & BLANTON 1
Rhynchohelea WIRTH & BLANTON 1
Schizonyxhelea CLASTRIER 2
Stilobezzia KIEFFER 65
Tribe Heteromyiini WIRTH
Clinohelea KIEFFER 12
Heteromyia SAY 11
Pellucidomyia MACFIE 4
Table 5. Continued.
Taxon Number of species
Physohelea GROGAN & WIRTH 2

Tribe Sphaeromiini NEWMAN
Austrosphaeromias SPINELLI 2
Groganhelea SPINELLI & DIPPOLITO 1
Johannsenomyia MALLOCH 2
Lanehelea WIRTH & BLANTON 2
Mallochohelea WIRTH 6
Neobezzia WIRTH & RATANAWORABHAN 8
Nilobezzia KIEFFER 3
Sphaerohelea SPINELLI & FELIPPE-BAUER 1
Tribe Palpomyiini ENDERLEIN
Amerohelea GROGAN & WIRTH 11
Bezzia KIEFFER 46
Clastrieromyia SPINELLI & GROGAN 4
Pachyhelea WIRTH 2
Palpomyia MEIGEN 46
Phaenobezzia HAESELBARTH 2
Tribe Stenoxenini COQUILLETT
Paryphoconus ENDERLEIN 40
Stenoxenus COQUILLETT 16
Unplaced genus
Didymophleps WEYENBERGH 1
Unplaced species 2
Total number of species (not including unplaced taxa) 1095
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. Most
females with serrate mandibles. Anepisternum divided into anterior and posterior portions by
anepisternal cleft. Postnotum lacking medial longitudinal groove. Wing with 1-2 radial veins
reaching margin, with two median vein branches (posterior one may be weak) reaching wing
margin. Wings overlapping each other over the abdomen in living individuals at rest (with ex-
ceptions in a few species of Stilobezzia). Foreleg shorter than hind leg. Tarsomere 1 of each leg
equal to or longer than tarsomere 2 (with a few exceptions in Forcipomyia).
Adults are similar to some Chironomidae but may be distinguished by the combina-
tion of a lack of a medial groove on the postnotum (otherwise only in Podonominae) and
a lack of a m-cu crossvein (present in Podonominae).
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, and
no open spiracles.
Eggs are not diagnosable at this time (but see key below).
KEY TO THE GENERA OF ADULT CERATOPOGONIDAE IN THE NEOTROPICAL REGION
There are currently 1095 species of Ceratopogonidae recorded from the Neotropical Region
(Table 5) but there are many more awaiting description, especially in the genera Forcipomyia,
Atrichopogon, Dasyhelea, Culicoides and Stilobezzia. Species of the family may be found at all altitudes
and in virtually every habitat where there is even a little moisture. The following key includes
genera known from the Neotropics and includes males, for the first time, of all genera (males
not known for Groganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, and Sphaerohelea).
This key is intended to work for both alcohol and slide mounted material. We have
indicated in bold type those character states that are best seen with a compound microscope
(i.e. with slide mounted material), although with skill and patience many of these character
states may also be observed in alcohol specimens. Some wing veins may be difficult to see
in some taxa; for alcohol specimens, tilting the wing may help to see faint wing veins and
with slide material, phase contrast microscopy provides the best resolution.
When the two claws at the end of a leg are the same size, they are referred to as being
equal; conversely when they differ in size they are called unequal. The costal ratio is the
length of the costa from the wing base (measured from the arculus), divided by the total
wing length. Terms for wing veins and cells follow the system of the Manual of Nearctic
Diptera (MCALPINE et al., 1981), with modifications proposed by SZADZIEWSKI (1996) and
are fully labeled here in Figs. 10C, D and 11I. As is widely used by other Dipterists, veins
are in capital letters and those of cells in small letters.
1 Eyes widely separated dorsally, lacking frontal suture (Fig. 6A); palpus with 4
segments (only one beyond elongate and/or swollen third) and with membranous
gap between segment 3 and 4+5 (Fig. 9A); wing without crossvein r-m (Fig. 10A);
female with radial cells fused into swollen unit (Fig. 10A); without palisade setae
on first tarsomere of hind leg (Fig. 14B); male gonostylus with an apical,
socketed peg (Fig. 19A); female cercus very elongate (Fig. 22E) ...........................
................................................................................................................... Leptoconops SKUSE
Eyes narrowly or widely separated dorsally, with frontal suture (Figs. 6B-G);
palpus with 2-5 segments but never with a membranous gap between the last two
segments (Figs. 9B-E); wing with crossvein r-m (Figs. 10B-J); radial cells distinct
or, if fused or absent (Figs. 11A, F), r-m clearly present; with (Fig. 14A) or without
(Fig. 14B) palisade setae on first tarsomere of hind leg; male gonostylus without
apical, socketed peg (Figs. 19B-F, 20A-F) although apex may be pointed (Fig.
19E); female cercus short (Figs. 22F, 23A-E) .............................................................. 2
Frontal suture
Palpus
Lacinia
Mandible
Labrum
Labium
A. Leptoconops
B. Culicoides C. Culicoides
Frontal suture
Clypeus
Labium
D. Rhynchohelea
E. Echinohelea
F. Parabezzia G. Fittkauhelea
Figure 6. Head capsules in anterior view. A: Female Leptoconops floridensis WIRTH (Nearctic Region).
B: Female Culicoides yukonensis HOFFMAN, mouthparts spread laterally (Nearctic Region). C: Male
Culicoides yukonensis, mouthparts spread laterally (Nearctic Region). D: Female Rhynchohelea
monilicornis WIRTH & BLANTON (Nearctic and Neotropical Regions). E: Female Echinohelea sp. F:
Female Parabezzia sp. G: Female Fittkauhelea amazonica WIRTH & BLANTON (Neotropical Region). (A-
C from MCALPINE et al., 1981; D-G courtesy of INBio, Costa Rica).
2(1) Wing with (Figs. 10B, E, G) or without (Figs. 10D, J, 11A-C) numerous
macrotrichia on membrane; hind first tarsomere with scattered setae or, at most, a
row of strong bristles (but these not forming a tight row, especially basally on the
tarsomere) (Fig. 14B); female with two small, equal claws on each leg (Figs. 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 species
have some macrotrichia more basally on the wing but these are scattered) (Figs.
11E, I); hind first tarsomere with row of palisade setae with at least more basal
setae with abutting bases, some groups with an additional pronounced basal stout
spine (Figs. 14A, G-I); females with two small, equal claws on each leg or with 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) or
narrow (Figs. 10B, E); paratergite narrow (as in Fig. 9G) or well-developed,
without setae or, if setae present, these number at least 10 and all are equal in size
and length ............................................................................................................................ 4
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 sensilla
coeloconica; male with short setae on first flagellomere, less than 1/2 the
length of those on following flagellomeres (Fig. 7C); male flagellomeres
sculptured (Fig. 7C); without tibial spur on foreleg; wing membrane without
distinct pattern of dark pigmentation (Fig. 10E) .......................... Dasyhelea KIEFFER
Eyes contiguous to widely separated medially (Figs. 6B, C); some antennal
flagellomeres with sensilla coeloconica; male with setae on first
flagellomere about as long as those on following flagellomeres; male
flagellomeres not sculptured; with tibial spur on foreleg; wing membrane
with or without distinct patterns of dark and/or light pigmentation (Figs. 10F-H);
CULICOIDINI .................................................................................................................. 6
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 without
distinct patterns of dark and/or light pigmentation (10F, G) ...................................
............................................................................................................... Culicoides LATREILLE
Wing with both radial cells obliterated (Fig. 10H); costa not reaching beyond 0.43
of wing length; females with reduced, vestigial mandibles; wing membrane
without distinct patterns of dark and/or light pigmentation (Fig. 10H) .................
.............................................................................................................. Paradasyhelea MACFIE
A. Forcipomyia
B. Forcipomyia
C. Dasyhelea
D. Rhynchohelea
Figure 7. Antennae. A: Female Forcipomyia glauca MACFIE (Palaearctic, Nearctic and Neotropical
Regions). B: Female Forcipomyia monilicornis (COQUILLETT) (Holarctic Region). C: Male Dasyhelea
pseudoincisurata WAUGH and WIRTH (Nearctic Region). D: Female Rhynchohelea monilicornis WIRTH &
BLANTON (Nearctic and Neotropical Regions) (all from MCALPINE et al., 1981).
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 is
unknown but likely with broadly separated eyes, antennal flagellomeres 10 and 11
fused and a katepisternal seta) ........................................................................................ 9
Wing with at least one well-developed radial cell (Figs. 10J, 11E, I); costal ratio
more than 0.50; female claws equal or unequal, some with only a single claw on
one or more of the legs .................................................................................................. 15
9(8) Male antennal flagellomeres 10 and 11 fused or with less than 13
flagellomeres; wing with or without radial cells; katepisternum with 1 or more
lateral setae (can be seen in alcohol specimens by holding specimen upright and
looking along plane of thoracic pleura in outline) (Fig. 9G) .................................. 10
Male antenna with flagellomeres 10 and 11 separate and with 13
flagellomeres present; wing without radial cell (cells completely fused) (Figs.
11A, B, D); katepisternum without setae (Fig. 9H) ............................................. 13
10(9) Female antenna with 12 flagellomeres (Fig. 7D); female labium stout, truncate,
with apical hooklike structures (Fig. 6D) ........................................................................
........................................................... Rhynchohelea WIRTH & BLANTON (male unknown)
Female antenna with 13 flagellomeres; female proboscis normal (Figs. 6B, C, E-G)
.............................................................................................................................................. 11
11(10) Male flagellum with 7-8 flagellomeres (Fig. 8A); palpus with 3 segments (Fig. 9C)
............................................................................................. Nannohelea GROGAN & WIRTH
Male flagellum with at least 9 flagellomeres (almost always with 13 but with 10
and 11 fused); palpus with 5 segments ........................................................................ 12
12(11) Anepisternum lacking setae; male antenna with flagellomeres 2-6 fused (6-11
also fused) ......................................................................... Brachypogon KIEFFER (in part)
Anepisternum with at least 1 seta on posterior margin (Fig. 9G); male antenna
with flagellomeres 2-5 articulated, not fused (flagellomeres 7-11 fused, sometimes
including flagellomere 6) ........................ Ceratoculicoides WIRTH & RATANAWORABHAN
13(9) Palpus with 5 segments; wing length 0.7-0.8 mm; male with well-developed
parameres .................................................................................... Schizonyxhelea CLASTRIER
Palpus with 2 segments (Fig. 9B); wing length < 0.6 mm; male without parameres
.............................................................................................................................................. 14
14(13) Fore tibia with apical spur; macrotrichia confined to margin and apex of wing
(in some, also a few on apices of M1 and M2); costal ratio > 0.40 (Fig. 11B); male
antenna with 6 separate flagellomeres (Fig. 8C); male with abdominal tergites 1-3
pale and 4-8 brown ........................................................... Baeohelea WIRTH & BLANTON
Fore tibia without apical spur; macrotrichia long and abundant over entire
wing; costal ratio < 0.36; male antenna with 12 flagellomeres but with 2-7, 8-9,
and 10-11 fused (Fig. 8B); male with abdomen uniformly brown ............................
................................................................................... Baeodasymyia CLASTRIER & RACCURT
A. Nannohelea
B. Baeodasymyia
C. Baeohelea
Sensilla coeloconica
D. Borkenthelea
Figure 8. Antennae. A: Male Nannohelea bourioni (CLASTRIER) (Palaearctic Region). B: Male

Baeodasymyia modesta CLASTRIER and RACCURT (Neotropical Region). C: Male Baeohelea nana WIRTH &
BLANTON (Neotropical Region). D: Male Borkenthelea harii (Neotropical Region). (A-B from WIRTH
& GROGAN, 1988; C courtesy of INBio, Costa Rica).
Fused 4+5
B. Baeohelea C. Nannohelea D. Leptohelea
A. Leptoconops E. Parabezzia
Paratergite
Anepisternum
Katepisternum
F. Atrichopogon G. Ceratoculicoides
Forecoxa
Midcoxa
H. Parabezzia
Figure 9. Female structures. A: Leptoconops bequaerti (KIEFFER) right palpus (Nearctic and Neotropical
Regions). B: Baeohelea nana WIRTH & BLANTON right palpus (Neotropical Region). C: Nannohelea sp.
right palpus. D: Leptohelea micronyx WIRTH & BLANTON right palpus (Neotropical Region). E: Parabezzia
sp. right palpus. F: Atrichopogon sp. thorax, in lateral view. G: Ceratoculicoides sp. thorax, in lateral view. H:
Parabezzia bystraki GROGAN & WIRTH thorax, in lateral view (Nearctic Region). (A, E courtesy of
INBio, Costa Rica; D from WIRTH & GROGAN, 1988; F from SZADZIEWSKI et al., 1997).
15(8) Wing with characteristic pattern of darkly mottled pigmentation, more or less
scattered on membrane (Fig. 12C); thorax and at least hind femur and tibia with
patterned pigmentation; male aedeagus with distinctive posteromedial loop (Fig.
21A) ....................................................................................................... Monohelea KIEFFER1
Wing without pigmentation or, if present, restricted to radial sector or with
discrete dots, stripes or bands (Figs. 11E, H), or if with substantial pattern of
pigmentation (Fig. 11H) then thorax and all femora and tibiae uniformly dark
brown; other taxa with variably pigmented thorax and legs; aedeagus, if present,
without posteromedial loop ........................................................................................... 16
16(15) Costa extending nearly to tip of wing; female wing with costa extending beyond
apex of second radial cell (Figs. 12B, D, E) (extension also in male wing of
Bahiahelea and Leptohelea) ................................................................................................. 17
Costa not extending beyond 0.85 of wing length (Figs. 11E, I); female wing with
costa ending at or extending slightly past apex of second radial cell ................... 25
17(16) Halter saclike, without constriction below knob (Fig. 13J) .........................................
...................................................................................... Diaphanobezzia INGRAM & MACFIE
Halter with knob and distinct stem .............................................................................. 18
18(17) Male ..................................................................................................................................... 19
Female ................................................................................................................................. 21
19(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 ..................... 20
20(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 ....... 22
22(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)
Clypeus laterally fused to head (Figs. 6F, G); wing with 1 radial cell (Fig. 12B); with
2 claws on each leg, although hind leg claws may be unequal ............................... 23
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 without
distinct necks .............................................................. Fittkauhelea WIRTH & BLANTON
1
M. fairchildi LANE & WIRTH does not key out well. Adults have banded wings, as in Downeshelea, but
the male has an aedeagus with a distinctive posteromedial loop.
A. Leptoconops B. Forcipomyia
r-m c First radial cell Second radial cell
R1 R3
r3
m1
M1
m2
anal cua1
A M2
CuP CuA2 CuA1
C. Atrichopogon D. Atrichopogon
E. Dasyhelea
F. Culicoides
G. Culicoides H. Paradasyhelea
J. Brachypogon
I. Cacaohelea
Figure 10. Female wings. A: Leptoconops sp. B: Forcipomyia fairfaxensis WIRTH (Nearctic Region). C:
Atrichopogon levis (COQUILLETT) (Nearctic Region). D: Atrichopogon sp. E: Dasyhelea pseudoincisurata
WAUGH & WIRTH (Nearctic Region). F: Culicoides insignis LUTZ (Nearctic and Neotropical Regions).
G: Culicoides copiosus ROOT & HOFFMAN (Nearctic Region). H: Paradasyhelea ingrami (Neotropical
Region) I: Cacaohelea youngi WIRTH & GROGAN (Neotropical Region). J: Brachypogon stigmalis
(COQUILLETT). (A-G, J, from MCALPINE et al., 1981; I from WIRTH & GROGAN, 1988).
A. Rhynchohelea B. Baeohelea
C. Leptohelea D. Schizonyxhelea
F. Brachypogon
E. Alluaudomyia
G. Echinohelea H. Allohelea
R1 R2 R3
I. Stilobezzia J. Borkenthelea
Figure 11. Female wings. A: Rhynchohelea monilicornis WIRTH & BLANTON (Nearctic and Neotropical
Regions). B: Baeohelea nana WIRTH & BLANTON (Neotropical Region). C: Leptohelea micronyx WIRTH &
BLANTON (Neotropical Region). D: Schizonyxhelea forattinii WIRTH & GROGAN (Neotropical Region).
E: Alluaudomyia bella (COQUILLETT) (Nearctic Region). F: Brachypogon paraensis WIRTH & BLANTON
(Neotropical Region). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Allohelea
johannseni (WIRTH) (Nearctic Region). I: Stilobezzia lutea (MALLOCH) (Nearctic Region). J: Borkenthelea
nothofagus SPINELLI & WIRTH (Neotropical Region). (A, E, G, I from MCALPINE et al., 1981; C, D, F-
H from WIRTH & GROGAN, 1988).
A. Austrohelea B. Parabezzia
C. Monohelea D. Bahiahelea
E. Parastilobezzia F. Downeshelea
G. Physohelea H. Clinohelea
I. Paryphoconus J. Stenoxenus
Figure 12. Wings. A: Female Austrohelea shannoni WIRTH & GROGAN (Neotropical Region). B:
Female Parabezzia petiolata MALLOCH (Nearctic Region). C: Female Monohelea hieroglyphica KIEFFER
(Neotropical Region). D: Female Bahiahelea brasiliensis WIRTH (Neotropical Region). E: Female
Parastilobezzia leei WIRTH & BLANTON (Neotropical Region). F: Female Downeshelea sp. G: Female
Physohelea oedidactyla (INGRAM & MACFIE) (Neotropical Region). H: Female Clinohelea bimaculata
(LOEW) (Nearctic Region). I: Female Paryphoconus angustipennis ENDERLEIN (Neotropical Region). J:
Male Stenoxenus coomani SGUY (Oriental Region) (A, E from WIRTH & GROGAN, 1988; B-C, H-J
from MCALPINE et al., 1981).
A. Stenoxenus B. Palpomyia
C. Bezzia D. Phaenobezzia
E. Neobezzia F. Groganhelea
G. Mallochohelea H. Amerohelea
J. Diaphanobezzia
I. Clastrieromyia
Figure 13. Female wings. A: Stenoxenus coomani SGUY (Oriental Region). B: Palpomyia plebeja (LOEW)
(Nearctic Region). C: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). D:
Phaenobezzia pistiae (INGRAM & MACFIE) (Afrotropical Region). E: Neobezzia blantoni WIRTH &
RATANAWORABHAN (Neotropical Region). F: Groganhelea rondoniensis SPINELLI & DIPPOLITO
(Neotropical Region). G: Mallochohelea smithi (Lewis) (Nearctic Region). H: Amerohelea sp.
(Neotropical Region) I: Clastrieromyia schnacki SPINELLI & GROGAN (Neotropical Region). J: Halter
Diaphanobezzia araucaria SPINELLI (Neotropical Region) (A-D, G from MCALPINE et al., 1981; H from
GROGAN & WIRTH, 1981; F from SPINELLI & DIPPOLITO, 1995; I from SPINELLI & GROGAN, 1986; E
from WIRTH & RATANAWORABHAN, 1972a; J from SPINELLI, 1996).
Palpus with 5 segments; eyes bare; spermathecae with distinct slender necks
.................................................................................................................... Bahiahelea WIRTH
25(16) Male ..................................................................................................................................... 26
Female ................................................................................................................................. 39
26(25) Hind leg with single long claw and small basal tooth (Fig. 14H) ..............................
................................................................................................................... Allohelea KIEFFER
Hind leg with two equal or subequal claws ................................................................ 27
27(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 in
outline) (Fig. 9G) ............................................................... Brachypogon KIEFFER (in part)
Katepisternum without setae .................................................................................... 28
28(27) Wing with 1 radial cell (Fig. 11E) ................................................................................. 29
Wing with 2 radial cells ................................................................................................... 33
29(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 alcohol
specimens) ......................................................................................................................... 30
30(29) Wing with pigmentation spots at least immediately basal to or over r-m and over
apex of radial cell (Fig. 11E), some with further pigmentation ................................
............................................................................................................ Alluaudomyia KIEFFER
Wing without pigmentation spots ................................................................................ 31
31(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 to
be split medially) (Fig. 20E) or with aedeagus absent but with a transverse
bar ventral to the aedeagus (Fig. 20D) ..................... Stilobezzia KIEFFER (in part)
Palpus with 4 segments (as in Fig. 9E); hind tarsomere 1 with only 1 row of
palisade setae; aedeagus continuous medially (as in Fig. 21C) ......................... 32
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 and
CuA2 (Fig. 12F); aedeagus continuous medially (Fig. 20F) ...................................
............................................................................................ Downeshelea WIRTH & GROGAN
Wing with or without pattern of pigmentation but if pattern present, then
without discrete patches on M2 and CuA2 (posterior portion of wing may be
generally more darkly pigmented) (Figs. 11J, 12A) and with aedeagus divided
medially (Figs. 20B, C) .................................................................................................. 34
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 moderately
elongate (Figs. 20D, E) .................................................................................................... 36
36(35) Flagellomeres 3-10 fused (Fig. 8D) ........................ Borkenthelea SPINELLI & GROGAN
Antenna with flagellomeres separate, not fused ........................................................ 37
37(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 ................... 38
38(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 aedeagus
appears to be split medially) (Fig. 20E) or with aedeagus absent but with a
transverse bar ventral to the aedeagus (Fig. 20D); antennal flagellomeres
without sensilla coeloconica ........................................................ Stilobezzia KIEFFER
Hind tarsomere 1 with only 1 row of palisade setae (as in Fig. 14A); wing without
macrotrichia; aedeagus undivided, somewhat triangular; some antennal
flagellomeres with sensilla coeloconica ............. Macrurohelea INGRAM & MACFIE
39(25) Katepisternum with 1 or more lateral setae (can be seen in alcohol specimens
by holding specimen upright and looking along plane of thoracic pleura in
outline) (Fig. 9G) ............................................................... Brachypogon KIEFFER (in part)
Katepisternum without seta ...................................................................................... 40
40(39) Wing with 1 radial cell (Figs. 11C, E) .......................................................................... 41
Wing with 2 radial cells (first may be small) (Figs. 10I, 11H, I) ............................ 43
41(40) Palpus with 4 segments (as in Fig. 9E); claws on hind legs equal .............................
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 of
radial cell (Fig. 11E); hind tarsomere 1 with only one row of palisade setae .........
Wing without pigmentation spots; hind tarsomere 1 with at least 1 1/2 rows of
palisade 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 ............................... 44
44(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 second
radial cell ............................................................................................................................ 45
Wing without patches of discrete pigmentation ....................................................... 47
45(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 ................................................................ 46
46(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 somewhat
broad; medial vein reduced to faint trace on basal portion; single claw on each leg
short, stout and sharp (Fig. 15F).. ................................ Cacaohelea WIRTH & GROGAN
First radial cell small to well-developed, with R3 at most thickened at apex (Fig.
11I); second radial cell narrow; medial veins clearly visible but with base of M2
sometimes absent; claw on at least hind leg elongate (Figs. 15A-C) ........................
................................................................................................ Stilobezzia KIEFFER (in part)
49(47) Radial cells separated by a short longitudinal vein (R2); wing with a few
macrotrichia restricted to extreme periphery at tip; antennal flagellomeres
without sensilla coeloconica ..................................... Isthmohelea INGRAM & MACFIE
Radial cells separated by a short transverse vein (R2); wing with or without
macrotrichia on apical 1/4; some antennal flagellomeres with sensilla
coeloconica (either only on flagellomere 1 or only on flagellomeres 8 and
12) ........................................................................................................................................ 50
50(49) Wing without macrotrichia; mandibles reduced, vestigial; eyes bare or
pubescent; claws small, equal in all legs ..................... Notiohelea GROGAN & WIRTH
Wing with macrotrichia on apical 1/4; mandibles toothed; eyes pubescent;
only fore- and midlegs with small, equal claws, or claws unequal on all legs (as in
Fig. 14F) ............................................................................................................................. 51
51(50) Radial cells subequal, small; R3 thickened; midleg claws unequal and somewhat
similar to those of hind leg; sensilla coeloconica only on flagellomere 1 ........
........................................................................................ Borkenthelea SPINELLI & GROGAN
Second radial cell about twice as long as first radial cell; R3 not thickened; midleg
with small, equal claws, much shorter than elongate single claw (with basal tooth)
of hind leg; sensilla coeloconica only on flagellomeres 8 and 12 .....................
............................................................................................. Austrohelea WIRTH & GROGAN
52(7) Tarsomere 5 of foreleg swollen (Figs. 15G, 17A, C); HETEROMYIINI ........... 53
Tarsomere 5 of foreleg not swollen ............................................................................. 56
53(52) M barely sessile, forking at r-m (Fig. 12G) .................. Physohelea GROGAN & WIRTH
M broadly sessile, forking before r-m (Fig. 12H) ...................................................... 54
54(53) Fore femur swollen, with numerous thick ventral spines (Fig. 17C) ........................
........................................................................................................................ Heteromyia SAY
Fore femur slender, with at most a few slender spines ........................................... 55
55(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 length
of tarsomere 1 (Fig. 15H); hind claw very long, either single or with a small basal
tooth, much longer than claws of fore and midlegs (Fig. 15H) ................................
.............................................................................................................. Pellucidomyia MACFIE
Fourth tarsomere of foreleg cordate of mid and hind leg bifid and spinose (Fig.
17B); female hind leg not much longer than other legs, with tarsomere 2 less than
twice the length of tarsomere 1; both hind claws about same length as those on
fore- and midleg ................................................................................... Clinohelea KIEFFER
H
H
Palisade setae B. Other subfamilies

A. Most Ceratopogoninae
H H
Empodium
C. Atrichopogon D. Forcipomyia
F M H
H
E. Dasyhelea F. Alluaudomyia
H H H
G. Echinohelea H. Allohelea I. Downeshelea

Figure 14. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Ceratopogon MEIGEN
first tarsomere. B: Forcipomyia sp. first tarsomere. C: Atrichopogon levis (COQUILLETT) (Nearctic
Region). D: Forcipomyia fusicornis (COQUILLETT) (Nearctic and Neotropical Regions). E: Dasyhelea
pseudoincisurata WAUGH & WIRTH (Nearctic Region). F: Alluaudomyia bella (COQUILLETT) (Nearctic
Region). G: Echinohelea lanei WIRTH (Nearctic and Neotropical Regions). H: Allohelea neotropica
WIRTH (Neotropical Region). I: Downeshelea sp. (C-E, G from MCALPINE et al., 1981; F, H-I from
WIRTH & GROGAN, 1988).
F H
A. Stilobezzia B. Stilobezzia
M H M H
F
F M H F
C. Stilobezzia D. Leptohelea E. Parastilobezzia
M H
F H H
M
F
M
F
H. Pellucidomyia
F. Cacaohelea G. Clinohelea
Figure 15. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Stilobezzia antennalis
(COQUILLETT) fifth tarsomere (Nearctic and Neotropical Regions). B: Stilobezzia antennalis
(COQUILLETT) fifth tarsomere (Nearctic and Neotropical Regions). C: Stilobezzia lutea (MALLOCH) fifth
tarsomeres (Nearctic Region). D: Leptohelea micronyx WIRTH & BLANTON fifth tarsomeres (Neotropical
Region). E: Parastilobezzia leei WIRTH & BLANTON fifth tarsomeres (Neotropical Region). F: Cacaohelea
youngi WIRTH & GROGAN (Neotropical Region). G: Clinohelea sp. H: Pellucidomyia ugandae MACFIE tarsi
(Afrotropical Region). (A-B from MCALPINE et al., 1981; C-F from WIRTH & GROGAN, 1988; G from
GROGAN & WIRTH, 1975; H from DE MEILLON & WIRTH, 1991).
F M H
F M
A. Neobezzia
H
C. Groganhelea
H
B. Neobezzia
E. Phaenobezzia
D. Groganhelea
Figure 16. Female leg structures (F = foreleg, M = midleg, H = hind leg). A: Neobezzia sp. fifth
tarsomeres. B: Neobezzia sp. tarsi. C: Groganhelea rondoniensis SPINELLI & DIPPOLITO tarsi (Neotropical
Region). D: Groganhelea rondoniensis SPINELLI & DIPPOLITO tarsi (Neotropical Region). E: Phaenobezzia
sp. hind leg. (A-B from SPINELLI & WIRTH, 1993; C-D from SPINELLI et al., 1995; E from WIRTH &
GROGAN, 1982).
F H
A. Clinohelea B. Clinohelea
F
C. Heteromyia D. Heteromyia
H
E. Heteromyia
H F
F. Stenoxenus G. Johannsenomyia
M H
H. Johannsenomyia I. Johannsenomyia
Figure 17. Female fifth tarsomeres (F = foreleg, M = midleg, H = hind leg). A: Clinohelea bimaculata
(LOEW) (Nearctic Region). B: Clinohelea bimaculata (LOEW) (Nearctic Region). C: Heteromyia fasciata
SAY (Nearctic Region). D: Heteromyia fasciata SAY (Nearctic Region). E: Heteromyia fasciata SAY
(Nearctic Region). F: Stenoxenus coomani SGUY (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).
56(52) Male genitalia large, with gonocoxites fused dorsally and ventrally (Fig. 21B);
female genitalia with sternite 8 large and plate like (Fig. 23A) ..................................
................................................................................................................. Echinohelea MACFIE
Male genitalia small or large, if large then gonocoxites not fused; female genitalia
with sternite 8 moderately developed .......................................................................... 57
57(56) Male (male of Groganhelea, Sphaerohelea are unknown) .............................................. 58
Female ................................................................................................................................. 72
58(57) Gonostylus much reduced in size (Figs. 21D, 22B) or fused with gonocoxite (Fig.
22D) .................................................................................................................................... 59
Gonostylus large, clearly present (Figs. 21F, 22A, C) (male Lanehelea are not
known but likely key to at least this point) ................................................................. 61
59(58) Wing with 2 clearly defined radial cells (Fig. 13I); gonostylus well- defined and
with pointed apex (Fig. 21D) .................................Clastrieromyia SPINELLI & GROGAN
Wing with 1 radial cell or with, at most, a faint indication of a division between
first and second radial cell (Fig. 13D); gonostylus either small and apically
rounded (Fig. 22B) or not clearly defined and fused with gonocoxite (Fig. 22D)
.............................................................................................................................................. 60
60(59) Anepisternum with numerous elongate setae; gonocoxite short and gonostylus
present but small Fig. 22B) ................................................. Phaenobezzia HAESELBARTH
Anepisternum without setae; gonocoxite and gonostylus fused and elongate (Fig.
22D) .......................................................................................................Nilobezzia KIEFFER
61(58) Genitalia with gonocoxite and tergite 9 elongate (Fig. 22C) .................................. 62
Genitalia more stout, with gonocoxite and tergite 9 not unusually elongate (Figs.
21F, 22A) ............................................................................................................................ 64
62(61) Wing with 1 radial cell (as in Fig. 13D) ....... Neobezzia WIRTH & RATANAWORABHAN
Wing with 2 radial cells (Fig. 13G) ............................................................................... 63
63(62) Forefemur with ventral thick spines .................................. Austrosphaeromias SPINELLI
Forefemur without ventral thick spines ....................................... Mallochohelea WIRTH
(Note. Some Mallochohelea species have males with non-elongate genitalia)
64(61) Tarsomere 5 of at least hind leg with stout, blunt spines (Fig. 17H); apex of tergite
9 with single row of thick, stout setae (Fig. 21C) .............. Johannsenomyia MALLOCH
Tarsomere 5 without thick spines, or if present, these more slender and sharply
pointed (Fig. 16A); tergite 9 with scattered setae, never arranged in a single row of
thick, stout setae ............................................................................................................... 65
65(64) With 1-2 stout setae on apex of the cercus (Fig. 21E); parameres fused, with
fusion restricted to most anteromedial portion (Fig. 21E) ..................................
............................................................................................. Amerohelea GROGAN & WIRTH
With several to many setae on the cercus; parameres separate or with fusion
various ............................................................................................................................... 66
66(65) Forefemur with stout spines (Fig. 18A) ...................................................................... 67
Forefemur without stout spines .................................................................................... 68
67(66) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part)
Wing with 2 radial cells (Fig. 13B) ................................... Palpomyia MEIGEN (in part)
68(66) Palpus with 4 segments ............................................................... Stenoxenus COQUILLETT
Palpus with 5 segments ................................................................................................... 69

69(68) Thorax slender and dorsoventrally flattened; paratergite with numerous small
setae; 1 radial cell (Fig. 12I) ..................................................... Paryphoconus ENDERLEIN
Thorax stouter and not dorsoventrally flattened; paratergite without setae; 1-2
radial cells .......................................................................................................................... 70
70(69) Wing with 1 radial cell (Fig. 13C) .......................................... Bezzia KIEFFER (in part)
Wing with 2 radial cells (Fig. 13B) ............................................................................... 71
71(70) Hind femur swollen (Fig. 18C); abdomen white, contrasting with dark thorax .....
.....................................................................................................................Pachyhelea WIRTH
Hind femur not swollen (Fig. 18A); abdomen light to dark brown, similar to dark
thorax ...................................................................................... Palpomyia MEIGEN (in part)
72(57) Wing with M2 strongly curving near base (Fig. 13A); palpus with 4 segments ......
.......................................................................................................... Stenoxenus COQUILLETT
Wing with M2 with not or only slightly curved near base (Figs. 13B-I); palpus with
5 segments ......................................................................................................................... 73
73(72) Tarsomere 5 armed ventrally with stout black blunt spines (Figs. 16A, B, 17G-I)
(these spines somewhat sharp pointed in Austrosphaeromias); abdomen without
tergal apodemes; sternite 8 with or without (Sphaerohelea only) postero-
lateral hair tufts, with long, medially overlapping, curved setae (best
examined at 100X) (Fig. 23B); ....................................... SPHAEROMIINI..........74
Tarsomere 5 unarmed, or if with some ventral spines, these are slender and with
sharp tips (Fig. 16E); abdomen with (Figs. 23C-E) or without tergal apodemes
(not visible in some teneral or young specimens); sternite 8 without hair tufts
.............................................................................................................................................. 81
74(73) Femora without spines; claws equal on foreleg, unequal on mid- and hind legs
(Figs. 17G-I) ............................................................................... Johannsenomyia MALLOCH
Femora with or without spines; claws equal on all legs or with only forelegs
unequal ............................................................................................................................... 75
75(74) Claws gently curved distally, with slender basal tooth on internal side ............... 76
Claws straight or flattened distally, with blunt or slender external basal tooth (Figs.
16A, B) ............................................................................................................................... 78
76(75) Forefemur swollen, with numerous thick ventral spines .............................................
.................................................................................................... Austrosphaeromias SPINELLI
Fore femur slender, unarmed ........................................................................................ 77
77(76) Costa extending to wing tip; tarsomere 5 with 2-3 pairs of ventral spines .............
............................................................................. Sphaerohelea SPINELLI & FELIPPE-BAUER
Costa not extending beyond 0.85 of wing length; tarsomere 5 with 4-5 pairs of
ventral spines ...................................................................... Lanehelea WIRTH & BLANTON
78(75) Wing with 1 radial cell; costa long, extending nearly to wing tip with costal ratio
over 0.87; femora without ventral spines .................................................................... 79
Wing with 1-2 radial cells; costa short, extending to less than 0.84 of wing length;
femora with or without ventral spines ......................................................................... 80
F M H
A. Palpomyia
H
F
B. Bezzia C. Pachyhelea
Figure 18. Female legs (F = foreleg, M = midleg, H = hind leg). A: Palpomyia plebeja (LOEW) (Nearctic
Region). B: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and Neotropical Regions). C: Pachyhelea
pachymera (WILLISTON) (Nearctic and Neotropical Regions) (all from MCALPINE et al., 1981).
Parameres
A. Leptoconops B. Forcipomyia
Parameres
Parameres
C. Dasyhelea D. Culicoides
Parameres
Parameres Aedeagus
E. Stenoxenus F. Bahiahelea
Figure 19. Male genitalia, ventral view. A: Leptoconops torrens (TOWNSEND) (Nearctic Region). B:
Forcipomyia acidicola (TOKUNAGA) (Holarctic Region). C: Dasyhelea pseudoincisurata WAUGH & WIRTH
(Nearctic Region). D: Culicoides haematopotus MALLOCH (Nearctic and Neotropical Regions). E:
Stenoxenus coomani SGUY (Oriental Region). F: Bahiahelea brasiliensis WIRTH (Neotropical Region) (A-
E from Manual of Nearctic Diptera; F from BORKENT, unpubl.).
A. Leptohelea B. Cacaohelea
C. Isthmohelea D. Stilobezzia
Aedeagus
Parameres
Gonocoxite
Gonostylus
Cercus
E. Stilobezzia F. Downeshelea
Figure 20. Male genitalia, ventral view. A: Leptohelea micronyx WIRTH & BLANTON (Neotropical
Region). B: Cacaohelea sp. C: Isthmohelea disjuncta INGRAM & MACFIE (Neotropical region). D:
Stilobezzia sp. E: Stilobezzia sp. (Neotropical Region). F: Downeshelea sp. (A, B, D-F courtesy of
INBio, Costa Rica; C from INGRAM & MACFIE, 1931).
Aedeagus
A. Monohelea B. Echinohelea
Gonostylus
C. Johannsenomyia D. Clastrieromyia
Paramer
E. Amerohelea F. Paryphoconus
Figure 21. Male genitalia, ventral view. A: Monohelea sp. B: Echinohelea sp. C: Johannsenomyia argentata
(LOEW) (Nearctic Region). D: Clastrieromyia dycei SPINELLI & GROGAN (Neotropical Region). E:
Amerohelea frontispina (DOW & TURNER) (Nearctic and Neotropical Regions). F: Paryphoconus
angustipennis ENDERLEIN (Neotropical Region). (A-E courtesy of INBio, Costa Rica; F from
MCALPINE et al., 1981).
79(78) External basal tooth of tarsal claws pointed (Fig. 16A); tarsus of hind leg not
elongate (Fig. 16B); wing with anal lobe poorly developed ........................................
..............................................................................Neobezzia WIRTH & RATANAWORABHAN
External basal tooth of tarsal claws blunt (Fig. 16C); tarsus of hind leg elongate
(Fig. 16D); wing with anal lobe well developed (Fig. 13F) .........................................
...................................................................................... Groganhelea SPINELLI & DIPPOLITO
80(78) Wing with 2 radial cells (Fig. 13G); wing grayish with veins darker brown, anterior
veins prominent; forefemur without ventral thick spines ........ Mallochohelea WIRTH
Wing with 1 radial cell; wing whitish, anterior veins pale; forefemur with ventral
thick spines ...........................................................................................Nilobezzia KIEFFER
81(73) Thorax slender and dorsoventrally flattened; paratergite with numerous small
setae; 1 radial cell (Fig. 12I) ..................................................... Paryphoconus ENDERLEIN
Thorax stouter and not dorsoventrally flattened; paratergite without setae; wing
with 1-2 radial cells .................................................................. PALPOMYIINI..........82
82(81) Abdomen with 1 pair of tergal apodemes arising near lateral margins of sternite 7;
1 spermatheca present, with no evidence of a reduced second one ........................
Abdomen with 1 or more pairs of tergal apodemes (Figs. 23C-E), if only 1 pair
present, these arise from more medially on posterior margin of sternite 7; 1-2
spermathecae but if apparently only one, a small reduced second one is present
.............................................................................................................................................. 83
83(82) Wing with 2 radial cells (Fig. 13B) ............................................................................... 84
Wing with 1 radial cell (Fig. 13C) ................................................................................. 86
84(83) Forefemur without stout spines; hind femur greatly swollen (Fig. 18C); abdomen
white, contrasting with dark thorax ....................................................Pachyhelea WIRTH
Forefemur with or without stout spines (Fig. 18A); hind femur not greatly swollen
(Fig. 18A); abdomen light to dark brown, similar to dark thorax ......................... 85
85(84) Flagellum short, only 1.2-1.4 times longer than the breadth of the head; wing
membrane unpigmented; anal lobe of wing well-developed (Fig. 13I); abdomen
with single pair of apodemes arising from the anterior margin of tergite 7 and
extending to about the anterior margin of tergite 6 (Fig. 23C) .................................
......................................................................................Clastrieromyia SPINELLI & GROGAN
Flagellum elongate, at least 2.5 times longer than the breadth of the head; wing
membrane pigmented; anal lobe of wing poorly developed (Fig. 13B); abdomen
with 2 or more pairs of elongate apodemes arising from the anterior margin of at
least tergites 6-7 and extending much further anteriorly into abdominal cavity
(Fig. 23E) ............................................................................................... Palpomyia MEIGEN
86(83) Costa moderately short with costal ratio 0.67-0.75 (Fig. 13C); forefemur with (Fig.
18B) or without stout spines; tarsomere 5 with or without stout ventral setae .....
....................................................................................................... Bezzia KIEFFER (in part)
Costa more elongate with costal ratio more than 0.80 (Fig. 13D); forefemur
without stout spines; tarsomere 5 with slender ventral setae with sharp, bent tips
(Fig. 16E) ................................................................................ Phaenobezzia HAESELBARTH
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, some
Ceratopogonidae have eggs which are either different from those of all other Diptera
(Dasyhelea, Stenoxenus and Paryphoconus) or are laid in a distinctive matrix (Alluaudomyia and
most Sphaeromiini).
1 Egg C-shaped (Fig. 24A) .................................................................................... Dasyhelea

Egg with, at most, a slight curvature (Figs. 24B-F) .................................................... 2
2(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) ................................................................................................... 3
3(2) Egg oval (Fig. 24B) ............................ Leptoconops (in part), Forcipomyiinae (in part)
Egg elongate, slender (Figs. 24C-E) ............................................................................... 4
4(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 common
genera. A world key to the genera of larvae and pupae is being prepared by the senior
author. It is important to realize that only 11 of 51 known genera of Neotropical
Ceratopogonidae have been described as larvae in this region (Table 2), although an
additional 11 genera are known as larvae from elsewhere.
1 Proleg present on prothorax (Figs. 25B, C); head capsule hypognathous to

prognathous (Figs. 25B, C); with strong setae on head capsule and body (Figs. 25B,
C) (Forcipomyiinae) ........................................................................................................... 2
Without proleg on thorax (Figs. 25A, D-F); head capsule somewhat hypognathous
or prognathous (Figs. 25A, D-F); without strong setae on head capsule and body
anterior to anal segment (Figs. 25A, D-F) .................................................................... 3
2(1) Body somewhat flattened dorsoventrally, with lateral tubercles (Fig. 25B) .............
............................................................................................................................. Atrichopogon
Gonostylus Parameres
Parameres A. Bezzia B. Phaenobezzia
Parameres
C. Mallochohelea D. Nilobezzia
Tenth segment
Cercus
E. Leptoconops F. Macrurohelea
Figure 22. Male genitalia, ventral view. A: Bezzia nobilis (WINNERTZ) (Palaearctic, Nearctic and
Neotropical Regions). B: Phaenobezzia pistiae (INGRAM & MACFIE) (Afrotropical Region). C:
Mallochohelea albibasis (MALLOCH) (Nearctic Region). D: Nilobezzia minor (WIRTH) (Nearctic Region).
Female genitalia, ventral view. E: Leptoconops bequaerti (KIEFFER) (Nearctic and Neotropical Regions).
F: Macrurohelea sp. (Neotropical Region). (A-D from MCALPINE et al., 1981; E courtesy of INBio,
Costa Rica; F from SPINELLI & WIRTH, 1993).
Sternite 8
Sternite 9
Sternite 10
Sternite 8 Cercus
A. Echinohelea B. Probezzia
Tergal apodeme
Tergal apodeme
C. Clastrieromyia
D. Paryphoconus E. Palpomyia
Figure 23. Female genitalia, ventral view. A: Echinohelea sp.(Neotropical Region). B: Probezzia concinna
(MEIGEN) (Holarctic Region). C: Clastrieromyia dycei SPINELLI & GROGAN (Neotropical Region). D:
Paryphoconus angustipennis ENDERLEIN (Neotropical Region). E: Palpomyia lineata (MEIGEN) (Holarctic
Region). (A courtesy of INBio, Costa Rica; B, D-E from MCALPINE et al., 1981).
A. Dasyhelea B. Forcipomyia
C. Culicoides D. Alluaudomyia
E. Probezzia F. Paryphoconus
Figure 24. Eggs. A: Dasyhelea flavifrons (GURIN) (Palearctic Region). B: Forcipomyia mcateei WIRTH
(Nearctic Region). C: Culicoides pulicaris (LINNAEUS) (Palearctic Region). D: Alluaudomyia needhami
THOMSEN (Nearctic Region). E: Probezzia concinna (GURIN) (Palearctic Region). F: Paryphoconus
angustipennis ENDERLEIN (Neotropical Region). (A from SZADZIEWSKI et al., 1997; B from CHAN &
LEROUX, 1965 (as F. saundersi); C from GUTSEVICH & GLUKHOVA, 1970; D from THOMSEN, 1937; E
from WESENBERG-LUND, 1914; F from MAYER, 1952).
A. Leptoconops
Prothoracic proleg
Posterior proleg
B. Atrichopogon
Prothoracic proleg Posterior proleg

C. Forcipomyia
D. Dasyhelea
Posterior proleg
E. Culicoides
F. Bezzia
Figure 25. Larval habitus. A: Leptoconops gallicus CLASTRIER (Palearctic Region), lateral view. B:
Atrichopogon fusculus (COQUILLETT) (Holarctic and Neotropical Regions), lateral view. C: Forcipomyia
sp., lateral view. D: Dasyhelea sp., lateral view. E: Culicoides sp., dorsal view. F: Bezzia sp., dorsal view
(A from CLASTRIER, 1973; B-F from MCALPINE et al., 1981, B as A. polydactylus).
Body cylindrical in cross-section, without lateral tubercles (Fig. 25C) ........ Forcipomyia
3(1) Head capsule directed anteroventrally; end of abdomen with series of hooks,
sometimes arranged as two prolegs, (often retracted and difficult to see in alcohol
material), never with elongate setae (Fig. 25D) ............................................. Dasyhelea
Head capsule prognathous; end of abdomen simple, without hooks, with or
without elongate setae (Figs. 25A, E, F) ....................................................................... 4
4(3) Head capsule somewhat poorly developed pigmented, with well-developed,
posteriorly directed apodemes extending into prothorax (Figs. 26A, B); abdomen
with segments secondarily divided (Fig. 25A) ............................................ Leptoconops
Head capsule well-developed, lacking apodemes posterior to margin of head
capsule (Figs. 26C-F); abdomen with segments normal, not secondarily divided
(Figs. 25E, F) (Ceratopogoninae) ................................................................................... 5
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) ........................... 6
6(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 GNEROS DE ADULTOS DE CERATOPOGONIDAE DE LA REGIN NEOTROPICAL

Existen hasta la fecha 1095 especies de Ceratopogonidae descriptas de la regin Neotropical
(Tabla 5), aunque muchas ms aguardan para a ser descriptas, particularmente en los gneros
Forcipomyia, Atrichopogon, Dasyhelea, Culicoides y Stilobezzia. Especies de esta familia pueden ser
halladas en todas las altitudes y virtualmente en todo hbitat donde exista por lo menos algo
de humedad. La siguiente clave incluye a los gneros conocidos de la regin Neotropical y,
por primera vez, a los machos de todos los gneros (no son conocidos los machos de
Groganhelea, Lanehelea, Notiohelea, Rhynchohelea WIRTH & BLANTON, y Sphaerohelea)
Esta clave est destinada a identificar tanto material conservado en alcohol como
material montado en lmina. Hemos sealado en negrita los caracteres que se observan
mejor mediante el uso de microscopio (por ej., en lminas), aunque con prctica y
paciencia muchos de estos caracteres tambin pueden ser observados en material
conservado en alcohol. En algunos grupos, algunas nervaduras de las alas puedes ser
dificiles de observar; con ejemplares conservados en alcohol, es posible reconocer dbiles
trazos de las nervaduras al examinarlos de perfil a la vez que se inclina levemente el ala,
mientras que con material montado en lmina, la mejor resolucin es la que brinda el
microscopio de contraste de fase.
Cuando las dos uas al final de la pata son del mismo tamao, se las refiere como
iguales; cuando son de diferente tamao, como desiguales. La relacin costal es el largo de
la costa desde la base del ala (medida desde el arculus), dividido por el largo total del ala.
La terminologa adoptada para las nervaduras y celdas de las alas sigue aquella del
Manual of Nearctic Diptera (MCALPINE et al., 1981), con las modificaciones propuestas
Apodemes
A. Leptoconops B. Leptoconops
C. Culicoides D. Culicoides
Seta o
Seta s
Projecting process
E. Nilobezzia F. Nilobezzia
Figure 26. Larval head capsules. A: Leptoconops gallicus CLASTRIER (Palearctic Region), dorsal view. B:
Leptoconops gallicus CLASTRIER (Palearctic Region), ventral view. C: Culicoides reconditus CAMPBELL &
PELHAM-CLINTON (Palearctic Region), dorsal view. D: Culicoides reconditus CAMPBELL & PELHAM-
CLINTON (Palearctic Region), ventral view. E: Nilobezzia formosa (LOEW) (Palearctic Region), dorsal
view. F: Nilobezzia formosa (LOEW) (Palearctic Region), ventral view. (A-B from CLASTRIER, 1973; C-
F from GLUKHOVA, 1979).
por SZADZIEWSKI (1996), y rotuladas conforme a las Figs. 10C, D y 11I. Al igual que otros
dipterlogos, las nervaduras son escritas aqu en letras maysculas y las celdas en letras
minsculas.
1 Ojos ampliamente separados, sin sutura frontal (Fig. 6A); palpo con 4 segmentos
(slo uno ms all del tercero, el cual es alargado o hinchado) y con intervalo
membranoso entre los segmentos 3 y 4+5 (Fig. 9A); ala sin nervadura transversal
r-m (Fig. 10A); hembra con las celdas radiales fusionadas en un estigma (Fig.
10A); sin hileras de setas empalizadas en el primer tarsmero de la pata posterior
(Fig. 14B); gonostilo del macho con una pa 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 dos
ltimos segmentos (Figs. 9B-E); ala con nervadura transversal r-m (Figs. 10B-J);
celdas radiales distintivas o, si estn fusionadas o ausentes (Figs. 11A, F), la
nervadura transversal r-m est claramente presente; con (Fig. 14A) o sin (Fig.
14B) hileras de setas empalizadas en el primer tarsmero de la pata posterior;
gonostilo del macho sin pa alveolada apical (Figs. 19B-F, 20A-F) aunque
el pice 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 en
la membrana; primer tarsmero posterior con setas dispersas o, a lo sumo, con
una hilera de pelos robustos (pero stos no forman un hilera apretada,
especialmente en la base del tarsmero) (Fig. 14B); hembra con dos uas
pequeas, iguales, en cada pata (Figs. 14C-E); con (Figs. 14C, D) o sin (Figs. 14E,
F) empodio bien desarrollado ......................................................................................... 3
Ala desnuda o con macrotriquias dispersas en la mitad distal del ala (unas pocas
especies tienen algunas macrotriquias en la base del ala, pero stas se hallan
dispersas) (Figs. 11E, I); primer tarsmero posterior con hilera de setas con al
menos ms setas basales cuyas bases contactan, algunos grupos con una fuerte
espina basal adicional (Figs. 14A, G-I); hembra con dos uas pequeas, iguales, en
cada pata, o con 1-2 uas en una o ms 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 radiales
bien desarrolladas, con la segunda celda radial larga y abierta moderadamente
(Figs. 10C, D); paratergito bien desarrollado, con una seta fuerte y 0-5 setas finas
ms cortas (Fig. 9F) ........................................................................ Atrichopogon KIEFFER
Costa corta o larga; si es larga, entonces la seguna celda radial es ancha (Figs. 10F,
G) o angosta (Figs. 10B, E); paratergito angosto (como en Fig. 9G) o bien
desarrollado, sin setas o, si hay setas presentes, son al menos 10 e iguales en
tamao y longitud .............................................................................................................. 4
4(3) Flagelmero apical con pezn terminal, el cual es estrechado basalmente (Figs.
7A, B); uas fuertemente curvas (Figs. 14C, D); empodio vestigial o bien
desarrollado (Figs. 14C, D) ........................................................... Forcipomyia MEIGEN
Flagelmero apical sin pezn terminal (aunque puede ser fuertemente

adelgazado) (Fig. 7C); uas slo ligeramente curvas (Fig. 14E); empodio
pobremente desarrollado .............................................................................................. 5
5(4) Ojos anchamente colindantes en la lnea media; flagelmeros antenales sin
sensilas celocnicas; macho con setas cortas en el primer flagelmero,
menores a la mitad del largo de aquellas de los flagelmeros subsiguientes
(7C); flagelmeros del macho esculturados (Fig. 7C); sin espoln en la pata
anterior; membrana del ala sin diseo distintivo de pigmentacin oscura (Fig.
10E) ......................................................................................................... Dasyhelea KIEFFER
Ojos contiguos a anchamente separados en la lnea media (Figs. 6B, C); algunos
flagelmeros antenales con sensilas celocnicas; macho con setas del
primer flagelmero ms o menos del mismo largo que aquellas de los
flagelmeros subsiguientes; flagelmeros del macho no esculturados; con
espoln tibial en la pata anterior; membrana del ala con o sin diseos
distintivos de pigmentacin 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 mandbulas dentadas; membrana alar con o sin
diseos distintivos de pigmentacin oscura y/o clara (Figs. 10F, G) .......................
............................................................................................................... Culicoides LATREILLE
Ala con ambas celdas radiales obliteradas (Fig. 10H); costa no alcanzndo ms all
de 0,43 del largo del ala; hembras con mandbulas reducidas, vestigiales;
membran alar sin diseos distintivos de pigmentacin oscura y/o clara (Fig. 10H)
.............................................................................................................. Paradasyhelea MACFIE
7(2) M bifurcndose ms all de r-m (v.g. bifurcacin mediana peciolada, en algunos de
base muy corta) (Fig. 11I), en algunos la base de M2 no est presente (Figs. 10I, J,
11A) ....................................................................................................................................... 8
M bifurcndose al nivel o anterior de r-m (v.g. bifurcacin mediana ssil, no
peciolada) (Figs. 11G, 12F-J, 13A-I); base de M2 siempre presente ...................... 52
8(7) Ala con celdas radiales reducidas (Figs. 11A, B, D, F); relacin costal menor que
0,65 (Figs. 11A, B, D, F); uas de la hembra ms o menos iguales (el macho de
Rhynchohelea es desconocido, pero probablemente con ojos anchamente separados,
flagelmeros antenales 10 y 11 fusionados, y con una seta katepisternal) ............ 9
Ala con al menos una celda radial bien desarrollada (Figs. 10J, 11E, I); relacin
costal mayor que 0,50; uas de la hembra iguales o desiguales, algunas con slo
una ua en una o ms de las patas ............................................................................... 15
9(8) Macho con los flagelmeros antenales 10 y 11 fusionados o con menos de 13
flagelmeros; ala con o sin celdas radiales; katepisterno con 1 o ms setas
laterales (pueden ser observadas en ejemplares preservados en alcohol,
tomndolos verticalmente y observndolos de perfil a lo largo del plano de la
pleura torcica) (Fig. 9G) ................................................................................................ 10
Antena del macho con flagelmeros 10 y 11 separados y con 13
flagelmeros 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 flagelmeros (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 flagelmeros; proboscis de la hembra normal (Figs.
6B, C, E-G) ........................................................................................................................ 11
11(10) Flagelo del macho con 7-8 flagelmeros (Fig. 8A); palpo con 3 segmentos (Fig.
9C) ...................................................................................... Nannohelea GROGAN & WIRTH
Flagelo del macho con al menos 9 flagelmeros (casi siempre con 13, pero con 10
y 11 fusionados); palpo con 5 segmentos ................................................................... 12
12(11) Anepisterno sin setas; antena del macho con flagelmeros 2-6 fusionados (6-
11 tambin fusionados) .............................................. Brachypogon KIEFFER (en parte)
Anepisterno con al menos 1 seta en el margen posterior (Fig. 9G); antena del
macho con flagelmeros 2-5 articulados, no fusionados (flagelmeros 7-11
fusionados, a veces incluso el flagelmero 6) ................................................................
...................................................................... Ceratoculicoides WIRTH & RATANAWORABHAN
13(9) Palpo con 5 segmentos; largo del ala 0,7-0,8 mm; macho con parmeros bien
desarrollados .............................................................................. Schizonyxhelea CLASTRIER
Palpo con 2 segmentos (Fig. 9B); largo del ala < 0,6 mm; macho sin parmeros
.............................................................................................................................................. 14
14(13) Tibia anterior con espoln apical; macrotriquias confinadas a los mrgenes
distales del ala (en algunos, tambin unas pocas en el pice de M1 y M2); relacin
costal > 0,40 (Fig. 11B); antena del macho con 6 flagelmeros separados (Fig. 8C);
macho con tergitos abdominales 1-3 claros y 4-8 pardos ...........................................
............................................................................................... Baeohelea WIRTH & BLANTON
Tibia anterior sin espoln apical; macrotriquias largas y abundantes sobre todo
el ala; relacin costal ratio < 0,36; antena del macho con 12 flagelmeros, pero
con 2-7, 8-9, y 10-11 fusionados (Fig. 8B); macho con abdomen uniformemente
pardo ........................................................................ Baeodasymyia CLASTRIER & RACCURT
15(8) Ala con diseo caracterstico de una pigmentacin oscura moteada, ms o menos
dispersa sobre la membrana (Fig. 12C); trax y al menos fmur y tibia posteriores
con diseo de pigmentacin; edeago del macho con lazo posteromediano
caracterstico (Fig. 21A) .................................................................... Monohelea KIEFFER1
Ala sin pigmentacin o, si es pigmentada, pigmentacin restricta al sector radial, o
con puntos, rayas o bandas discretas (Figs. 11E, H), o si el patrn de pigmentacin
es substancial (Fig. 11H) entonces trax y todos los fmures y tibias
uniformemente pardos oscuros; otros taxa con trax y patas pigmentados de
manera variable; edeago, si est presente, sin rizo posteromediano16
16(15) Costa extendindose hasta cerca del extremo del ala; costa del ala de la hembra
extendindose ms all del pice de la segunda celda radial (Figs. 12B, D, E)
(extensin tambin 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 en
Downeshelea, pero el macho posee un edeago con lazo posteromediano caracterstico.
Costa no extendindose ms all de 0,85 del largo del ala (Figs. 11E, I); costa del
ala de la hembra finalizando en el pice de la segunda celda radial o extendindose
ms all ligeramente ......................................................................................................... 25
17(16) Halterio en forma de saco, sin constriccin por debajo del capitelo (Fig. 13J) .....
...................................................................................... Diaphanobezzia INGRAM & MACFIE
Halterio con capitelo y pedicelo bien definidos ........................................................ 18
18(17) Macho ................................................................................................................................ .19
Hembra ............................................................................................................................... 21
19(18) Ala con costa no extendindose ms all del pice de la segunda celda radial ......
Ala con costa extendindose apreciablemente ms all del pice de la segunda
celda radial ......................................................................................................................... 20
20(19) Con flagelmeros 10-13 ms alargados que los flagelmeros precedentes;
penacho antenal bien desarrollado; parmeros fusionados en una esfera
voluminosa (Fig. 19F) ............................................................................ Bahiahelea WIRTH
Con flagelmeros 2-13 ms o menos iguales en longitud; penacho antenal corto;
parmeros alargados, delgados ...................................... Leptohelea WIRTH & BLANTON
21(18) Palpo con tres segmentos, ltimo segmento corto y pequeo (Fig. 9D); uas
pequeas, iguales, sin diente basal interno (Fig. 15D) ..................................................
.............................................................................................. Leptohelea WIRTH & BLANTON
Palpo con 4-5 segmentos, ltimo segmento alargado (Fig. 9E); uas variables .....
.............................................................................................................................................. 22
22(21) Clpeo no fusionado a la cabeza (como en la Fig. 6B); ala con la primera celda
radial pequea, segunda alargada (Fig. 12E); con una ua (con diente basal
grueso) en cada pata (Fig. 15E) ............................ Parastilobezzia WIRTH & BLANTON
Clpeo fusionado lateralmente a la cabeza (Figs. 6F, G); ala con 1 celda radial (Fig.
12B); con 2 uas en cada pata, a pesar de que las uas de la pata posterior pueden
ser desiguales ..................................................................................................................... 23
23(22) Ojos moderadamente separados, sin pubescencia (Fig. 6F); uas sin diente basal
interno ................................................................................................ Parabezzia MALLOCH
Ojos anchamente separados, con o sin pubescencia (Fig. 6G); uas con diente
basal interno ...................................................................................................................... 24
24(23) Palpo con 4 segmentos; ojos pubescentes; espermatecas sin cuellos
diferenciados ............................................................... Fittkauhelea WIRTH & BLANTON
Palpo con 5 segmentos; ojos sin pubescencia; espermatecas con cuellos
diferenciados delgados ...................................................................... Bahiahelea WIRTH
25(16) Macho ................................................................................................................................. 26
Hembra ............................................................................................................................... 39
26(25) Pata posterior con una ua larga y diente basal pequeo (Fig. 14H) .......................
................................................................................................................... Allohelea KIEFFER
Pata posterior con dos uas iguales o subiguales ...................................................... 27
27(26) Katepisterno con 1 o ms setas laterales (pueden ser observados en ejemplares
preservados en alcohol tomndolos verticalmente y observndolos de perfil a lo
largo del plano de la pleura torcica) (Fig. 9G) ....... Brachypogon KIEFFER (en parte)
Katepisterno sin setas .................................................................................................. 28

28(27) Ala con 1 celda radial (Fig. 11E) .................................................................................. 29
Ala con 2 celdas radiales ................................................................................................. 33
29(28) Coxa anterior con setas espinosas largas, o setas delgadas afinadas distalmente
(Fig. 9H) ............................................................................................. Parabezzia MALLOCH
Coxa anterior con slo setas cortas, simples (a veces perdidas en ejemplares en
alcohol) ............................................................................................................................... 30
30(29) Ala con manchas pigmentadas, al menos inmediatamente basal a, o sobre r-m y
sobre el pice de la celda radial (Fig. 11E), algunos con pigmentacin adicional
Ala sin manchas pigmentadas ........................................................................................ 31
31(30) Palpo con 5 segmentos; tarsmero 1 de la pata posterior con al menos 1 1/2
hileras de setas empalizadas; edeago compuesto por 2 escleritos laterales (por
lo que el edeago parece estar fisurado en la lnea media) (Fig. 20E) o con
edeago ausente pero con una barra transversa ventral al edeago (Fig. 20D)
..............................................................................................Stilobezzia KIEFFER (en parte)
Palpo con 4 segmentos (como en Fig. 9E); tarsmero 1 de la pata posterior con
slo 1 hilera de setas empalizadas; edeago continuo en la lnea media (como en
Fig. 21C) ............................................................................................................................. 32
32(31) Ojos sin pubescencia; genitalia con parmeros fusionados, muy cortos .........
Ojos pubescentes; genitalia sin parmeros ........ Fittkauhelea WIRTH & BLANTON
33(28) Ala con reas pigmentadas, incluyendo reas poco notorias, discontnuas en M2
and CuA2 (Fig. 12F); edeago continuo en la lnea media (Fig. 20F) ...................
............................................................................................ Downeshelea WIRTH & GROGAN
Ala con o sin diseo de pigmentacin, pero si est presente, entonces sin reas
poco notorias, discontnuas en M2 and CuA2 (porcin posterior del ala puede ser
ms oscura) (Figs. 11J, 12A) y con edeago dividido en la lnea media (Figs. 20B, C)
.............................................................................................................................................. 34
34(33) Antena con los 5 flagelmeros apicales ms alargados que los 8 flagelmeros
basales (antena tipo-hembra), sin penacho ................. Cacaohelea WIRTH & GROGAN
Antena con los 3 flagelmeros apicales ms alargados que los 10 flagelmeros
basales (antena tipo-macho), con o sin penacho ....................................................... 35
35(34). Celdas radiales separadas por una nervadura longitudinal corta (R2); ala sin
macrotriquias; esternito 9 alargado (Fig. 20C) ........... Isthmohelea INGRAM & MACFIE
Las 2 celdas radiales cercanas una de la otra, separadas por una nervadura
transversal corta (R2) (Figs. 11J, 12A); ala con o sin macrotriquias; esternito 9
corto o moderadamente alargado (Figs. 20D, E) ...................................................... 36
36(35) Flagelmeros 3-10 fusionados (Fig. 8D) ............... Borkenthelea SPINELLI & GROGAN
Antena con flagelmeros separados, no fusionados ................................................ 37
37(36) Largo combinado de los tarsmeros 2-5 de la pata posterior aproximadamente
1,4-1,7 veces mayor que el tarsmero 1 ..................... Austrohelea WIRTH & GROGAN
Largo combinado de los tarsmeros 2-5 de la pata posterior aproximadamente del
mismo tamao que el tarsmero 1 ............................................................................... 38
38(37) Tarsmero 1 de la pata posterior con al menos 1 1/2 hileras de setas empalizadas;
ala con o sin macrotriquias; edeago compuesto por 2 escleritos laterales (por
lo que el edeago parece estar fisurado en la lnea media) (Fig. 20E) o con
edeago ausente pero con una barra transversa ventral al edeago (Fig. 20D);
flagelmeros antenales sin sensilas celocnicas .................. Stilobezzia KIEFFER
Tarsmero 1 de la pata posterior con slo 1 hilera de setas empalizadas (como en
Fig. 14A); ala sin macrotrquias; edeago no dividido, ms o menos triangular;
algunos flagelmeros antenales con sensilas celocnicas .................................
39(25) Katepisterno con 1 o ms setas laterales (pueden ser observados en ejemplares
preservados en alcohol tomndolos verticalmente y observndolos de perfil a lo
largo del plano de la pleura torcica) (Fig. 9G) ......... Brachypogon KIEFFER (en parte)
Katepisterno sin setas .................................................................................................. 40
40(39) Ala con 1 celda radial (Figs. 11C, E) ............................................................................ 41
Ala con 2 celdas radiales (la primera puede ser pequea) (Figs. 10I, 11H, I) ........ 43
41(40) Palpo con 4 segmentos (como en Fig. 9E); uas de la pata posterior iguales ........
...................................................... gnero no descripto cercano a Parabezzia MALLOCH
Palpo con 5 segmentos; uas de la pata posterior desiguales o representadas por
un taln (Figs. 14F, 15A-C) ............................................................................................ 42
42(41) Ala con manchas pigmentadas inmediatamente basal a, o sobre r-m y sobre el
pice de la celda radial (Fig. 11E); tarsmero 1 de la pata posterior con slo 1
hilera de setas empalizadas ........................................................... Alluaudomyia KIEFFER
Ala sin manchas pigmentadas; tarsmero 1 de la pata posterior con al menos 1 1/2
hilera de setas empalizadas ............................................. Stilobezzia KIEFFER (en parte)
43(40) Dcimo segmento de la hembra alargado y doblado anteroventralmente (Fig. 22F)
Dcimo segmento de la hembra no alargado, dirigido posteriormente ............... 44
44(43) Alas con reas pigmentadas, con 2-3 bandas oscuras (Fig. 11H), o al menos con
reas pigmentadas en r-m e inmediatamente posterior al pice de la segunda celda
radial .................................................................................................................................... 45
Ala sin reas pigmentadas discretas ............................................................................. 47
45(43) Patas anterior y media, cada una con una ua (cada una con un diente basal ms
pequeo) ............................................................................Stilobezzia KIEFFER (en parte)
Patas anterior y media, cada una con 2 uas iguales ................................................ 46
46(45) Ua de la pata posterior con diente basal (Fig. 14H) .................... Allohelea KIEFFER
Ua de la pata posterior sin diente basal (Fig. 14I) ..... Downeshelea WIRTH & GROGAN
47(44) Patas anterior y media, cada una con 1 ua (algunos con diente basal ms
pequeo) ............................................................................................................................ 48
Patas anterior y media, cada una con uas pares, iguales, o desiguales (como en
Fig. 14F) ............................................................................................................................. 49
48(47) Primera celda radial pequea, R3 engrosada (Fig. 10I); segunda celda radial ms o
menos ancha; nervadura mediana reducida a un trazo poco perceptible en la base;
una ua corta, robusta, puntiaguda en cada pata (Fig. 15F) .......................................
.............................................................................................. Cacaohelea WIRTH & GROGAN
Primera celda radial pequea a bien desarrollada, R3 a lo sumo engrosada en el

pice (Fig. 11I); segunda celda radial angosta; nervadura mediana claramente
visible, pero algunas veces la base de M2 est ausente; ua, de al menos la pata
posterior, alargada (Figs. 15A-C) .................................. Stilobezzia KIEFFER (en parte)
49(47) Celdas radiales separadas por una nervadura longitudinal corta (R2); ala con
macrotriquias escasas, restrictas al pice extremo; flagelmeros antenales sin
sensilas celocnicas ...................................................... Isthmohelea INGRAM & MACFIE
Celdas radiales separadas por una nervadura transversal corta (R2); ala con o sin
macrotriquias en el 1/4 apical; algunos flagelmeros antenales con sensilas
celocnicas (slo en el flagelmero 1 o slo en flagelmeros 8 y 12) ........ 50
50(49) Ala sin macrotriquias; mandbulas reducidas, vestigiales; ojos con o sin
pubescencia; uas pequeas, iguales, en todas las patas ...........................................
............................................................................................... Notiohelea GROGAN & WIRTH
Ala con macrotriquias en el 1/4 apical; mandbulas dentadas; ojos
pubescentes; slo patas anterior y media con uas pequeas, iguales, o uas
desiguales en todas las patas (como en Fig. 14F) ...................................................... 51
51(50) Celdas radiales subiguales, pequeas; R3 engrosada; uas de la pata media
desiguales y ms o menos similares a las de la pata posterior; slo flagelmero 1
con sensilas celocnicas ....................................... Borkenthelea SPINELLI & GROGAN
Segunda celda radial aproximadamente el doble de largo que primera celda radial;
R3 no engrosada; pata media con uas pequeas, iguales, ms cortas que la ua
nica, alargada (con diente basal) de la pata posterior; slo flagelmeros 8 y 12
con sensilas celocnicas ............................................ Austrohelea WIRTH & GROGAN
52(7) Tarsmero 5 de la pata anterior hinchado (Figs. 15G, 17A, C); ................................
................................................................................................. HETEROMYIINI..........53
Tarsmero 5 de la pata anterior no hinchado ............................................................ 56
53(52) M ssil, bifurcndose a nivel de r-m (Fig. 12G) ......... Physohelea GROGAN & WIRTH
M anchamente ssil, bifurcndose anteriormente a r-m (Fig. 12H) ...................... 54
54(53) Fmur anterior hinchado, con numerosas espinas gruesas ventrales (Fig. 17C) ....
........................................................................................................................ Heteromyia SAY
Fmur anterior delgado, a lo sumo con unas pocas espinas delgadas .................. 55
55(54) Tarsmero 4 de todas las patas cilndrico o ligeramente cordiforme, sin espinas
fuertes; pata posterior de la hembra llamativamente alargada, con tarsmero 2 al
memos 3/4 del largo del tarsmero 1 (Fig. 15H); ua posterior muy larga, nica o
con diente basal pequeo, ms larga que las uas de las patas anterior y media (Fig.
15H) .................................................................................................... Pellucidomyia MACFIE
Tarsmero 4 de la pata anterior cordiforme, tarsmeros 4 de las patas media y
posterior bfido y espinoso (Fig. 17B); pata posterior de la hembra no mucho ms
larga que las otras patas, con tarsmero 2 menor que el doble del largo del
tarsmero 1; ambas uas posteriores aproximadamente del mismo largo que
aquellas de las patas anterior y media .............................................. Clinohelea KIEFFER
56(52) Genitalia masculina grande, con gonocoxitos fusionados dorsalmente y
ventralmente (Fig. 21B); genitalia femenina con esternito 8 alargado, laminar (Fig.
23A) ........................................................................................................ Echinohelea MACFIE
Genitalia masculina pequea o grande, si es grande entonces los gonocoxitos no

estn fusionados; genitalia femenina con esternito 8 moderadamente desarrollado
.............................................................................................................................................. 57
57(56) Macho (macho de Groganhelea, Sphaerohelea desconocidos) ...................................... 58
Hembra ............................................................................................................................... 72
58(57) Gonostilo muy reducido en tamao (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 ubicara en este sitio de la clave) .............. 61
59(58) Ala con 2 celdas radiales claramente definidas (Fig. 13I); gonostilo bien definido,
con pice puntiagudo (Fig. 21D) .......................... Clastrieromyia SPINELLI & GROGAN
Ala con 1 celda radial o con, al menos, un indicio poco perceptible de divisin
entre primera y segunda celda radial (Fig. 13D); gonostilo pequeo y redondeado
apicalmente (Fig. 22B) o no claramente definido y fusionado con el gonocoxito
(Fig. 22D) ........................................................................................................................... 60
60(59) Anepisterno con numerosas setas alargadas; gonocoxito corto y gonostilo
presente pero pequeo (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 ms robusta, con gonocoxito y tergito 9 no inusualmente alargados
(Figs. 21F, 22A) ................................................................................................................. 64
62(61) Ala con 1 celda radial (como en Fig. 13D) ...... Neobezzia WIRTH & RATANAWORABHAN
Ala con 2 celdas radiales (Fig. 13G) ............................................................................ 63
63(62) Fmur anterior con espinas guesas ventrales ................... Austrosphaeromias SPINELLI
Fmur anterior sin espinas gruesas ventrales .............................. Mallochohelea WIRTH
(Nota. Los machos de algunas especies de Mallochohelea no tienen genitalia alargada)
64(61) Tarsmere 5 de al menos la pata posterior con espinas robustas, romas (Fig. 17H);
pice del tergito 9 con una hilera de setas gruesas, robustas (Fig. 21C) ..................
....................................................................................................... Johannsenomyia MALLOCH
Tarsmero 5 sin espinas gruesas, o si estn presentes, son ms delgadas y
finamente puntiagudas (Fig. 16A); tergito 9 con setas dispersas, nunca dispuestas
en una hilera de setas gruesas, robustas ...................................................................... 65
65(64) Con 1-2 setas robustas en el pice de los cercos (Fig. 21E); parmeros
fusionados, fusin restricta a la porcin ms anteromedia (Fig. 21E) ............
Con algunas a muchas setas en los cercos; parmeros separados o con
diferente grado de fusin ........................................................................................... 66
66(65) Fmur anterior con espinas robustas (Fig. 18A) ....................................................... 67
Fmur anterior sin espinas robustas ............................................................................ 68
67(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) Trax delgado y aplanado dorsoventralmente; paratergito con numerosas setas

pequeas; 1 celda radial (Fig. 12I) .......................................... Paryphoconus ENDERLEIN
Trax ms robusto y no aplanado dorsoventralmente; paratergito sin setas; 1-2
celdas radiales .................................................................................................................... 70
70(69) Ala con 1 celda radial (Fig. 13C) ........................................ Bezzia KIEFFER (en parte)
Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 71
71(70) Fmur posterior hinchado (Fig. 18C); abdomen blanco, contrastando con el trax
oscuro ........................................................................................................Pachyhelea WIRTH
Fmur posterior no hinchado (Fig. 18A); abdomen claro a pardo oscuro, similar al
trax oscuro ........................................................................ Palpomyia MEIGEN (en parte)
72(57) Ala con M2 fuertemente curvada cerca de la base (Fig. 13A); palpo con 4
segmentos ...................................................................................... Stenoxenus COQUILLETT
Ala con M2 no curvada cerca de la base, o slo ligeramente (Figs. 13B-I); palpo
con 5 segmentos ............................................................................................................... 73
73(72) Tarsmero 5 armado ventralmente con espinas robustas, negras, romas (Figs.
16A, B, 17G-I) (estas espinas algo puntiagudas en Austrosphaeromias); abdomen sin
apodemas tergales; esternito 8 con o sin (slo Sphaerohelea) mechones de
pelos posterolaterales, con setas largas, curvadas, superpuestas en la lnea
media (mejor visin a 100X) (Fig. 23B); ...................... SPHAEROMIINI..........74
Tarsmero 5 inerme, o si existen algunas espinas ventrales, son delgadas y
puntiagudas (Fig. 16E); abdomen con (Figs. 23C-E) o sin apodemas tergales (no
visibles en algunos ejemplares jvenes o pobremente esclerotizados); esternito 8
sin mechones de pelos ................................................................................................ 81
74(73) Fmures sin espinas; uas iguales en la pata anterior, desiguales en las patas media
y posterior (Figs. 17G-I) .......................................................... Johannsenomyia MALLOCH
Fmures con o sin espinas; uas iguales en todas las patas o slo desiguales en la
pata anterior ...................................................................................................................... 75
75(74) Uas suavemente curvas distalmente, con diente basal delgado en el margen
interno ................................................................................................................................ 76
Uas rectas o aplanadas distalmente, con diente basal delgado o romo en el
margen externo (Figs. 16A, B) ...................................................................................... 78
76(75) Fmur anterior hinchado, con numerosas espinas ventrales gruesas ........................
.................................................................................................... Austrosphaeromias SPINELLI
Fmur anterior delgado, inerme ................................................................................... 77
77(76) Costa extendindose hasta el extremo distal del ala; tarsmero 5 con 2-3 pares de
espinas ventrales .............................................. Sphaerohelea SPINELLI & FELIPPE-BAUER
Costa no extendindose ms all de 0,85 del largo del ala; tarsmero 5 con 4-5
pares de espinas ventrales ............................................... Lanehelea WIRTH & BLANTON
78(75) Ala con 1 celda radial; costa larga, extendindose casi hasta el extremo distal del
ala; relacin costal mayor que 0,87; fmures sin espinas ventrales ....................... 79
Ala con 1-2 celdas radiales; costa corta, no llegando a 0,84 del largo del ala;
fmures con o sin espinas ventrales ............................................................................. 80
79(78) Diente basal externo de las uas tarsales puntiagudo (Fig. 16A); tarso de la pata
posterior no alargado (Fig. 16B); ala con lbulo anal pobremente desarrollado ...
..............................................................................Neobezzia WIRTH & RATANAWORABHAN
Diente basal externo de las uas tarsales romo (Fig. 16C); tarso de la pata
posterior alargado (Fig. 16D); ala con lbulo anal bien desarrollado (Fig. 13F) ....
...................................................................................... Groganhelea SPINELLI & DIPPOLITO
80(78) Ala con 2 celdas radiales (Fig. 13G); ala griscea, con nervaduras anteriores pardas
oscuras; fmur anterior sin espinas gruesas ventrales ............... Mallochohelea WIRTH
Ala con 1 celda radial; ala blancuzca, nervaduras anteriores claras; fmur anterior
con espinas gruesas ventrales ...........................................................Nilobezzia KIEFFER
81(73) Trax delgado y aplanado dorsoventralmente; paratergito con numerosas setas
pequeas; 1 celda radial (Fig. 12I) .......................................... Paryphoconus ENDERLEIN
Trax ms robusto y no aplanado dorsoventralmente; paratergito sin setas; ala con
1-2 celdas radiales; ................................................................... PALPOMYIINI..........82
82(81) Abdomen con 1 par de apodemas tergales que surgen cerca de los mrgenes
laterales del esternito 7; 1 espermateca presente, sin trazos ni evidencias de una
segunda reducida ............................................................. Amerohelea GROGAN & WIRTH
Abdomen con 1 o ms pares de apodemas tergales (Figs. 23C-E), si slo est
presente 1 par, estos apodemas surgen ms cerca de la lnea media en el margen
posterior del esternito 7; 1-2 espermatecas, pero si existe slo 1, una segunda
reducida est presente ..................................................................................................... 83
83(82) Ala con 2 celdas radiales (Fig. 13B) ............................................................................. 84
Ala con 1 celda radial (Fig. 13C) .................................................................................. 86
84(83) Fmur anterior sin espinas robustas; fmur posterior muy hinchado (Fig. 18C);
abdomen blanco, contrastando con el trax oscuro ........................Pachyhelea WIRTH
Fmur anterior con o sin espinas robustas (Fig. 18A); fmur posterior no muy
hinchado (Fig. 18A); abdomen claro a pardo oscuro, similar al trax oscuro ........... 85
85(84) Flagelo corto, slo 1,2-1,4 veces ms largo que el ancho de la cabeza; membrana
alar sin pigmentacin; ala con lbulo anal bien desarrollado (Fig. 13I); abdomen
con un par de apodemas que surgen del margen anterior del tergito 7 y se
extienden hasta el margen anterior del tergito 6 (Fig. 23C) ........................................
......................................................................................Clastrieromyia SPINELLI & GROGAN
Flagelo alargado, a menos 2,5 veces ms largo que el ancho de la cabeza;
membrana alar pigmentada; ala con lbulo anal pobremente desarrollado (Fig.
13B); abdomen con 2 o ms pares de apodemas alargados que surgen del margen
anterior de al menos los tergitos 6-7 y se extienden anteriormente dentro de
cavidad abdominal (Fig. 23E) ............................................................ Palpomyia MEIGEN
86(83) Costa moderadamente corta, relacin costal 0,67-0,75 (Fig. 13C); fmur anterior
con (Fig. 18B) o sin espinas robustas; tarsmero 5 con o sin setas ventrales
robustas .................................................................................... Bezzia KIEFFER (en parte)
Costa ms larga, relacin costal ratio mayor que 0,80 (Fig. 13D); fmur anterior
sin espinas robustas; tarsmero 5 con setas ventrales delgadas con extremos
afinados y doblados (Fig. 16E) ........................................... Phaenobezzia HAESELBARTH
CLAVE PARA LOS HUEVOS DE GNEROS DE CERATOPOGONIDAE DE LA REGIN NEOTROPICAL

Solamente han sido descriptos los huevos de pocas especies de ceratopognidos (Table 1).
La clave siguiente es, por lo tanto, muy tentativa.
No se hallan disponibles claves para huevos de Diptera en las cuales los huevos de
Ceratopogonidae pudan ser distinguidos de aquellos pertenecientes a otras familias. Sin
embargo, algunos Ceratopogonidae poseen huevos que son diferentes de todos los dems
(Dasyhelea, Stenoxenus y Paryphoconus) o las oviposiciones tienen lugar en una matriz
caracterstica (Alluaudomyia y la mayora de los Sphaeromiini).
1 Huevo en forma de C (Fig. 24A) ...................................................................... Dasyhelea

Huevo, a lo sumo, con curvatura ligera (Figs. 24B-F) ............................................... 2
2(1) Huevo con collar caracterstico subapical y constriccin terminal a manera de
pezn en el extremo anterior (Fig. 24F) .............................. Stenoxenus y Paryphoconus
Huevo sin ornamentacin en ninguno de los extremos, pequeas espculas
pueden estar presentes (Figs. 24A-E) ............................................................................ 3
3(2) Huevo oval (Fig. 24B) .................. Leptoconops (en parte), Forcipomyiinae (en parte)
Huevo alargado, delgado (Figs. 24C-E) ........................................................................ 4
4(3) Huevos ordenados en una banda alargada, dispuestos en ngulo con respecto al
eje longitudinal de la banda (Fig. 24E) ............................................................................
...... mayora de Sphaeromiini (conocidos para Johannsenomyia, Mallochohelea, Macropeza)
Huevos colocados separadamente o en grupos; si estn colocados en banda, los
huevos 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 GNEROS DE CERATOPOGONIDAE DE LA REGIN NEOTROPICAL

Los inmaduros de Ceratopogonidae se hallan entre los ms pobremente conocidos entre
todas las familias de nematceros (Table 2). Se brinda aqu una clave tentativa para
algunos gneros comunes. El primer autor de esta contribucin est preparando una clave,
a nivel genrico, para larvas y pupas de gneros de todo el Mundo. Es importante destacar
que slo 11 de los 51 gneros conocidos de Ceratopogonidae Neotropicales han sido
descriptos como larvas sobre la base de material de esta regin (Table 2), mientra que 11
otros gneros se conocen como larvas a partir de material de otras regiones.
1 Pseudpodo presente en el protrax (Figs. 25B, C); cpsula ceflica hipognata a

prognata (Figs. 25B, C); con setas fuertes en cpsula ceflica y cuerpo (Figs. 25B,
C) (Forcipomyiinae) ........................................................................................................... 2
Sin pseudpodo en el trax (Figs. 25A, D-F); cpsula ceflica algo hipognata o
prognata (Figs. 25A, D-F); sin setas fuertes en cpsula ceflica y cuerpo
anteriormente al segmento anal (Figs. 25A, D-F) ....................................................... 3
2(1) Cuerpo algo aplanado dorsoventralmente, con tubrculos laterales (Fig. 25B) .....
............................................................................................................................. Atrichopogon
Cuerpo cilndrico en seccin transversal, sin tubrculos laterales (Fig. 25C) .........
............................................................................................................................... Forcipomyia
3(1) Cpsula ceflica dirigida anteroventralmente; finalizacin del abdomen con serie
de ganchos, a veces dispuestos como dos pseudpodos (a menudo retrados y
dificil de observar en ejemplares preservados en alcohol), nunca con setas
alargadas (Fig. 25D) ............................................................................................. Dasyhelea
Cpsula ceflica prognata; finalizacin del abdomen simple, sin ganchos, con o sin
setas alargadas (Figs. 25A, E-F) ...................................................................................... 4
4(3) Cpsula ceflica con pigmentacin pobremente desarrollada, con apodemas bien
desarrollados dirigidos posteriormente y que se extienden en el protrax (Figs.
26A, B); abdomen con segmentos divididos secundariamente (Fig. 25A) ..............
............................................................................................................................... Leptoconops
Cpsula ceflica bien desarrollada, sin apodemas posteriores al margen de cpsula
ceflica (Figs. 26C-F); abdomen con segmentos normales, no divididos
secundariamente (Figs. 25E, F) (Ceratopogoninae) .................................................... 5
5(4) Cpsula ceflica con todas las setas simples (Figs. 26C, D) ............... Culicoides
Cpsula ceflica con una o ms setas s, u y la posterior o plumosa (Figs.
26E, F, seta u no visible, generalmente lateral sobre la cpsula ceflica) .............. 6
6(5) Margen posteroventral de la cpsula ceflica con proceso que se proyecta
posteriormente (Fig. 26F) ........................................................ Sphaeromiini (en parte)
Margen posteroventral de la cpsula ceflica como una lnea recta o algo curvada,
sin proceso que se proyecta posteriormente (Fig. 26E) ..............................................
................................................. Sphaeromiini (en parte), Palpomyiini, Ceratopogonini
8
SYNOPSIS OF THE CERATOPOGONIDAE OF
THE NEOTROPICAL REGION
References providing available keys to the species of the various genera are also given in
the catalog in Table 4.
Allohelea KIEFFER. Only a single species occurs in the Neotropical Region, A. neotropica
WIRTH from Central America and Jamaica, but six species are known from the Nearc-
tic (WIRTH, 1991a). Males are the only members in the family with massive, enlarged
hind legs and a single hind claw, as in the predaceous female.
Alluaudomyia KIEFFER. Nineteen species occur in the Neotropical Region. Of these species,
seventeen were reviewed by SPINELLI & WIRTH (1984c). These small, brightly pat-
terned predaceous midges have larvae that inhabit ponds, lakes and streams where
they are unique in swimming on the surface film.
Amerohelea GROGAN & WIRTH. Eleven species have been recorded from the Neotropical
Region. Of these species, ten were reviewed by GROGAN & WIRTH (1981). Females
differ from other members of the tribe Palpomyiini in having only a single spermath-
eca and a single pair of internal abdominal tergal apodemes that arise near the lateral
margins of segment 7.
Atrichopogon KIEFFER. One hundred species are known from the Neotropical Region with
18 recently described from Costa Rica (BORKENT & PICADO, 2004) and five from 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 the
Neotropical Region and we suspect that hundreds of species remain to be described.
Adults are among the most commonly encountered diurnal biting midges and can be
found on a variety of flowers (mostly small, white or green) or resting on vegetation.
The adult males of at least some species form swarms. Larvae are aquatic to 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 blister
beetles (WIRTH 1956a, b; BORKENT & ROCHA-FILHO, 2006), while others feed on dead
insects, are pollinovorous or are non-feeding (autogenous). The larvae and pupae
have important taxonomic features. EWEN & SAUNDERS (1958) reared nine species
from several localities in the Neotropics.
Austrohelea WIRTH & GROGAN. Only a single species occurs in the Neotropical Region, A.
shannoni WIRTH & BLANTON from the temperate Nothofagus forests of Argentina and
Chile. The genus exhibits a transantarctic distribution, with six other species known
from Australia and New Zealand (WIRTH & GROGAN, 1988). At least three unnamed
species have been collected from Patagonia.
Austrosphaeromias SPINELLI. The genus is known only from two species from southern Ar-
gentina and Chile (SPINELLI, 1997a). It is the only genus of the tribe Sphaeromiini in
the Neotropical Region inhabiting temperate Nothofagus forests.
Baeodasymyia CLASTRIER & RACCURT. Five species of these minute, highly reduced biting
midges are known in the Neotropical Region (BORKENT & CRAIG, 1999). Adults walk
with a distinctive rapid, stuttering gait (best seen with a hand lens). The immatures are
restricted to the headwaters of springs and small seeps.
Baeohelea WIRTH & BLANTON. Only a single species (B. nana WIRTH & BLANTON) of this
minute, highly reduced predaceous genus is known in the Neotropical Region, distrib-
uted from Costa Rica to Ecuador. Baeohelea and its sister group Baeodasymyia are the
only two genera in the family to have their palpi reduced to two segments.
Bezzia KIEFFER. Forty-six species occur in the Neotropical Region. Most species are shining
dark black, but many have striking banded leg patterns. This is the most common
genus in the tribe Palpomyiini in the region and the second most common predaceous
genus after Stilobezzia. The larva of at least one species in Costa Rica lives in 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 temperate
Nothofagus forests of southern Argentina and Chile (SPINELLI & GROGAN, 2001). Males
are unique in the tribe Ceratopogonini in having flagellomeres 3-10 fused.
Brachypogon KIEFFER. Twenty-five species of these small, black midges are known from the
Neotropical Region. Four new species from the Peruvian Amazonia are being described
(SPINELLI & MARINO, in prep.). One undescribed species is common in some rocky coastal
areas in Costa Rica and an additional 11 new species are known from Costa Rica. This
suggests that numerous further species remain to be discovered in this region.
Cacaohelea WIRTH & GROGAN. Only a single described species of these tiny midges is known,
and the original specimens came from a cacao plantation in Costa Rica (WIRTH &
GROGAN, 1988). Further material, including that of a second undescribed species, has
been collected in Costa Rica in areas devoid of cacao.
Ceratoculicoides WIRTH & RATANAWORABHAN. Within the region, only one species is known
from Mexico, with a second, unnamed species from Costa Rica known from two
female adults (HUERTA & BORKENT, 2005). All remaining species are restricted to the
Holarctic Region (WIRTH & GROGAN, 1988).
Clastrieromyia SPINELLI & GROGAN. The genus is known from four Neotropical species (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 the
anterior margin of tergite 7 and extending to about the anterior margin of tergite 6.
Clinohelea KIEFFER. Fourteen species of this predaceous genus occur in the Neotropical
Region (SPINELLI & DURET, 1993). The elegant adults have long, slender legs and large
dark maculations on their wings.
Culicoides LATREILLE. Two hundred and sixty-six species of this vertebrate-biting genus have
been recorded in the region. Of these, only 16 are serious pests of humans or 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 important
ecological 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 some
species 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 recorded
from the Neotropics, but many more remain undescribed. At least eleven new species
are known from Patagonia (DAZ & SPINELLI, in prep.). Adults have reduced, non-
biting mouthparts but are common on many flowers (particularly those that are small
and white) where they feed on nectar. Adults may be common in some coastal 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 grazers
on algae and detritus and are generally present in natural small aquatic habitats (e.g.
tree holes, rock pools) or artificial ones (e.g. cemetery flower-holding containers).
This very diverse and common genus is in urgent need of revision.
Diaphanobezzia INGRAM & MACFIE. The genus is known from four species from Argentinean
Patagonia. The peculiar halter, without a constriction below the knob, is unique within
the family.
Downeshelea WIRTH & GROGAN. Eighteen species occur in the Neotropical Region (LANE &
WIRTH, 1964). They have distinctive large blotches on their wings and may be 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 elongate
antennal 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. amazonica
WIRTH & 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 the
Neotropical Region. Adults of some species are ectoparasites of large insects (as
described in the section on Bionomics above) but those of the subgenus Lasiohelea
feed on vertebrate blood. Although adult females are common and many have biting
mouthparts, the hosts for most of the species remain unknown. The adult males of at
least some species form swarms. Most larvae have elongate dorsal setae (also present
in Atrichopogon), are aquatic to terrestrial, and are found in a diverse array of habitats
such as in mosses, under the bark of rotting trees, inside decaying fruits, and in aquat-
ic epiphytes. Many species remain to be described.
Groganhelea SPINELLI & DIPPOLITO. This genus of the tribe Sphaeromiini is known from a
single species, G. rondoniensis SPINELLI & DIPPOLITO, from Brazilian Amazonia and 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 femora
with dense ventral spines and extremely elongated hind legs, each with a single, very
long claw which they project upward when at rest. Occasionally adults may be abundant.
Isthmohelea INGRAM & MACFIE. The genus is known from a single species, I. disjuncta INGRAM
& MACFIE, from its type-locality, in the Valdivian rain forests of southern Chile. The
male has a strikingly elongate abdominal segment 9.
Johannsenomyia MALLOCH. Only two species are known from the Neotropical Region. The
larvae inhabit rivers and streams.
Lanehelea WIRTH & BLANTON. This genus of the tribe Sphaeromiini is known from two
species from Colombia. Males are unknown.
Leptoconops SKUSE. Twelve species of this vertebrate blood sucking genus occur in the 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 for
females to lay their eggs. The larvae are pink and primarily inhabit wet, protected
sands on beaches of oceans and lakes.
Leptohelea WIRTH & BLANTON. The genus is known from a single female specimen, L. micr-
onyx WIRTH & BLANTON (WIRTH & BLANTON, 1970a), from Colombia and one male
from Costa Rica. The maxillary palpus of adults is 3-segmented. Males are included
here, in a key, for the first time.
Macrurohelea INGRAM & MACFIE. Twelve species occur in the Neotropical Region, particular-
ly in Patagonia. SPINELLI & GROGAN (1990) provided a key to males and females of
eleven of these species. The genus exhibits a transantarctic distribution, being known
from 3 other species from Australia (GROGAN & WIRTH, 1985). Females are unique in
the family in having the tenth segment elongated and bent anteroventrally.
Mallochohelea WIRTH. Six species are known from the Neotropical Region. None are common.
Monohelea KIEFFER. Eighteen species occur in the Neotropical Region, the majority of them
included in a key to males and females by LANE & WIRTH (1964). Adults have wings
with characteristic black and gray maculations, brightly banded legs and females have
single large hind claws.
Nannohelea GROGAN & WIRTH. Only one named species of this minute predaceous genus,
N. clastrieri GROGAN & WIRTH, occurs in the Neotropical Region, from Colombia (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 Neotropical
Region. WIRTH & RATANAWORABHAN (1972a) provided a key to females of seven of
these species.
Nilobezzia KIEFFER. Three species occur in the Neotropical Region. LANE (1961b) provided
a key to males and females. Adults may be locally common, especially around lowland
permanent ponds.
Notiohelea GROGAN & WIRTH. The genus is known from two species from the temperate
Nothofagus forests of southern Argentina and Chile (GROGAN & WIRTH, 1979b; SPINELLI
& GROGAN, 1990). Males are unknown.
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 on
small 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. leei
WIRTH & BLANTON, from Costa Rica and Colombia (WIRTH & GROGAN, 1988).
Paryphoconus ENDERLEIN. Forty species occur in the Neotropical Region. SPINELLI & WIRTH
(1984b) provided a key to females of thirty eight of these species. The adults are
medium-sized to large (female wing lengths 1.5-7 mm) and are generally rare. The few
abundant collections from the Neotropical Region are from near rivers.
Pellucidomyia MACFIE. Four species occur in the Neotropical Region (WIRTH & 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 southern
Argentina and Chile (GROGAN & WIRTH, 1979a). It is the only genus of the tribe
Heteromyiini in the Neotropical Region inhabiting temperate Nothofagus forests.
Rhynchohelea WIRTH & BLANTON. A single species of this monotypic genus, R. monilicornis
WIRTH & BLANTON, is known from Florida and Costa Rica (WIRTH & GROGAN, 1988).
Within the Neotropical Region, this minute relative of Brachypogon is known from only
one female specimen from Costa Rica (BORKENT, 1992).
Schizonyxhelea CLASTRIER. Two species of these small, reduced relatives of Stilobezzia are
known from the Neotropical Region (BORKENT, 2000b). The larvae of one species
were collected from a moderately-sized spring in Costa Rica.
Sphaerohelea SPINELLI & FELIPPE-BAUER. A single species of this monotypic genus, S. biestroi
SPINELLI & FELIPPE-BAUER, is known from northeastern Argentina (SPINELLI & 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 is
closely related to Paryphoconus, the only two genera in the tribe Stenoxenini, and the eggs
of both genera have a terminal nipple-like constriction with a fringed subapical collar.
Female Stenoxenus have a strikingly different M2 wing vein (Fig. 13A) compared to their
males and other members of the family. Adults may be common near rivers.
Stilobezzia KIEFFER. Sixty-four species of these common, often brightly colored and 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 yet
to be discovered in that small country. At least twelve new species in the subgenus
Acanthohelea, from Nothofagus forest in Patagonia, are being described (CARZOLA &
SPINELLI, in prep.).
Undescribed genus near Parabezzia MALLOCH. A single species is known as male and female
adults from Costa Rica. It will be fully described in a future publication.
9
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Proc. Entomol. Soc. Washington 88: 455-460.
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SPINELLI, G.R. & W.L. GROGAN (1994): Two new Neotropical species of the subgenus Isohelea of
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SPINELLI, G.R. & W.L. GROGAN (1998): A revision of the Neotropical predaceous midges of Brachypogon
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WIRTH, W.W. (1951b): New species and records of Virginia Heleidae (Diptera). - Proc. Entomol. Soc.
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WIRTH, W.W. (1953a): Biting midges of the heleid genus Stilobezzia in North America. - Proc. U.S.
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WIRTH, W.W. (1955): Three new species of Culicoides from Texas (Diptera, Heleidae). - J. Washington
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WIRTH, W.W. (1956b): The biting midges ectoparasitic on blister beetles (Diptera, Heleidae). - Proc.
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WIRTH, W.W. (1956c): The heleid midges involved in the pollination of rubber trees in America
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WIRTH, W.W. (1959a): Pachyhelea, a new genus of American Ceratopogonidae related to Palpomyia
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WIRTH, W.W. (1959b): New species and records of Heleidae from Brazil (Diptera). - Deut. Entomol.
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WIRTH, W.W. (1960): The genus Pellucidomyia MACFIE (Diptera, Ceratopogonidae). - Bull. Brooklyn
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WIRTH, W.W. (1962): A reclassification of the Palpomyia-Bezzia-Macropeza groups, and a revision of the
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WIRTH, W.W. (1965a): A new Johannsenomyia from Brazil (Diptera: Ceratopogonidae). - Proc. Entomol.
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WIRTH, W.W. (1965b): A revision of the genus Parabezzia MALLOCH (Diptera, Ceratopogonidae). -
WIRTH, W.W. (1965c): Two new species of Macrurohelea from Chile (Diptera, Ceratopogonidae). -
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WIRTH, W.W. (1966): A new Jamaican blood-sucking midge from lacewings (Diptera, Ceratopogonidae).
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
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WIRTH, W.W. (1971): A review of the stick-ticks, neotropical biting midges of the Forcipomyia
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WIRTH, W.W. (1977): A review of the pathogens and parasites of the biting midges (Diptera:
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WIRTH, W.W. (1978): New species and records of intertidal biting midges of the genus Dasyhelea
KIEFFER from the Gulf of California (Diptera: Ceratopogonidae). - Pacific Insects 18: 191-198.
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WIRTH, W.W. (1983a): The North American predaceous midges of the Bezzia bicolor group (Diptera:
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WIRTH, W.W. (1991a): The predaceous midge genus Allohelea KIEFFER in the western hemisphere
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WIRTH, W.W. (1991b): New and little-known species of Forcipomyia (Diptera: Ceratopogonidae)
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WIRTH, W.W. (1991c): Forcipomyia bicolor and related species of the subgenus Lepidohelea in Brazil
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WIRTH, W.W. (1992): A new genus of Ceratopogonini (Diptera: Ceratopogonidae) from Brazil. -
WIRTH, W.W. (1994a): The subgenus Atrichopogon (Lophomyidium) with a revision of the Nearctic
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WIRTH, W.W. (1994b): The western hemisphere species of the predaceous midge genus Echinohelea,
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WIRTH, W.W. & W.R. ATCHLEY (1973): A review of the North American Leptoconops (Diptera:
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WIRTH, W.W. & P. BARRETO (1978): New species of Culicoides biting midges (Diptera: Ceratopogonidae)
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WIRTH, W.W. & F.S. BLANTON (1953a): Studies in Panama Culicoides (Diptera: Heleidae): I, Descriptions
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WIRTH, W.W. & F.S. BLANTON (1953b): Studies in Panama Culicoides (Diptera, Heleidae). II. Descriptions
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WIRTH, W.W. & F.S. BLANTON (1953c): Studies in Panama Culicoides (Diptera, Heleidae). III. A new
species related to phlebotomus (WILLISTON). - Entomol. News 64: 113-120.
WIRTH, W.W. & F.S. BLANTON (1955a): Studies in Panama Culicoides (Diptera, Heleidae) IV. Descriptions
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W IRTH , W.W. & F.S. B LANTON (1955b): Studies in Panama Culicoides (Diptera, Heleidae) V.
Descriptions of three new species of the subgenus Oecacta POEY. - Bull. Brooklyn Entomol.
Soc. 50: 121-127.
WIRTH, W.W. & F.S. BLANTON (1956a): A new species of salt-marsh sand fly from Florida, the Bahamas,
Panama and Ecuador: its distribution and taxonomic differentiation from Culicoides furens (POEY)
(Diptera, Heleidae). - Fla. Entomol. 39: 157-162.
WIRTH, W.W. & F.S. BLANTON. (1956b): Studies in Panama Culicoides VII. The species of the pulicaris
and cova-garciai group (Diptera, Heleidae). - Proc. Entomol. Soc. Washington 58: 211-227.
WIRTH, W.W. & F.S. BLANTON (1956c): Studies in Panama Culicoides (Diptera, Heleidae). IX. Two new
species related to leoni BARBOSA and reevesi WIRTH. - Bull. Brooklyn Entomol. Soc. 51: 45-52.
WIRTH, W.W. & F.S. BLANTON (1959): Biting midges of the genus Culicoides from Panama (Diptera:
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WIRTH, W.W. & F.S. BLANTON (1967): The North American Culicoides of the guttipennis group (Diptera:
WIRTH, W.W. & F.S. BLANTON (1968a): A revision of the neotropical biting midges of the hylas group
of Culicoides (Diptera: Ceratopogonidae). - Fla. Entomol. 51: 201-215.
WIRTH, W.W. & F.S. BLANTON (1968b): A new Culicoides from Guyana (Diptera: Ceratopogonidae). -
Fla. Entomol. 51: 251-252.
WIRTH, W.W. & F.S. BLANTON (1970a): New genera of Neotropical Ceratopogonidae (Diptera). - Fla.
Entomol. 53: 7-14.
W IRTH , W.W. & F.S. B LANTON (1970b): New species of Neotropical Culicoides (Diptera:
WIRTH, W.W. & F.S. BLANTON (1970c): Notes on Brachypogon KIEFFER (Diptera, Ceratopogonidae), a new
species, and two new Neotropical genera of the tribe Ceratopogonini. - Fla. Entomol. 53: 93-104.
WIRTH, W.W. & F.S. BLANTON (1970d): A review of the Culicoides nigrigenus group, with two new
species (Diptera: Ceratopogonidae). - Entomol. News 81: 141-151.
WIRTH, W.W. & F.S. BLANTON (1971): New Neotropical sandflies of the Culicoides debilipalpis group
(Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 34-43.
WIRTH, W.W. & F.S. BLANTON (1972a): Lanehelea, a new neotropical genus of Sphaeromiini (Diptera:
Ceratopogonidae). - Studia Entomol. 15: 433-438.
WIRTH, W.W. & F.S. BLANTON (1972b): A new Patagonian biting midge of the the genus Monohelea,
Isthmohelea (Diptera: Ceratopogonidae). - Fla. Entomol. 55: 173-176.
WIRTH, W.W. & F.S. BLANTON (1973): A review of the maruins or biting midges of the genus Culicoides
(Diptera: Ceratopogonidae) in the Amazon Basin. - Amazoniana 4: 405-470.
WIRTH, W.W. & F.S. BLANTON (1974a): A new Florida sand fly closely related to Culicoides haematopotus
MALLOCH (Diptera: Ceratopogonidae). - Fla. Entomol. 57: 23-26.
WIRTH, W.W. & F.S. BLANTON (1974b): The West Indian sandflies of the genus Culicoides (Diptera:
Ceratopogonidae). - Agri. Res. Serv., U.S. Dept. Agric., Tech. Bull. 1474: iv + 98 pp.
W IRTH , W.W. & F.S. B LANTON (1978): Two new species of neotropical Culicoides (Diptera:
Ceratopogonidae). - Pan-Pac. Entomol. 54: 236-240.
W IRTH , W.W. & J.L. C ASTNER (1990): New neotropical species of stick-tick (Diptera:
Ceratopogonidae) from katydids. - Fla. Entomol. 73: 157-160.
WIRTH, W.W. & M.I. DOW (1971): Studies on the genus Forcipomyia III. Blantonia, a new subgenus in
the Trichohelea complex (Diptera: Ceratopogonidae). - Fla. Entomol. 54: 289-295.
WIRTH, W.W. & M.I. DOW (1972): Studies on the genus Forcipomyia 4. Rhynchoforcipomyia, a new
neotropical subgenus in the Trichohelea complex (Diptera: Ceratopogonidae). - Ann. Entomol.
Soc. Am. 65: 862-872.
WIRTH, W.W. & A.L. DYCE (1985): The current taxonomic status of the Culicoides vectors of bluetongue
viruses. - In: BARBER, T.L. & M.M. JOCHIM (eds.): 151-162. Bluetongue and related orboviruses.
Alan R. Liss, New York: 746 pp.
WIRTH, W.W. & M.L. FELIPPE-BAUER (1989): The neotropical biting midges related to Culicoides paraensis
(Diptera: Ceratopogonidae). - Mem. Inst. Oswaldo Cruz 84:551-565.
WIRTH, W.W. & W.L. GROGAN (1977): Taxonomic notes on the genus Heteromyia SAY, and a new
species from Nicaragua (Diptera: Ceratopogonidae). - Fla. Entomol. 60: 177-185.
WIRTH, W.W. & W.L. GROGAN (1981): Natural History of Plummers Island, Maryland XXV. Biting
midges (Diptera: Ceratopogonidae). 3. The species of the tribe Stilobezziini. - Bull. Biol. Soc.
WIRTH, W.W. & W.L. GROGAN (1982): The predaceous midges of the genus Phaenobezzia in North
America (Diptera: Ceratopogonidae). - Mem. Entomol. Soc. Washington 10:179-192.
WIRTH, W.W. & W.L. GROGAN (1983): The Nearctic species of the Bezzia bivittata group (Diptera:
Ceratopogonidae). - Proc. Biol. Soc. Washington 96: 489-523.
WIRTH, W.W. & W.L. GROGAN (1988): The predaceous midges of the world (Diptera: Ceratopogonidae;
Tribe Ceratopogonini). - Flora and Fauna Handbook 4. E.J. Brill, Leiden: xv + 160 pp.
WIRTH, W.W. & A.A. HUBERT (1960): Ceratopogonidae (Diptera) reared from cacti, with a review of
the copiosus group of Culicoides. - Ann. Entomol. Soc. Am. 53: 639-658.
WIRTH, W.W. & V.H. LEE (1967): New species of Culicoides from high altitudes in the Colombian
Andes (Diptera: Ceratopogonidae). - Proc. U.S. Nat. Mus. 124: 1-22.
WIRTH, W.W. & D.H. MESSERSMITH (1971): Studies on the genus Forcipomyia. 1. The North American
parasitic midges of the subgenus Trichohelea (Diptera: Ceratopogonidae). - Ann. Entomol. Soc.
Am. 64: 15-26.
WIRTH, W.W. & B.A. MULLENS (1992): Culicoides boydi (Diptera: Ceratopogonidae): a potential vector
of hemorrhagic disease viruses to desert bighorn sheep in southern California. - J. Med. Entomol.
29: 1006-1010.
WIRTH, W.W. & N.C. RATANAWORABHAN (1971a): Ceratoculicoides, a new genus related to Ceratopogon
MEIGEN (Diptera, Ceratopogonidae). - Proc. Entomol. Soc. Washington 73: 170-177.
WIRTH, W.W. & N.C. RATANAWORABHAN (1971b): Notes on Neotropical Pellucidomyia (Diptera,
WIRTH, W.W. & N.C. RATANAWORABHAN (1972a): Neobezzia, a new neotropical biting midge genus of
the tribe Sphaeromiini (Diptera: Ceratopogonidae). - J. Kansas Entomol. Soc. 45: 476-490.
WIRTH, W.W. & N.C. RATANAWORABHAN (1972b): A revision of the tribe Stenoxenini (Diptera:
WIRTH, W.W. & N.C. RATANAWORABHAN (1978): Studies on the genus Forcipomyia. V. Key to subgenera
and description of a new subgenus related to Euprojoannisia BRTHES (Diptera: Ceratopogonidae).
WIRTH, W.W. & S.J. SORIA (1975): A new Neotropical Forcipomyia midge closely related to F. (F.)
genualis (LOEW) (Diptera: Ceratopogonidae). - Rev. Theobroma 5: 19-27.
WIRTH, W.W. & S.J. SORIA (1980): Studies on the genus Forcipomyia VI. The Neotropical species of the
subgenus Warmkea (Diptera: Ceratopogonidae). - Rev. Theobroma 9: 137-161.
WIRTH, W.W. & S.J. SORIA (1981): Two Culicoides biting midges reared from inflorescences of Calathea
in Brazil and Colombia, and a key to the species of the discrepans groups (Diptera:
Ceratopogonidae). - Rev. Theobroma 11: 107-117.
WIRTH, W.W. & G.R. SPINELLI (1992a): Immature stages of Forcipomyia seminole WIRTH and a related
new neotropical species (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 349-356.
WIRTH, W.W. & G.R. SPINELLI (1992b): American predaceous midges of the subgenus Eukraiohelea
of Stilobezzia (Diptera: Ceratopogonidae). - Fla. Entomol. 75: 342-349.
WIRTH, W.W. & G.R. SPINELLI (1993a): The American species of the annulatipes group of the subgenus
Lepidohelea, genus Forcipomyia (Diptera: Ceratopogonidae). - Insecta Mundi 6: 109-125 (1992).
WIRTH, W.W. & G.R. SPINELLI (1993b): The North American species of the Forcipomyia (Lepidohelea)
bicolor subgroup (Diptera: Ceratopogonidae). - Proc. Entomol. Soc. Washington 95: 611-634.
WIRTH, W.W. & W.T. WAUGH (1976): Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae)
associated with culture of Cocoa. - Studia Entomol. 19: 223-236.
WIRTH, W.W. & R.W. WILLIAMS (1957): The biting midges of the Bermuda Islands, with descriptions
of five new species. - Proc. Entomol. Soc. Washington 59: 5-14.
WIRTH, W.W., DYCE, A.L. & G.R. SPINELLI (1988): An atlas of wing photographs, with a summary of
the numerical characters of the Neotropical species of Culicoides (Diptera: Ceratopogonidae). -
Contrib. Am. Entomol. Inst. 25: 1-72.
YOUNG, A.M. (1982): Effects of shade cover and availability of midge breeding sites on pollinating
midge populations and fruit set in two cocoa farms. - J. Appl. Ecol. 19: 47-63.
YOUNG, A.M. (1983): Seasonal differences in abundance and distribution of cocoa-pollinating midges
in relation to flowering and fruit set between shaded and sunny habitats of the La Lola cocoa
farm in Costa Rica. - J. Appl. Ecol. 20: 801-831.
YOUNG, A.M. (1986a): Cocoa Pollination. - Cocoa Growers 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.
10
SUBJECT INDEX *
abercrombyi MACFIE (Forcipomyia) 52 aliciae HUERTA & BORKENT (Ceratoculicoides) 80

acanthodes MACFIE (Bezzia) 31, 94 alleni CLASTRIER & WIRTH (Forcipomyia) 54
Acanthohelea KIEFFER 40, 85, 153 Allohelea KIEFFER 32, 76, 101, 115-116, 139,
acidicola (TOKUNAGA) (Forcipomyia) 57, 125 141, 148
acotylus LUTZ (Culicoides) 71 Alluaudomyia KIEFFER 32, 76, 101, 115-116, 129,
aculeata INGRAM & MACFIE (Palpomyia) 95 140-141, 146, 148
adamsoni MACFIE (Atrichopogon) 44 almeidai (LANE) (Palpomyia) 95
aductus DIPPOLITO & SPINELLI (Stenoxenus) 99 almirantei WIRTH & BLANTON (Culicoides) 74
aegealitis SPINELLI & WIRTH (Dasyhelea) 59 altivolans MACFIE (Atrichopogon) 44
aemulus MACFIE (Paryphoconus) 97 alvarezi ORTZ (Culicoides) 75
aenipes (MACFIE) (Mallochohelea) 91 amapaensis LANE (Paryphoconus) 97
aeria SAUNDERS (Forcipomyia) 26, 59 amazonica CLASTRIER (Stilobezzia) 85
aeronautica MACFIE (Forcipomyia) 58 amazonica SPINELLI & WIRTH (Alluaudomyia) 76
affinis FELIPPE-BAUER & SPINELLI (Monohelea) 83 amazonica WIRTH & BLANTON (Fittkauhelea) 81,
aguirrei TAVARES & SOUZA (Monohelea) 83 104, 150
aitkeni DE MEILLON & WIRTH (Forcipomyia) 55 amazonica WIRTH (Forcipomyia) 54
aitkeni SPINELLI & WIRTH (Bezzia) 93 amazonius MACFIE (Culicoides) 69
aitkeni WIRTH & BLANTON (Culicoides) 67 ameliae BROWNE (Culicoides) 62
aitkeni WIRTH (Echinohelea) 81 americana KIEFFER (Stilobezzia) 86
alahialinus BARBOSA (Culicoides) 70 Amerohelea GROGAN & WIRTH 32, 41, 92, 102, 114,
alambiculorum MACFIE (Culicoides) 65 122, 128, 143, 145, 148
albibasis (MALLOCH) (Mallochohelea) 91, 130 amnicola (MACFIE) (Neobezzia) 30, 91
albicoxa LANE & FORATTINI (Stilobezzia) 86 amnigena (MACFIE) (Stilobezzia) 86
albidiventris (KIEFFER) (Pachyhelea) 95, 152 Amossovia GLUKHOVA 62
albinensis INGRAM & MACFIE (Atrichopogon) 44 ancora (COQUILLETT) (Dasyhelea) 59
albitarsis WIRTH & RATANAWORABHAN (Neobezzia) 91 andensis INGRAM & MACFIE (Dasyhelea) 59
albocincta (Stilobezzia) 86 andicola WIRTH & LEE (Culicoides) 63
albopennis LANE (Clinohelea) 88 andinus WIRTH & LEE (Culicoides) 73
albopicta INGRAM & MACFIE (Dasyhelea) 59 angustipennis ENDERLEIN (Paryphoconus) 31, 97, 113,
albuquerquei LANE (Bezzia) 93 127, 131-132
albuquerquei WIRTH & BLANTON (Culicoides) 70 Anilomyia VARGAS 40, 62
alexanderi WIRTH (Parabezzia) 30, 84 anitae HUERTA & IBAEZ-BERNAL (Forcipomyia) 52
*
Figures in bold
annettae SPINELLI & BORKENT (Culicoides) 27, 67 Austrohelea WIRTH & GROGAN 32, 40, 77, 101,
annulatipes MACFIE (Forcipomyia) 52 116-117, 140, 142, 148
annuliductus WIRTH (Culicoides) 27, 66 austroparaensis SPINELLI (Culicoides) 66
anomalicornis KIEFFER (Paryphoconus) 97 Austrosphaeromias SPINELLI 40, 90, 102, 122-123,
antefurcatus WIRTH & BLANTON (Culicoides) 71 143-144, 149
antennalis (COQUILLETT) (Stilobezzia) 30, 86, 119 Avaritia FOX 63
antequerae (LYNCH ARRIBLZAGA) (Heteromyia) 89 avilaensis ORTZ & MIRSA (Culicoides) 70
antiguensis SAUNDERS (Forcipomyia) 26, 56 azureus WIRTH & BLANTON (Culicoides) 70
antioquiae CLASTRIER & WIRTH (Forcipomyia) 54 bachmanni SPINELLI (Culicoides) 66
antunesi FORATTINI (Culicoides) 73 Baeodasymyia CLASTRIER & RACCURT 32, 40-41, 77,
Apelma KIEFFER 22-27, 56, 58 101, 107, 138, 149
apicalis SPINELLI & WIRTH (Paryphoconus) 97 Baeohelea WIRTH & BLANTON 40-41, 77, 101, 107,
apricans (INGRAM & MACFIE) (Austrosphaeromias) 90 138, 149
apunctipennis SPINELLI & GROGAN (Brachypogon) 78 bahamensis (JOHNSON) (Dasyhelea) 27, 59
aragaoi TAVARES & LUNA DIAS (Culicoides) 66 Bahiahelea WIRTH 40, 78, 101, 110, 115, 138-139
araucana SPINELLI & WIRTH (Bezzia) 93 bahiensis BARBOSA (Culicoides) 64
araucaria SPINELLI (Diaphanobezzia) 80, 114 bahiensis WIRTH & SPINELLI (Forcipomyia) 52
archboldi DE MEILLON & WIRTH (Forcipomyia) 55 bajensis WIRTH (Dasyhelea) 59
archboldi WIRTH & BLANTON (Culicoides) 72 balboa (LANE & WIRTH) (Downeshelea) 80
archboldi WIRTH (Atrichopogon) 47 balsapambensis ORTZ & LEN (Culicoides) 72
archibaldoi TAVARES & SOUZA (Monohelea) 83 balseiroi SPINELLI & GROGAN (Parabezzia) 30, 84
arcuatus WIRTH & RATANAWORABHAN (Stenoxenus) 99 balseiroi SPINELLI (Atrichopogon) 44
arenosa CLASTRIER & RACCURT (Parabezzia) 84 bambusicola LUTZ (Culicoides) 27, 64
argentata (LOEW) (Johannsenomyia) 90, 121, 127 banksi (GERRY) (Nilobezzia) 92
argenteola MACFIE (Forcipomyia) 25, 49 barbatus BORKENT & PICADO (Atrichopogon) 44
argentina LANE & DURET (Clinohelea) 88 barbosai WIRTH & BLANTON (Culicoides) 17, 27, 70
argentinensis KIEFFER (Dasyhelea) 59 barrettoi LANE & DURET (Clinohelea) 88
aridus SPINELLI & MARINO (Atrichopogon) 44 barrettoi LANE (Palpomyia) 95
arubae FOX & HOFFMAN (Culicoides) 17, 27, 75 barrettoi LANE (Paryphoconus) 97
assimilis INGRAM & MACFIE (Atrichopogon) 46 barthi TAVARES & SOUZA (Culicoides) 70
asuturus BORKENT & PICADO (Atrichopogon) 44 basifemoralis WIRTH & SPINELLI (Forcipomyia) 52
ateles (MACFIE) (Phaenobezzia) 97 bassoi RONDEROS & SPINELLI (Leptoconops) 43
atelis WIRTH (Culicoides) 74 batesi LANE (Paryphoconus) 98
atlantica WIRTH & WILLIAMS (Bezzia) 94 batesi WIRTH & BLANTON (Culicoides) 67
atlantis WIRTH & WILLIAMS (Dasyhelea) 27, 59 baueri WIRTH (Forcipomyia) 58
Atrichopogon KIEFFER 15, 18, 20, 22, 24, 32, 44, bayano WIRTH (Culicoides) 27, 66
54, 101, 103, 105, 109, 111, 129, 134, 136, beaveri WIRTH & BARRETO (Culicoides) 71
147-148, 150 beccus BORKENT & PICADO (Atrichopogon) 45
atrichopogon LANE & FORATTINI (Stilobezzia) 85 beckae WIRTH (Stilobezzia) 86
atripalpis WIRTH & BLANTON (Culicoides) 71 beebei FOX (Culicoides) 62
attenuata SAUNDERS (Forcipomyia) 26, 52 bejaranoi DURET & LANE (Heteromyia) 89
aureus ORTZ (Culicoides) 74 belemensis CLASTRIER & WIRTH (Forcipomyia) 54
auricoma KIEFFER (Atrichopogon) 44 belemensis WIRTH & BLANTON (Culicoides) 71
australis WIRTH (Dasyhelea) 59 bella (COQUILLETT) (Alluaudomyia) 29, 76, 112, 118
Beltranmyia VARGAS 63 boliviensis KIEFFER (Palpomyia) 95-96

benarrochi ORTZ & MIRSA (Culicoides) 73 boliviensis SPINELLI & WIRTH (Culicoides) 73
bequaerti (KIEFFER) (Leptoconops) 17, 25, 33, 43, bonaerensis SPINELLI (Brachypogon) 78
109, 130, 151 borgmeieri WIRTH & WAUGH (Dasyhelea) 60
bermudae WIRTH & WILLIAMS (Dasyhelea) 60 borinqueni FOX & HOFFMAN (Culicoides) 28, 65
bermudensis WILLIAMS (Culicoides) 27, 63 Borkenthelea SPINELLI & GROGAN 40, 78, 101, 116-
Bezzia KIEFFER 32, 86, 91-93, 97-98, 102, 122- 117, 140, 142, 149
123, 128, 133, 143-145, 149 borkenti SPINELLI & CAZORLA (Brachypogon) 79
bicellii (LANE) (Mallochohelea) 91 bourioni (CLASTRIER) (Nannohelea) 83, 108
bicinctipes INGRAM & MACFIE (Stilobezzia) 85 Brachyconops WIRTH & ATCHLEY 43
bicolor LANE (Stilobezzia) 30, 86 Brachypogon KIEFFER 32, 78, 101, 107, 115-116,
bicolor LUTZ (Forcipomyia) 52, 59 138-139, 141, 149, 152
bicolor SAUNDERS (Forcipomyia) 26, 58-59 brachyrhyncha WIRTH & DOW (Forcipomyia) 56
bicornis FELIPPE-BAUER & QUINTELAS (Downeshelea) 80 brasiliae (LANE) (Neobezzia) 91
bicuspis BORKENT & PICADO (Atrichopogon) 45 brasiliana CLASTRIER & WIRTH (Forcipomyia) 54
bidentata FELIPPE-BAUER & SPINELLI (Monohelea) 83 brasilianum FORATTINI (Culicoides) 68
biestroi SPINELLI & FELIPPE-BAUER (Sphaerohelea) brasiliensis (LANE) (Nilobezzia) 92
92, 152 brasiliensis (LUTZ) (Leptoconops) 43-44
biestroi SPINELLI & RONDEROS (Culicoides) 67 brasiliensis LANE (Monohelea) 83
biestroi SPINELLI (Alluaudomyia) 29, 76 brasiliensis MACFIE (Atrichopogon) 45
bifida WIRTH & SPINELLI (Forcipomyia) 52 brasiliensis MACFIE (Forcipomyia) 26, 52
bifidus EWEN (Atrichopogon) 25, 45 brasiliensis MACFIE (Palpomyia) 95
bifidus SPINELLI & GROGAN (Brachypogon) 78 brasiliensis MACFIE (Stenoxenus) 99
bimaculata (LOEW) (Clinohelea) 113, 121 brasiliensis SPINELLI & GROGAN (Parabezzia) 84
bimaculata LANE & FORATTINI (Stilobezzia) 86 brasiliensis WIRTH (Bahiahelea) 78, 113, 125
bimaculatus FLOCH & ABONNENC (Culicoides) 68 bredini WIRTH & BLANTON (Culicoides) 65
bimaculatus SPINELLI & GROGAN (Brachypogon) 78 breelandi CLASTRIER & WIRTH (Forcipomyia) 54
birabeni CAVALIERI (Culicoides) 70 brevicornis (KIEFFER) (Bezzia) 30, 93
bispinosa KIEFFER (Stilobezzia) 86 brevilabellata CLASTRIER & WIRTH (Forcipomyia) 54
bivittata (COQUILLETT) (Bezzia) 30, 93 brevipalpis (INGRAM & MACFIE) (Paradasyhelea) 75
blantoni (LANE & WIRTH ) (Johannsenomyia) 90 brevipalpis MACFIE (Atrichopogon) 45
blantoni (LANE & WIRTH) (Downeshelea) 80 bricenoi ORTZ (Culicoides) 70
blantoni (LANE) (Pellucidomyia) 89 broadheadi CLASTRIER & WIRTH (Forcipomyia) 54
blantoni LANE & FORATTINI (Stilobezzia) 86 bromeliae SAUNDERS (Forcipomyia) 25, 49
blantoni SORIA & BYSTRAK (Forcipomyia) 25, 49 bromeliae SPINELLI & WIRTH (Bezzia) 30, 93
blantoni SPINELLI & WIRTH (Bezzia) 30, 93 bromelicola (LUTZ) (Forcipomyia) 26, 55-56
blantoni VARGAS & WIRTH (Culicoides) 64 brownei SPINELLI (Culicoides) 68
blantoni WIRTH & RATANAWORABHAN (Neobezzia) brunnea WIRTH (Parabezzia) 84
91, 114 brunneipennis SPINELLI & WIRTH (Paryphoconus) 98
blantoni WIRTH & RATANAWORABHAN (Stenoxenus) 99 bystraki GROGAN & WIRTH (Parabezzia) 109
blantoni WIRTH (Echinohelea) 81 Cacaohelea WIRTH & GROGAN 40-41, 79, 101, 115,
blantoni WIRTH (Parabezzia) 84 117, 126, 140-141, 149
Blantonia WIRTH & DOW 48 cacaoi WIRTH & WAUGH (Dasyhelea) 60
boliviensis (KIEFFER) (Palpomyia) 96 cacaophila RONDEROS & SPINELLI (Forcipomyia) 52
cacozelus MACFIE (Culicoides) 65 catarinensis MARINO & SPINELLI (Forcipomyia) 51

caerulea SAUNDERS (Forcipomyia) 25, 48 catarinensis SPINELLI & WIRTH (Alluaudomyia) 76
calatheae WIRTH (Forcipomyia) 49 catarinensis SPINELLI & WIRTH (Bezzia) 93
calcarata (COQUILLETT) (Forcipomyia) 25, 49 caucaensis WIRTH & LEE (Culicoides) 72
calchaqui SPINELLI & MARINO (Forcipomyia) 57 caudata INGRAM & MACFIE (Macrurohelea) 82
calchaqui SPINELLI (Brachypogon) 78 caudispina (Dipetalonema) 33
caldasi BROWNE (Culicoides) 72 cayoensis SPINELLI & GROGAN (Parabezzia) 84
caliginosa INGRAM & MACFIE (Forcipomyia) 50 cayoensis SPINELLI & WIRTH (Bezzia) 93
caliginosella WIRTH (Forcipomyia) 50 cebacoi (LANE & WIRTH) (Downeshelea) 80
callangana KIEFFER (Palpomyia) 95 Centrorhynchus LUTZ 51-52, 61, 64
Caloforcipomyia SAUNDERS 48 Ceratobezzia KIEFFER 41, 88, 98
caloptera KIEFFER (Heteromyia) 89 Ceratoculicoides WIRTH & RATANAWORABHAN 41,
calvescens MACFIE (Dasyhelea) 27, 60 101, 107, 109, 138, 149
camposi ORTZ & LEN (Culicoides) 71 Ceratopogon MEIGEN 27, 44, 46-47, 49-51, 54,
cancer HOGUE & WIRTH (Culicoides) 28, 70 56, 59-62, 67-70, 75-76, 79, 83, 86-87, 92-
capitata WIRTH & GROGAN (Bezzia) 93 98, 118
caprilesi FOX (Culicoides) 74 cerifera SAUNDERS (Forcipomyia) 26, 53
caribbea WIRTH & DOW (Forcipomyia) 48 cervicalis (Onchocerca) 33
caribbeana CLASTRIER & RACCURT (Parabezzia) 84 chacoensis SPINELLI & WIRTH (Culicoides) 65
caribbeana SAUNDERS (Forcipomyia) 26, 56 chaconi MACFIE (Stilobezzia) 30, 86
caribbeana SPINELLI & WIRTH (Alluaudomyia) 29, 76 chaquensis DURET & LANE (Heteromyia) 89
caribbeana SPINELLI & WIRTH (Dasyhelea) 60 charrua FELIPPE-BAUER & SPINELLI (Downeshelea) 80
caribbeanus EWEN (Atrichopogon) 25, 45 charruus SPINELLI & Martinez (Culicoides) 28, 68
caribe LANE & FORATTINI (Stilobezzia) 86 chaverrii SPINELLI & BORKENT (Culicoides) 28, 62
caridei (BRTHES) (Culicoides) 17, 64 chiapasi (LANE & WIRTH) (Downeshelea) 80
carioca (TAVARES & SILVA PEREIRA) (Downeshelea) 80 chilensis (PHILIPPI) (Forcipomyia) 51
carioca LANE (Bezzia) 93 chilensis FORATTINI (Leptoconops) 44, 151
carioca LANE (Palpomyia) 95 chilensis GROGAN & WIRTH (Notiohelea) 83
carnatus BORKENT & PICADO (Atrichopogon) 45 chilensis INGRAM & MACFIE (Atrichopogon) 46
carpenteri WIRTH & BLANTON (Culicoides) 71 chilensis INGRAM & MACFIE (Dasyhelea) 60
carpinteroi MARINO & SPINELLI (Atrichopogon) 45 chilensis INGRAM & MACFIE (Palpomyia) 96
carrerai (LANE) (Palpomyia) 95 chilensis SPINELLI & RONDEROS (Bezzia) 93
carrerai LANE (Stenoxenus) 99 chirusi (LANE & WIRTH) (Downeshelea) 80
carsiomelas WIRTH & BLANTON (Culicoides) 71 christopheri BORKENT (Baeodasymyia) 29, 77
carvalhoi WIRTH & BLANTON (Culicoides) 73 chrysonotus WIRTH & BLANTON (Culicoides) 62
casali CAVALIERI & CHIOSSONE (Atrichopogon) 45 cincta (COQUILLETT) (Dasyhelea) 27, 59, 60
casali CAVALIERI & CHIOSSONE (Leptoconops) 44 claripennis LYNCH ARRIBLZAGA (Ceratopogon) 99
castanea LANE (Heteromyia) 89 clastrieri DESSART (Forcipomyia) 55
castanea MACFIE (Palpomyia) 95 clastrieri GROGAN & WIRTH (Nannohelea) 83, 151
castillae FOX (Culicoides) 72 clastrieri SPINELLI & GROGAN (Parabezzia) 84
castneri CLASTRIER & WIRTH (Forcipomyia) 54 Clastrieromyia SPINELLI & GROGAN 32, 41, 95, 102,
castroi (TAVARES & SILVA PEREIRA) (Downeshelea) 80 122, 128, 143, 145, 149
catarina CLASTRIER & WIRTH (Forcipomyia) 54 clavata WILLISTON (Heteromyia) 30, 89
catarinensis LANE (Palpomyia) 95 clavipennis SPINELLI & WIRTH (Bezzia) 93
clavipes WIRTH & RATANAWORABHAN (Neobezzia) 91 Culicoides LATREILLE 17-18, 22, 24, 32-34, 40, 42-
Clinohelea KIEFFER 32, 41, 88, 101, 117, 119, 43, 60-65, 67, 69, 100-101, 103, 105, 133-
142, 149 134, 137, 147, 149, 150
coheni CLASTRIER & WIRTH (Forcipomyia) 54 cummingi SPINELLI & BORKENT (Culicoides) 71
colombiae (LANE & WIRTH) (Downeshelea) 80 cunasi LANE & WIRTH (Monohelea) 83
colombiae WIRTH (Forcipomyia) 58 cylindrica WIRTH & DOW (Forcipomyia) 56
colombiana CLASTRIER & WIRTH (Forcipomyia) 54 cylindricornis WIRTH & BLANTON (Culicoides) 74
colossus BORKENT & PICADO (Atrichopogon) 45 daedaloides WIRTH & BLANTON (Culicoides) 71
columbiana (KIEFFER) (Dasyhelea) 60 daedalus MACFIE (Culicoides) 71
columbiana KIEFFER (Palpomyia) 96 dalcyi GROGAN & WIRTH (Amerohelea) 92
columbianus KIEFFER (Atrichopogon) 45 dalessandroi WIRTH & BARRETO (Culicoides) 70
comechingon SPINELLI & MARINO (Atrichopogon) 45 damascenoi LANE & DURET (Clinohelea) 88
commatis WIRTH & BLANTON (Culicoides) 71 danaisi (FLOCH & ABONNENC) (Forcipomyia) 58
concinna (MEIGEN) (Probezzia) 131-132 darlingtonae WIRTH & BLANTON (Culicoides) 66
concoloripes MACFIE (Bezzia) 94 darwini MARINO & SPINELLI (Forcipomyia) 53
conifera MACFIE (Palpomyia) 96 Dasyhelea KIEFFER 15, 22, 24, 32, 40, 59, 75, 101,
contubernalis ORTZ & LEN (Culicoides) 69 103, 105, 129, 133-134, 137, 146-147, 150
convexipenis WIRTH & SPINELLI (Forcipomyia) 52 dasyophrus MACFIE (Culicoides) 72
coomani SGUY (Stenoxenus) 113-114, 121, 125 davidi SPINELLI (Culicoides) 68
copanensis UTMAR & WIRTH (Forcipomyia) 48 daviesi WIRTH & BLANTON (Culicoides) 70
copiosus ROOT & HOFFMAN (Culicoides) 65, 111 deanei FELIPPE-BAUER & QUINTELAS (Downeshelea) 80
coquilletti KIEFFER (Stilobezzia) 30, 85-86 deanei FELIPPE-BAUER & WIRTH (Culicoides) 75
coracina KIEFFER (Stilobezzia) 87 debilipalpis LUTZ (Culicoides) 17, 28, 66
cornuta SAUNDERS (Forcipomyia) 26, 52 decor (WILLISTON) (Culicoides) 62
coroicoensis WIRTH (Palpomyia) 96 delpontei CAVALIERI & CHIOSSONE (Atrichopogon) 45
correntina DURET & LANE (Heteromyia) 89 delpontei CAVALIERI (Forcipomyia) 50
correntina RONDEROS & DAZ (Dasyhelea) 27, 60 denisae CLASTRIER (Culicoides) 66
costalis MACFIE (Atrichopogon) 45 dentifemur SPINELLI & WIRTH (Bezzia) 93
costalis WIRTH (Parabezzia) 84 depilis MACFIE (Atrichopogon) 45
costaricae MACFIE (Atrichopogon) 45 desutterae CLASTRIER & WIRTH (Forcipomyia) 54
costaricae WIRTH & RATANAWORABHAN (Neobezzia) 91 diabolicus HOFFMAN (Culicoides) 68
Cotocripus BRTHES 40, 64 Diaphanobezzia INGRAM & MACFIE 40, 80, 101,
coutinhoi BARRETTO (Culicoides) 68 110, 139, 150
covagarciai ORTZ (Culicoides) 62 Dicrohelea KIEFFER 90
crassicrus KIEFFER (Palpomyia) 96 dicrourus WIRTH & BLANTON (Culicoides) 28, 70
crepuscularis MALLOCH (Culicoides) 28, 63 Didymophleps WEYENBERGH 99, 102
crescentis WIRTH & BLANTON (Culicoides) 71 didymothecae MACFIE (Atrichopogon) 45
cristata CLASTRIER & WIRTH (Forcipomyia) 54 diffusus SPINELLI (Culicoides) 68
crucifer CLASTRIER (Culicoides) 66 diminuta LANE & FORATTINI (Stilobezzia) 87
crudelis (MAYER) (Forcipomyia) 53-54 diminutus BARBOSA (Culicoides) 69
crudelis KNAB (Forcipomyia) 54 dimorphus KIEFFER (Stenoxenus) 99
cryptogamus MACFIE (Atrichopogon) 45 Diphaomyia VARGAS 64
cuacuahuitlus HUERTA & BORKENT (Brachypogon) 79 discoloripes MACFIE (Forcipomyia) 52
cuiabai WIRTH (Culicoides) 70 discrepans ORTZ & MIRSA (Culicoides) 70
disjuncta INGRAM & MACFIE (Isthmohelea) 82, 126, 151 equatoriensis BARBOSA (Culicoides) 66
distispinulosa SPINELLI & WIRTH (Alluaudomyia) 29, 76 eriophora (WILLISTON) (Forcipomyia) 54
diversus FELIPPE-BAUER (Culicoides) 66 esmeralda LANE & FORATTINI (Stilobezzia) 87
dominicana DE MEILLON & WIRTH (Forcipomyia) 26, 56 espinolai F ELIPPE -B AUER & L OURENCO - DE -
dominicanus WIRTH & BLANTON (Culicoides) 62 OLIVEIRA (Culicoides) 66
dominicii ORTZ (Culicoides) 71 esteparia MARINO & SPINELLI (Forcipomyia) 49
domizii SPINELLI (Atrichopogon) 45 estevezae RONDEROS & SPINELLI (Culicoides) 72
donajii VARGAS (Culicoides) 72 estevezae SPINELLI & WIRTH (Alluaudomyia) 76
donskoffi CLASTRIER & WIRTH (Forcipomyia) 54 ethelae SPINELLI & GROGAN (Brachypogon) 78
dorsalis WIRTH & DOW (Forcipomyia) 56 eublepharus MACFIE (Culicoides) 72
dorsofasciata (LUTZ) (Stilobezzia) 30, 86 eucnemus MACFIE (Atrichopogon) 45
dowi BYSTRAK & WIRTH (Forcipomyia) 49 eukosma MACFIE (Forcipomyia) 48
Downeshelea WIRTH & GROGAN 80, 101, 110, 113, Eukraiohelea INGRAM & MACFIE 86
115-116, 118, 126, 138, 140-141, 150 Euprojoannisia BRTHES 49
dryadum MACFIE (Stilobezzia) 87 euthystyla WIRTH & SPINELLI (Forcipomyia) 52
Drymodesmyia VARGAS 65 evansi WIRTH & BLANTON (Culicoides) 64
duartei TAVARES & LUNA DIAS (Culicoides) 73 excentricus LANE (Stenoxenus) 99
dubia MACFIE (Forcipomyia) 52 fairchildi LANE & WIRTH (Monohelea) 83, 110, 138
dubitans LANE (Stilobezzia) 30, 87 fairchildi WIRTH & BLANTON (Culicoides) 71
dunklei CLASTRIER & WIRTH (Forcipomyia) 54 fairfaxensis WIRTH (Forcipomyia) 111
dunni WIRTH & BLANTON (Culicoides) 71 falcifera SAUNDERS (Forcipomyia) 25, 49
dureti LANE & FORATTINI (Stilobezzia) 87 fallax KIEFFER (Clinohelea) 88
dureti RONDEROS & SPINELLI (Culicoides) 66 farri WIRTH & BLANTON (Culicoides) 62
dycei SPINELLI & GROGAN (Clastrieromyia) 31, 95, farri WIRTH (Forcipomyia) 56
127, 131 fasciata GROGAN & WIRTH (Amerohelea) 92
eadsi WIRTH & BLANTON (Culicoides) 66 fasciata SAY (Heteromyia) 89, 121
echinodes MACFIE (Atrichopogon) 45 fasciatus (WALKER) (Heteromyia) 89
Echinohelea MACFIE 32, 40, 81, 101, 104, 122, 127, felippebauerae CLASTRIER & WIRTH (Forcipomyia) 54
131, 142, 150 femoralis LANE & FORATTINI (Stilobezzia) 87
Echinoideshelea WIRTH 81 fernandezi ORTZ (Culicoides) 72
ecuadorensis SPINELLI & GROGAN (Brachypogon) 78 fernandoi TAVARES & SOUZA (Culicoides) 68
ecuadorensis SPINELLI & WIRTH (Paryphoconus) 98 ferreyrai RONDEROS & SPINELLI (Culicoides) 68
edeni WIRTH & BLANTON (Culicoides) 28, 64 fiebrigi KIEFFER (Atrichopogon) 45
edmistoni WIRTH & SPINELLI (Forcipomyia) 26, 52 fiebrigi KIEFFER (Stilobezzia) 30, 87
edwardsi (SAUNDERS) (Forcipomyia) 26, 50, 56 fieldi WIRTH & BLANTON (Culicoides) 73
edwardsi INGRAM & MACFIE (Forcipomyia) 50 filarifer HOFFMAN (Culicoides) 22, 33, 68
edwardsi INGRAM & MACFIE (Stilobezzia) 85 filibranchia (LUTZ) (Dasyhelea) 27, 60
edwardsiana WIRTH (Forcipomyia) 50 filiducta SPINELLI & WIRTH (Bezzia) 93
efferus FOX (Culicoides) 62 filiductus WIRTH (Culicoides) 66, 93
eldridgei WIRTH & BARRETO (Culicoides) 66 fimbriatus MACFIE (Atrichopogon) 45
elegantula (JOHANNSEN) (Stilobezzia) 86 Fittkauhelea WIRTH & BLANTON 41, 81, 101, 110,
elutus MACFIE (Culicoides) 64 115, 139-140, 150
endemicus SPINELLI & MARINO (Atrichopogon) 45 fittkaui SPINELLI & WIRTH (Alluaudomyia) 76
enderleini LANE (Paryphoconus) 98 fittkaui SPINELLI & WIRTH (Paryphoconus) 98
fittkaui WIRTH & BLANTON (Culicoides) 74 fusca SPINELLI & WIRTH (Bezzia) 93
fittkaui WIRTH & RATANAWORABHAN (Neobezzia) 91 fuscipennis (LANE & WIRTH) (Downeshelea) 80
flava (WILLISTON) (Forcipomyia) 51 fuscipennis SPINELLI & GROGAN (Macrurohelea) 82
flavicauda MACFIE (Dasyhelea) 27, 60 fuscipennis WIRTH (Parabezzia) 84
flavicaudae MACFIE (Atrichopogon) 45 fusciradialis SPINELLI & WIRTH (Paryphoconus) 98
flavidus (JOHANNSEN) (Paryphoconus) 31, 98 fuscivenosus (LUTZ) (Brachypogon) 79
flavifemoris MACFIE (Forcipomyia) 53 fusculus (COQUILLETT) (Atrichopogon) 25, 47, 133
flavifrons (GURIN) (Dasyhelea) 132 fuscus LANE (Paryphoconus) 98
flavipes LUTZ (Atrichopogon) 45, 95 fusicornis (COQUILLETT) (Forcipomyia) 56, 118
flavivenulus COSTA LIMA (Culicoides) 68 fusipalpis WIRTH & BLANTON (Culicoides) 68
flinti SPINELLI & WIRTH (Bezzia) 93 gabaldoni ORTZ (Culicoides) 73
flinti SPINELLI & WIRTH (Paryphoconus) 98 galapagensis (COQUILLETT) (Forcipomyia) 54
flinti WIRTH (Culicoides) 66 galindoi GROGAN & WIRTH (Amerohelea) 92
flochabonnenci ORTZ & MIRSA (Culicoides) 72 galindoi WIRTH & BLANTON (Culicoides) 73
florenciae MESSERSMITH (Culicoides) 72 galindoi WIRTH & SORIA (Forcipomyia) 59
floridensis BECK (Culicoides) 28, 75 galliarii MARINO & SPINELLI (Forcipomyia) 49
floridensis DOW & WIRTH (Forcipomyia) 57 gallicus CLASTRIER (Leptoconops) 133, 135
floridensis WIRTH (Leptoconops) 44, 104 gamboai BORKENT & PICADO (Atrichopogon) 45
fluminensis FELIPPE-BAUER & QUINTELAS gentilii SPINELLI & GROGAN (Macrurohelea) 82
(Downeshelea) 80 genualis (LOEW) (Forcipomyia) 25, 50
fluminensis LANE (Bezzia) 93 germanus MACFIE (Culicoides) 66
fluminensis LANE (Stilobezzia) 87 gibbera (COQUILLETT) (Bezzia) 93
flumineus MACFIE (Atrichopogon) 45 gibsoni WIRTH (Culicoides) 72
fluvialis MACFIE (Culicoides) 72 ginesi ORTZ (Culicoides) 66
fluviatilis (LUTZ) (Culicoides) 75 glabellus WIRTH & BLANTON (Culicoides) 73
forattinii ORTZ (Culicoides) 74 glaber MACFIE (Atrichopogon) 45
forattinii WIRTH & GROGAN (Schizonyxhelea) 30, glabra (COQUILLETT) (Bezzia) 30, 93
85, 112 glabrior MACFIE (Culicoides) 66
Forcipomyia MEIGEN 15, 17, 18, 19-21, 22, 24, 32- Glaphiromyia VARGAS 65
33, 40, 47-49, 53, 55-58, 101-103, 105, 118, glauca MACFIE (Forcipomyia) 25, 48, 106
133, 134, 136, 147-148, 150 glauca MACFIE (Stilobezzia) 30, 87-88
formosa (LOEW) (Nilobezzia) 135 globulifer MACFIE (Atrichopogon) 45
foxi ORTZ (Culicoides) 17, 68 globulosa SPINELLI & WIRTH (Bezzia) 93
franklini CLASTRIER & WIRTH (Forcipomyia) 54 goeldii WIRTH & BLANTON (Culicoides) 74
franklini SPINELLI (Culicoides) 68 goianensis LANE (Bezzia) 93
freitasi WIRTH & BLANTON (Culicoides) 64 goianensis LANE (Paryphoconus) 98
frontispina (DOW & TURNER) (Amerohelea) 92, 127 gordoni MACFIE (Atrichopogon) 45
fuliginosa (MEIGEN) (Forcipomyia) 26, 53-54 gorgasi WIRTH & BLANTON (Culicoides) 70
fulvus JOHANNSEN (Stenoxenus) 99 gracile (Dipetalonema) 33
fur (JOHNSON) (Forcipomyia) 56 grahambelli FORATTINI (Culicoides) 66
furcifera MACFIE (Forcipomyia) 48 grandcolasi CLASTRIER & WIRTH (Forcipomyia) 54
furens (POEY) (Culicoides) 17, 28, 33, 70, 150 grandis LANE & FORATTINI (Stilobezzia) 87
furva INGRAM & MACFIE (Stilobezzia) 85 grandis MACFIE (Paryphoconus) 98
fusca (PHILIPPI) (Forcipomyia) 40, 50 granditergitus BORKENT & PICADO (Atrichopogon) 45
granditibialis BORKENT & PICADO (Atrichopogon) 45 hieroglyphica KIEFFER (Monohelea) 82-83, 113
griseola WIRTH (Dasyhelea) 60 hippolytae MACFIE (Dasyhelea) 60
Groganhelea SPINELLI & DIPPOLITO 41, 90, 102- hirsuta INGRAM & MACFIE (Stilobezzia) 85
103, 122, 128, 134, 143, 145, 150 hirtipes (KIEFFER) (Dasyhelea) 60
grogani SPINELLI & WIRTH (Bezzia) 94 hispida BORKENT (Palpomyia) 96
guadeloupensis DELCOLLE & RIEB (Dasyhelea) 60 hitchcocki SPINELLI & WIRTH (Culicoides) 65
guadeloupensis FLOCH & ABONNENC (Culicoides) 72 hobbsi WIRTH & SPINELLI (Forcipomyia) 53
guaimiesi LANE & WIRTH (Monohelea) 83 hoffmani FOX (Culicoides) 28, 67
guamai WIRTH & BLANTON (Culicoides) 74 Hoffmania FOX 67
guamai WIRTH & DOW (Forcipomyia) 57 Holoconops KIEFFER 43
guarani LANE (Palpomyia) 31, 96 homofacies SPINELLI (Atrichopogon) 46
guarani MARINO & SPINELLI (Forcipomyia) 50 hondurensis HOFFMAN (Leptoconops) 43
guarani RONDEROS & SPINELLI (Culicoides) 72 hondurensis SPINELLI & BORKENT (Culicoides) 64
guarani RONDEROS & SPINELLI (Forcipomyia) 52 hondurensis SPINELLI & GROGAN (Parabezzia) 84
guarani SPINELLI (Alluaudomyia) 76 hondurensis SPINELLI & WIRTH (Bezzia) 94
guerrai WIRTH & BLANTON (Culicoides) 72 hondurensis SPINELLI & WIRTH (Dasyhelea) 60
guianae (MACFIE) (Stilobezzia) 85 horacioi LANE (Clinohelea) 30, 88
guianae (WIRTH) (Downeshelea) 80 horticola LUTZ (Culicoides) 67
guianae MACFIE (Paryphoconus) 98 hortorum WEYENBERGH (Didymophleps) 99
guianensis MACFIE (Atrichopogon) 45 hugoi SPINELLI & GROGAN (Brachypogon) 79
gustavoi BORKENT (Baeodasymyia) 77 hylas MACFIE (Culicoides) 28, 67-68
guttatus (COQUILLETT) (Culicoides) 67-68 iberaensis SPINELLI & CAZORLA (Palpomyia) 96
gutturosa (Onchocerca ) 33 ichesi RONDEROS & SPINELLI (Culicoides) 66
guyana CLASTRIER & WIRTH (Forcipomyia) 54 ignacioi FORATTINI (Culicoides) 68
guyana CLASTRIER (Palpomyia) 96 imitator ORTZ (Culicoides) 67
guyana CLASTRIER (Schizonyxhelea) 85 impar (JOHANNSEN) (Brachypogon) 79
guyanensis FLOCH & ABONNENC (Culicoides) 28, 74 impusilloides SPINELLI & WIRTH (Culicoides) 63
Haematomyidium GOELDI 66-67 inacayali SPINELLI & MARINO (Atrichopogon) 46
haematopotus MALLOCH (Culicoides) 28, 64, 125 inaequalis SPINELLI & GROGAN (Parabezzia) 84
haitiensis CLASTRIER & RACCURT (Parabezzia) 84 inamollae FOX & HOFFMAN (Culicoides) 67-68
haitiensis DELCOLLE (Culicoides) 65 incubans (MACFIE) (Forcipomyia) 56
harii SPINELLI & GROGAN (Borkenthelea) 78, 108 incultus EWEN (Atrichopogon) 25, 46
harpa SPINELLI & BORKENT (Forcipomyia) 57 inermicollis KIEFFER (Palpomyia) 96
harpegonata WIRTH & SORIA (Forcipomyia) 25, 50 ingrami SPINELLI & GROGAN (Paradasyhelea) 75, 111
harrisi MACFIE (Atrichopogon) 45 insigniforceps MACFIE (Forcipomyia) 52
Hartomyia MALLOCH 85 insignipalpis MACFIE (Atrichopogon) 46
hatoensis UTMAR & WIRTH (Forcipomyia) 48 insignipalpis MACFIE (Forcipomyia) 54
hayesi MATTA (Culicoides) 62 insignis LUTZ (Culicoides) 18, 22, 29, 33, 67-68, 111
heliconiae FOX & HOFFMAN (Culicoides) 68 insigniventris MACFIE (Atrichopogon) 46
herediae WIRTH & SPINELLI (Forcipomyia) 26, 53 insinuatus ORTZ & LEN (Culicoides) 17, 67
hermani SPINELLI & BORKENT (Culicoides) 63 insularis SPINELLI & GROGAN (Brachypogon) 79
hermosa UTMAR & WIRTH (Forcipomyia) 48 insularis SPINELLI & GROGAN (Palpomyia) 96
hertigi WIRTH & BLANTON (Culicoides) 65 intermedia SAUNDERS (Forcipomyia) 26, 52
Heteromyia SAY 32, 89, 101, 117, 142, 151-152 intrepida MACFIE (Forcipomyia) 58
iquitosensis CLASTRIER & WIRTH (Forcipomyia) 54 lamprogaster EDWARDS (Heteromyia) 89

iriartei FOX (Culicoides) 64 Lanehelea WIRTH & BLANTON 41, 90, 102-103, 122-
irwini GROGAN & WIRTH (Macrurohelea) 82 123, 134, 143-144, 151
irwini SPINELLI & WIRTH (Culicoides) 64 lanei ORTZ (Culicoides) 74
Isoecacta GARRETT 76 lanei WIRTH & RATANAWORABHAN (Pellucidomyia) 89
Isohelea KIEFFER 79 lanei WIRTH & RATANAWORABHAN (Stenoxenus) 99
Isthmohelea INGRAM & MACFIE 40, 82, 101, 115, lanei WIRTH (Echinohelea) 29, 81, 112, 118
117, 140, 142, 151 lanei WIRTH (Johannsenomyia) 90
ixodoides (FIEBRIG-GERTZ) (Forcipomyia) 18, 53-54 lanei WIRTH (Monohelea) 83
jamaicensis EDWARDS (Culicoides) 65 lanei WIRTH (Paryphoconus) 31, 98
jamaicensis SPINELLI & WIRTH (Bezzia) 94 Lasiohelea KIEFFER 17, 20, 22, 26, 51-52, 58-59, 150
jamaicensis SPINELLI & WIRTH (Dasyhelea) 60 latifemoris (INGRAM & MACFIE) (Pachyhelea) 95
jamaicensis WIRTH (Echinohelea) 81 latipennis SPINELLI & WIRTH (Paryphoconus) 98
jamaicensis WIRTH (Forcipomyia) 58 leei SPINELLI & WIRTH (Alluaudomyia) 76
jamaicensis WIRTH (Parabezzia) 84 leei SPINELLI & WIRTH (Bezzia) 94
jipajapae WIRTH (Forcipomyia) 58 leei SPINELLI & WIRTH (Paryphoconus) 98
jocosa SAUNDERS (Forcipomyia) 26, 56 leei WIRTH & BLANTON (Lanehelea) 90-91
johannseni (LANE) (Palpomyia) 96 leei WIRTH & BLANTON (Parastilobezzia) 84, 113,
johannseni (WIRTH) (Allohelea) 112 119, 152
Johannsenomyia MALLOCH 32, 90-91, 95, 102, 122- leei WIRTH & RATANAWORABHAN (Forcipomyia) 57
123, 129, 143-144, 146, 151 leei WIRTH (Echinohelea) 81
johnsoni COQUILLETT (Stenoxenus) 31, 99 lenkoi LANE (Bezzia) 94
jubata SPINELLI & WIRTH (Bezzia) 94 lenti TAVARES & LUNA DIAS (Culicoides) 70
jurbergi FELIPPE-BAUER (Culicoides) 64 leoni BARBOSA (Culicoides) 73
kampa FELIPPE-BAUER (Culicoides) 67 leopoldoi ORTZ (Culicoides) 72
kawensis CLASTRIER & WIRTH (Forcipomyia) 54 Lepidohelea KIEFFER 26, 48, 52
keilini (SAUNDERS) (Forcipomyia) 26, 56 Leptoconops SKUSE 16-17, 22, 25, 32-33, 43-44, 101,
kiefferi LANE (Heteromyia) 89 103, 111, 129, 134, 136, 146-147, 151
kiefferi LANE (Paryphoconus) 98 Leptohelea WIRTH & BLANTON 40-41, 82, 101, 110,
kiefferi LANE (Stilobezzia) 87 138-139, 151
kintzi WIRTH & BLANTON (Culicoides) 74 lesliei WIRTH (Forcipomyia) 26, 58-59
knowltoni BECK (Culicoides) 63 levis (COQUILLETT) (Atrichopogon) 44, 111, 118
knowltoni CLASTRIER & WIRTH (Leptoconops) 43 lichyi FLOCH & ABONNENC (Culicoides) 73
koenigi DELCOLLE & RIEB (Dasyhelea) 60 limai BARRETTO (Culicoides) 73
kremeri SPINELLI & GROGAN (Clastrieromyia) 95 limitrofe SPINELLI & FELIPPE-BAUER (Mallochohelea) 91
kuanoskeles MACFIE (Forcipomyia) 53 limnetis INGRAM & MACFIE (Forcipomyia) 58
kuscheli WIRTH & BLANTON (Culicoides) 74 limnochares (MACFIE) (Palpomyia) 96
kuscheli WIRTH (Macrurohelea) 82 limonensis ORTZ & LEN (Culicoides) 67
lacajae MACFIE (Atrichopogon) 46 limpidus WIRTH & RATANAWORABHAN (Stenoxenus) 99
lacrimatorii MACFIE (Forcipomyia) 53 lineata (MEIGEN) (Palpomyia) 131
lacustris INGRAM & MACFIE (Dasyhelea) 27, 60 lituratus (WILLISTON) (Atrichopogon) 46
lacustris LANE (Palpomyia) 31, 96 lobatoi FELIPPE-BAUER (Culicoides) 73
lacustris RONDEROS (Culicoides) 71 lobatus BORKENT & PICADO (Atrichopogon) 46
lahillei (ICHES) (Culicoides) 17, 29, 33, 67 longicornis (WILLISTON) (Palpomyia) 97
longicornis EWEN (Atrichopogon) 25, 46 manaosensis LANE & FORATTINI (Stilobezzia) 85

longispina SAUNDERS (Forcipomyia) 25, 49 mapuche MARINO & SPINELLI (Forcipomyia) 53
lopesi BARRETTO (Culicoides) 73 mapuche SPINELLI (Brachypogon) 79
Lophomyidium CORDERO 47 marginalis WIRTH & RATANAWORABHAN (Stenoxenus) 99
lota (WILLISTON) (Forcipomyia) 49 marini SPINELLI & DIPPOLITO (Forcipomyia) 50
loughnani EDWARDS (Culicoides) 65 marinkellei WIRTH & LEE (Culicoides) 64
louriei (MACFIE) (Forcipomyia) 59 marmosetae (Dipetalonema) 33
lutea (MALLOCH) (Stilobezzia) 112, 119 marshi WIRTH & BLANTON (Culicoides) 63
lutealaris WIRTH & BLANTON (Culicoides) 63 martinezi WIRTH & BLANTON (Culicoides) 74
luteigenua WIRTH & SPINELLI (Forcipomyia) 26, 53 maruim LUTZ (Culicoides) 17, 29, 69
luteisquamosa WIRTH (Forcipomyia) 54 Mataemyia VARGAS 40, 69
luteogrisea WIRTH & WILLIAMS (Dasyhelea) 60 mathisi SPINELLI & WIRTH (Bezzia) 94
luteovenus ROOT & HOFFMAN (Culicoides) 29, 64 maya FELIPPE-BAUER (Monohelea) 83
lutzi COSTA LIMA (Culicoides) 68 maya SPINELLI & WIRTH (Paryphoconus) 98
lutzi LANE (Palpomyia) 96 maya SPINELLI & WIRTH (Phaenobezzia) 97
lydiae BORKENT (Baeodasymyia) 77 mayeri FORATTINI & LANE (Forcipomyia) 54
lyrinotatus WIRTH & BLANTON (Culicoides) 74 mayeri ORTZ (Monohelea) 83
Macfiehelea LANE 89 mayeri WIRTH (Paryphoconus) 31, 98
macfiei (COSTA LIMA) (Forcipomyia) 56 mazaruni MACFIE (Bezzia) 94
macfiei LANE (Echinohelea) 81 mcateei WIRTH (Forcipomyia) 132
macfiei LANE (Paryphoconus) 98 mckeeveri BRICKLE & HAGAN (Culicoides) 65
macfiei LANE (Stilobezzia) 87 mediomunda MINAYA (Dasyhelea) 27, 60
macfiei SPINELLI & GROGAN (Paradasyhelea) 75 Megaconops WIRTH & ATCHLEY 44
Macfiella FOX 40, 69 megatheca SPINELLI & WIRTH (Bezzia) 94
macheti CLASTRIER & LEGRAND (Forcipomyia) 58 melleus (COQUILLETT) (Culicoides) 29, 75
macieli TAVARES & RUIZ (Culicoides) 29, 70 mellichroa MACFIE (Palpomyia) 96
macrostigma WIRTH & BLANTON (Culicoides) 74 mendozae INGRAM & MACFIE (Atrichopogon) 46
Macrurohelea INGRAM & MACFIE 40, 82, 101, 116, menzeli CLASTRIER & WIRTH (Forcipomyia) 54
130, 141, 151 merceri CAZORLA & SPINELLI (Stilobezzia) 87
maculata LANE (Stilobezzia) 87 mesotibialis SPINELLI & WIRTH (Bezzia) 94
maculata MACFIE (Dasyhelea) 60 messersmithi WIRTH & DOW (Forcipomyia) 56-57
maculicrus INGRAM & MACFIE (Palpomyia) 96 Metaforcipomyia SAUNDERS 26, 53
maculipennis (COQUILLETT) (Monohelea) 83 metagonatus WIRTH & BLANTON (Culicoides) 63
maculipennis CLASTRIER (Atrichopogon) 46 mexicanus HUERTA (Atrichopogon) 46
maculithorax (WILLISTON) (Culicoides) 70 michaeli BORKENT (Baeodasymyia) 29, 77
maculitibia LANE & FORATTINI (Stilobezzia) 86 microchela KIEFFER (Palpomyia) 96
maculosa INGRAM & MACFIE (Forcipomyia) 53 Microconops KIEFFER 43
magna (SAUNDERS) (Forcipomyia) 26, 56 Microhelea KIEFFER 4, 18, 19-21, 53
magnipalpis WIRTH & BLANTON (Culicoides) 73 micronyx WIRTH & BLANTON (Leptohelea) 82, 109,
magnus BORKENT & PICADO (Atrichopogon) 46, 95 112, 119, 126, 151
maia LANE & FORATTINI (Stilobezzia) 85 microtoma (KIEFFER) (Forcipomyia) 54
malariologiensis PERRUOLO (Culicoides) 75 minasensis FELIPPE-BAUER (Culicoides) 65
Mallochohelea WIRTH 32, 91, 102, 122, 128-129, minisquamosa WIRTH (Forcipomyia) 54
143, 145-146, 151 minitheca MARINO & SPINELLI (Forcipomyia) 50
minor (WIRTH) (Nilobezzia) 130 nelsoni GROGAN & WIRTH (Amerohelea) 92

mirsae ORTZ (Culicoides) 65 nemoralis (MACFIE) (Mallochohelea) 91
misionensis SPINELLI (Brachypogon) 79 Neobezzia WIRTH & RATANAWORABHAN 31, 41, 91,
misionensis SPINELLI (Paryphoconus) 98 102, 120, 122, 128, 143, 145, 151
miyamotoi WIRTH & BLANTON (Culicoides) 74 Neoceratopogon MALLOCH 76
modesta CLASTRIER & RACCURT (Baeodasymyia) 77, 108 neoparaensis TAVARES & SOUZA (Culicoides) 67
modesta LANE (Stilobezzia) 87 neopulicaris WIRTH (Culicoides) 29, 64
mojingaensis WIRTH & BLANTON (Culicoides) 69-70 neotropica (MACFIE) (Nilobezzia) 92
monicae SPINELLI & BORKENT (Culicoides) 63 neotropica CLASTRIER & WIRTH (Forcipomyia) 55
monicae SPINELLI & GROGAN (Brachypogon) 79 neotropica WIRTH (Allohelea) 76, 118, 148
monilicornis (COQUILLETT) (Forcipomyia) 57, 106 neotropica WIRTH (Echinohelea) 81
monilicornis WIRTH & BLANTON (Rhynchohelea) 84- neotropicalis (LANE) (Paryphoconus) 98
85, 104, 106, 112, 152 nerudai SPINELLI & GROGAN (Borkenthelea) 78
Monohelea KIEFFER 32, 46, 77, 80-82, 101, 110, neunguis GROGAN & WIRTH (Parabezzia) 84
127, 138, 151 niger LANE (Stenoxenus) 99
monotheca SPINELLI & GROGAN (Macrurohelea) 82 nigerrima INGRAM & MACFIE (Stilobezzia) 85
monticola INGRAM & MACFIE (Dasyhelea) 60 nigra KIEFFER (Heteromyia) 89
monticola WIRTH & LEE (Culicoides) 73 nigrescens MACFIE (Forcipomyia) 48
moorei CLASTRIER & WIRTH (Forcipomyia) 55 nigrigenus WIRTH & BLANTON (Culicoides) 62-63
morenoi MARINO & SPINELLI (Forcipomyia) 53 nigrimaxillata CLASTRIER & WIRTH (Forcipomyia) 55
morrisoni GROGAN & WIENERS (Dasyhelea) 60 nigripes (MACFIE) (Mallochohelea) 91
mortuifolii SAUNDERS (Forcipomyia) 25, 49 nigripes MACFIE (Clinohelea) 30, 88
multidentata RONDEROS & SPINELLI (Forcipomyia) 52 nigripes MACFIE (Paryphoconus) 98
multilineata (LUTZ) (Downeshelea) 80 nigritibialis SPINELLI & WIRTH (Bezzia) 94
multipicta INGRAM & MACFIE (Forcipomyia) 50 nigroflava LANE & FORATTINI (Stilobezzia) 87
multispinosa SPINELLI & GROGAN (Palpomyia) 96 nigroflava LANE (Palpomyia) 96
musae CLASTRIER & DELCOLLE (Forcipomyia) 26, 56 nigroscutellata LANE (Palpomyia) 96
mutabilis (COQUILLETT) (Dasyhelea) 27, 59-60 Nilobezzia KIEFFER 32, 91, 102, 122, 128, 143,
muzoni MARINO & SPINELLI (Forcipomyia) 50 145, 151
muzoni SPINELLI & DURET (Clinohelea) 88 nobilis (WINNERTZ) (Bezzia) 31, 93-94, 114, 124, 130
Mycterotypus NO 43 nodosa SAUNDERS (Forcipomyia) 26, 58
nana (MACFIE) (Forcipomyia) 58 nothofagus SPINELLI & GROGAN (Borkenthelea) 78, 112
nana WIRTH & BLANTON (Baeohelea) 77-78, 108- Notiohelea GROGAN & WIRTH 40, 83, 101, 103,
109, 112, 149 117, 134, 142, 151
Nannohelea GROGAN & WIRTH 83, 101, 107, 109, novaeteutoniae CLASTRIER & WIRTH (Forcipomyia) 55
138, 151 novaeteutoniae MACFIE (Atrichopogon) 46
nanus MACFIE (Atrichopogon) 46 nubeculosa SPINELLI & WIRTH (Alluaudomyia) 76
narthekophora MACFIE (Forcipomyia) 59 nubeculosus MACFIE (Atrichopogon) 46
navaiae BYSTRAK & WIRTH (Forcipomyia) 49 nubifer MACFIE (Paryphoconus) 98
nebulosus MACFIE (Atrichopogon) 46 nuncupata MACFIE (Forcipomyia) 50
necrophila SPINELLI & RODRGUEZ (Dasyhelea) 24, obesa COSTA LIMA (Forcipomyia) 55
27, 60 obfuscatus INGRAM & MACFIE (Atrichopogon) 46
needhami THOMSEN (Alluaudomyia) 132 obnoxius FOX (Culicoides) 74
neivai LANE (Clinohelea) 88 obnubilus INGRAM & MACFIE (Atrichopogon) 46
obscura LANE & FORATTINI (Stilobezzia) 87 panamensis BARBOSA (Culicoides) 65

obscurus EWEN (Atrichopogon) 25, 46 panamensis LANE & FORATTINI (Stilobezzia) 30, 87
ocumare (ORTZ) (Atrichopogon) 46 panamensis RONDEROS & SPINELLI (Leptoconops) 44
ocumarensis ORTZ (Culicoides) 69 panamensis WIRTH (Echinohelea) 81
Oecacta POEY 70 panamensis WIRTH (Parabezzia) 84
oedidactyla (INGRAM & MACFIE) (Physohelea) 90, 113 panamericanus FOX (Culicoides) 71
oklahomensis KHALAF (Culicoides) 29, 62 pancensis BROWNE (Culicoides) 75
oligarthra SAUNDERS (Forcipomyia) 26, 56 Parabezzia MALLOCH 32, 83, 101, 104, 109, 110,
oliveirai (LANE) (Pellucidomyia) 89 115-116, 139-141, 150, 152-153
oliveirai LANE (Palpomyia) 96 paracaudata GROGAN & WIRTH (Macrurohelea) 82
oliveirai LANE (Paryphoconus) 31, 98 paracincta WIRTH (Dasyhelea) 27, 60
oliveri FOX & HOFFMAN (Culicoides) 69 Paradasyhelea MACFIE 32, 40, 75, 101, 105, 137, 152
opilionivora (LANE) (Forcipomyia) 58 paraensis (GOELDI) (Culicoides) 17, 24, 29, 33, 66-67
orellana (ROBACK) (Heteromyia) 89 paraensis (LANE) (Nilobezzia) 92
orjuelai WIRTH & LEE (Culicoides) 63 paraensis LANE (Palpomyia) 96
ornata LANE & FORATTINI (Stilobezzia) 85 paraensis LANE (Stenoxenus) 99
ornata WIRTH (Monohelea) 83 paraensis WIRTH & BLANTON (Brachypogon) 79, 112
ornaticrus INGRAM & MACFIE (Stilobezzia) 85 parahybae MACFIE (Dasyhelea) 60
ornatipennis CLASTRIER (Atrichopogon) 46 paraignacioi SPINELLI (Culicoides) 69
ornatipennis MACFIE (Echinohelea) 81 paramaruim WIRTH & BLANTON (Culicoides) 69
ornatipennis MACFIE (Forcipomyia) 51 paranaensis SPINELLI & WIRTH (Paryphoconus) 98
ortizi FOX (Culicoides) 71 parascopus WIRTH & BLANTON (Culicoides) 65
ozzardi (Dipetalonema) 33 Parastilobezzia WIRTH & BLANTON 40-41, 84, 101,
pabloi BROWNE (Culicoides) 72 110, 139
pachycnemus MACFIE (Atrichopogon) 46 Paryphoconus ENDERLEIN 32, 41, 97, 102, 123, 128-
Pachyhelea WIRTH 32, 41, 95, 102, 123, 128, 144- 129, 144-146, 152
145, 152 patagonica INGRAM & MACFIE (Dasyhelea) 60
Pachyleptus WALKER 89 patagonica INGRAM & MACFIE (Forcipomyia) 50
pachymera (WILLISTON) (Pachyhelea) 31, 95, 124, 152 patagonica INGRAM & MACFIE (Palpomyia) 96
pachymerus LUTZ (Culicoides) 74 patagonica INGRAM & MACFIE (Stilobezzia) 85
Padrosia RAFINESQUE 61 patagonica SPINELLI & GROGAN (Diaphanobezzia) 80
painteri FOX (Culicoides) 68 patagoniensis RONDEROS (Leptoconops) 43
pallescens LANE & FORATTINI (Stilobezzia) 87 patagoniensis RONDEROS & SPINELLI (Culicoides) 17, 64
pallida SPINELLI & GROGAN (Parabezzia) 84 patulipalpis WIRTH & BLANTON (Culicoides) 72
pallidipennis SPINELLI & GROGAN (Brachypogon) 79 paucienfuscatus BARBOSA (Culicoides) 74
pallidipes KIEFFER (Atrichopogon) 46 paulistana FORATTINI & RABELLO (Dasyhelea) 27, 61
palpalis MACFIE (Atrichopogon) 46 paulistensis LANE (Palpomyia) 96
palpalis MACFIE (Culicoides) 69 paulistensis LANE (Paryphoconus) 98
Palpomyia MEIGEN 23, 30, 32, 79-80, 86, 89, 90, paulistensis LANE (Stilobezzia) 87
92, 95, 102, 122-123, 128, 143-145, 152 pectinatus MACFIE (Atrichopogon) 46
pampaensis SPINELLI & WIRTH (Culicoides) 67 Pedilohelea DE MEILLON & WIRTH 48, 55
pampana LANE (Palpomyia) 96 pellucida INGRAM & MACFIE (Diaphanobezzia) 80
pampoikilus MACFIE (Culicoides) 71 Pellucidomyia MACFIE 32, 89, 101, 117, 142, 152
panamensis (LANE & WIRTH) (Downeshelea) 80 penicillatus DELCOLLE & RIEB (Atrichopogon) 46
penthesileae MACFIE (Dasyhelea) 27, 60 propinqua (WILLISTON) (Forcipomyia) 50

penultimata WIRTH (Forcipomyia) 55 propinquus MACFIE (Culicoides) 75
perstans (Dipetalonema ) 33 propriipennis MACFIE (Culicoides) 72
peruviana CLASTRIER & WIRTH (Forcipomyia) 55 Protersesthes KIEFFER 43
peruvianus FELIPPE-BAUER (Culicoides) 67 pseudocrescentis TAVARES & LUNA DIAS (Culicoides) 72
peruvianus KIEFFER (Atrichopogon) 46 Pseudoculicoides MALLOCH 59
petiolata MALLOCH (Parabezzia) 83, 113 pseudodiabolicus FOX (Culicoides) 17, 69
petrocchiae SHANNON & DEL PONTE (Leptoconops) pseudofasciata GROGAN & WIRTH (Amerohelea) 92
44, 151 pseudofulvus SPINELLI (Stenoxenus) 99
Phaenobezzia HAESELBARTH 32, 97, 102, 120, 122, pseudogibbera SPINELLI & WIRTH (Bezzia) 94
128, 143, 145, 152 pseudoincisurata WAUGH and WIRTH (Dasyhelea) 106,
phaeonotus WIRTH & BLANTON (Culicoides) 71 111, 118, 125
Phasmidohelea MAYER 53-55 pseudolacustris DIPPOLITO & SPINELLI (Palpomyia) 96
philotherma MACFIE (Dasyhelea) 61 pseudoparaensis SPINELLI & GROGAN (Brachypogon) 79
phlebotomus (WILLISTON) (Culicoides) 17-18, 29, 33, 69 pseudovenustula SPINELLI & WIRTH (Bezzia) 94
Physohelea GROGAN & WIRTH 40, 90, 102, 117, pseudunguis SPINELLI & GROGAN (Parabezzia) 84
142, 152 Psychophaena PHILIPPI 40, 70-71
Phytohelea REMM 26, 55 Pterobosca MACFIE 20, 56
picadoae SPINELLI & BORKENT (Culicoides) 72 pulchripennis MACFIE (Culicoides) 73
piceiventris KIEFFER (Atrichopogon) 46 pulchripennis MACFIE (Sphaeromias) 76
pichindensis BROWNE (Culicoides) 73 pulchripes KIEFFER (Bezzia) 31, 94
picta (COQUILLETT) (Stilobezzia) 87 pulicaris (LINNAEUS) (Culicoides) 132
pictipennis (PHILIPPI) (Culicoides) 70-71 pullata (WIRTH) (Mallochohelea) 91
pictipennis CLASTRIER (Atrichopogon) 46 pumila MACFIE (Dasyhelea) 61
pictoni MACFIE (Forcipomyia) 26, 51 punctipennis (WILLISTON) (Bezzia) 93-94
pifanoi ORTZ (Culicoides) 72, 74 punctipes WIRTH (Stilobezzia) 87
pilosa SPINELLI & GROGAN (Notiohelea) 83 punctiradialis SPINELLI & WIRTH (Alluaudomyia) 77
pilosus WIRTH & BLANTON (Culicoides) 65 punctivenosa WIRTH & GROGAN (Alluaudomyia) 77
pinamarensis SPINELLI (Forcipomyia) 51 punctulata LANE (Stilobezzia) 87
piroskyi CAVALIERI (Forcipomyia) 50 puracensis WIRTH & DOW (Forcipomyia) 57
pistiae (INGRAM & MACFIE) (Phaenobezzia) 97, 114, 130 puracensis WIRTH & LEE (Culicoides) 17, 63
platensis (BRTHES) (Forcipomyia) 49 pusilla (LUTZ) (Dasyhelea) 27, 61
plaumanni CLASTRIER & WIRTH (Forcipomyia) 55 pusilloides WIRTH & BLANTON (Culicoides) 63
plaumanni SPINELLI & WIRTH (Alluaudomyia) 76 pusillus LUTZ (Culicoides) 22, 33, 63
plaumanni SPINELLI (Culicoides) 69 pygmaea (WILLISTON) (Dasyhelea) 61
plebeja (LOEW) (Palpomyia) 114, 124 pyrsonota MACFIE (Dasyhelea) 61
poikilonotus MACFIE (Culicoides) 65 quartibrunneus BORKENT & PICADO (Atrichopogon) 46
pollex BORKENT & FORSTER (Dasyhelea) 61 quasicornuta SAUNDERS (Forcipomyia) 26, 52
polypori WIRTH & BLANTON (Culicoides) 69 quasidentica FELIPPE-BAUER & QUINTELAS
polystictus KIEFFER (Culicoides) 75 (Downeshelea) 81
poncai LANE & WIRTH (Monohelea) 83 quasiingrami MACFIE (Forcipomyia) 49
popayanensis WIRTH & LEE (Culicoides) 63 quasiparaensis CLASTRIER (Culicoides) 67
poulaineae INGRAM & MACFIE (Forcipomyia) 50-51 quatei SPINELLI & GROGAN (Borkenthelea) 78
prima CLASTRIER (Alluaudomyia) 76 quatei WIRTH (Forcipomyia) 50
quechua MARINO & SPINELLI (Forcipomyia) 151 saintjusti TAVARES & RUIZ (Culicoides) 68
rabelloi LANE (Stilobezzia) 30, 87 Saliohelea WIRTH & RATANAWORABHAN 48, 57
raccurti SPINELLI & GROGAN (Parabezzia) 84 saltaensis SPINELLI & WIRTH (Culicoides) 65
rachoui TAVARES & SOUZA (Culicoides) 67 saltanensis LANE & DURET (Clinohelea) 88
raleighi MACFIE (Forcipomyia) 25, 50 saltensis (CAVALIERI) (Forcipomyia) 52
rangeli ORTZ & MIRSA (Culicoides) 72 sanctaeclarae MACFIE (Atrichopogon) 47
raposoensis CLASTRIER & WIRTH (Forcipomyia) 55 sanctaeclarae WIRTH (Forcipomyia) 57
raposoensis SPINELLI & WIRTH (Bezzia) 94 sanctibernardini KIEFFER (Stilobezzia) 86
raposoensis WIRTH & BARRETO (Culicoides) 64 sanctilaurentii KIEFFER (Atrichopogon) 47
raposoi DE MEILLON & WIRTH (Forcipomyia) 55 sanmartini WIRTH & BARRETO (Culicoides) 67
rava INGRAM & MACFIE (Stilobezzia) 86 santaemarthae (KIEFFER) (Dasyhelea) 61
recifei BARBOSA (Culicoides) 69 santanderi BROWNE (Culicoides) 73
recifensis BARBOSA (Culicoides) 74 saundersi EWEN (Atrichopogon) 25, 47, 132
reconditus CAMPBELL & PELHAM-CLINTON sayhuequei MARINO & SPINELLI (Forcipomyia) 58
(Culicoides) 135 scabra (COQUILLETT) (Palpomyia) 96
redactus BORKENT & PICADO (Atrichopogon) 46 Schizoforcipomyia CHAN & LEROUX 57
remigatus EWEN (Atrichopogon) 25, 46 Schizonyxhelea CLASTRIER 30, 32, 41, 85, 101, 107,
remigera UTMAR & WIRTH (Forcipomyia) 48 138, 152
reticulatus LUTZ (Culicoides) 29, 74 schmitzi SPINELLI & GROGAN (Brachypogon) 79
rettenmeyerorum CLASTRIER & WIRTH (Forcipomyia) 55 schmitzorum DIPPOLITO & SPINELLI (Bezzia) 94
reynoldsi INGRAM & MACFIE (Dasyhelea) 61 schnacki SPINELLI & GROGAN (Clastrieromyia) 95, 114
Rhynchoforcipomyia WIRTH & DOW 40, 48, 56 schnacki SPINELLI (Alluaudomyia) 29, 77
Rhynchohelea WIRTH & BLANTON 84, 101, 103, 107, schwarzii (COQUILLETT) (Nilobezzia) 30, 92
134, 137-138, 152 scissurae MACFIE (Dasyhelea) 61
ricardoi RONDEROS & SPINELLI (Leptoconops) 44 scopus ROOT & HOFFMAN (Culicoides) 65
richardsi MACFIE (Echinohelea) 81 scorzai ORTZ (Culicoides) 75
ringueleti SPINELLI (Brachypogon) 79 scutata LANE & FORATTINI (Stilobezzia) 87
riojana SPINELLI & MARINO (Forcipomyia) 58 seminole WIRTH (Forcipomyia) 26, 52-53
rioplatensis MARINO & SPINELLI (Forcipomyia) 26, 51 sentior (INGRAM & MACFIE) (Austrosphaeromias) 90
rodriguezi ORTZ (Culicoides) 72 sequax (WILLISTON) (Atrichopogon) 47
roldani SPINELLI & WIRTH (Bezzia) 31, 94 setifer (LUTZ) (Culicoides) 64
ronderosae SPINELLI & BORKENT (Culicoides) 65 setiger MACFIE (Stenoxenus) 99
ronderosi GROGAN & WIRTH (Amerohelea) 92 setigera SAUNDERS (Forcipomyia) 25, 49
rondoniensis SPINELLI & DIPPOLITO (Groganhelea) setigera SPINELLI & WIRTH (Bezzia) 94
90, 114, 120, 150 setosa WIRTH (Macrurohelea) 82
roraimensis FELIPPE-BAUER & SPINELLI (Monohelea) 83 setosicrus (KIEFFER) (Forcipomyia) 49
rostratus WIRTH & BLANTON (Culicoides) 25, 63 setosilateralis BORKENT & PICADO (Atrichopogon) 47
roubaudi CLASTRIER & DELCOLLE (Forcipomyia) 58 seudoobfuscatus SPINELLI (Atrichopogon) 47
rozeboomi BARBOSA (Culicoides) 68-69 sexpunctata SPINELLI & WIRTH (Alluaudomyia) 77
rubriceps KIEFFER (Clinohelea) 88 sexvittata WIRTH (Forcipomyia) 51
rufa KIEFFER (Heteromyia) 89 shannoni (INGRAM & MACFIE) (Forcipomyia) 58
ruizi FORATTINI (Culicoides) 69 shannoni (WIRTH & BLANTON) (Austrohelea) 77,
rusticus MACFIE (Atrichopogon) 47 113, 148
sabalitensis UTMAR & WIRTH (Forcipomyia) 48 shannoni INGRAM & MACFIE (Dasyhelea) 61
silvicola MACFIE (Stilobezzia) 87 Stilobezzia KIEFFER 17, 24, 32, 40, 84-86, 101-
similans LANE & FORATTINI (Stilobezzia) 87 103, 115-117, 126, 134, 140-142, 149, 152
similis SPINELLI & GROGAN (Macrurohelea) 82 stonei (WIRTH) (Downeshelea) 80-81
similis SPINELLI & MARINO (Atrichopogon) 47 stubalensis FOX (Culicoides) 74
similis SPINELLI (Amerohelea) 92 stylifer (LUTZ) (Forcipomyia) 26, 51-52
simplex LANE & FORATTINI (Stilobezzia) 87 suarezi RODRGUEZ & WIRTH (Culicoides) 63
sinclairi BORKENT (Dasyhelea) 61 suarezi SPINELLI & RONDEROS (Dasyhelea) 61
sinuosa DOW & WIRTH (Forcipomyia) 58 subaspera (COQUILLETT) (Palpomyia) 31, 96
smarti MACFIE (Echinohelea) 81 subflavus MACFIE (Paryphoconus) 98
smithi (LEWIS) (Mallochohelea) 114 subfusca MACFIE (Bezzia) 94
snowi LANE (Bezzia) 31, 94 subfuscula INGRAM & MACFIE (Palpomyia) 97
soibelzoni MARINO & SPINELLI (Forcipomyia) 57 subsessilis KIEFFER (Stilobezzia) 86
somuncurensis MARINO & SPINELLI (Forcipomyia) 51 succinea INGRAM & MACFIE (Stilobezzia) 86
sonorensis WIRTH & RATANAWORABHAN Synthyridomyia SAUNDERS 57
(Paryphoconus) 98 taeniatus MACFIE (Atrichopogon) 47
sordidipes (MACFIE) (Amerohelea) 30, 92 talarum SPINELLI (Atrichopogon) 47
soriai WIRTH & WAUGH (Dasyhelea) 61 tamboensis WIRTH & LEE (Culicoides) 72
soriai WIRTH (Forcipomyia) 51 tamioi LANE (Palpomyia) 97
spangleri DE MEILLON & WIRTH (Forcipomyia) 55 tanycornis BORKENT (Palpomyia) 97
spangleri WIRTH (Parabezzia) 84 tapantiensis BORKENT & PICADO (Atrichopogon) 47
spathicerca WIRTH (Dasyhelea) 61 taragui MARINO (Forcipomyia) 50
spatulifera SAUNDERS (Forcipomyia) 25, 49 taragui SPINELLI (Paryphoconus) 98
spatuliformis SPINELLI & GROGAN (Brachypogon) 79 tarapaca SPINELLI & WIRTH (Culicoides) 65
spatuligera MACFIE (Forcipomyia) 51 tavaresi FELIPPE-BAUER & WIRTH (Culicoides) 74
Sphaerohelea SPINELLI & FELIPPE-BAUER 41, 92, tehuelche MARINO & SPINELLI (Forcipomyia) 58
102-103, 122-123, 134, 143-144, 152 telesfordi SPINELLI & GROGAN (Brachypogon) 79
spilmani DE MEILLON & WIRTH (Forcipomyia) 55 telmatophilus (MACFIE) (Paryphoconus) 98
spinellii GROGAN & WIRTH (Amerohelea) 82 tenuiannulata SPINELLI & WIRTH (Alluaudomyia) 77
spinellii WIRTH & GROGAN (Alluaudomyia) 77 tenuicrus KIEFFER (Palpomyia) 97
spinellii WIRTH & GROGAN (Diaphanobezzia) 80 tenuiforceps MACFIE (Forcipomyia) 57
spinifemur (LANE) (Palpomyia) 96 tenuilobus WIRTH & BLANTON (Culicoides) 73
spinifemur WIRTH & BLANTON (Lanehelea) 91 tenuisquamipes WIRTH (Forcipomyia) 51
spinosa LUTZ (Palpomyia) 96 teretipalpis WIRTH & BARRETO (Culicoides) 71
spinosa SAUNDERS (Forcipomyia) 26, 59 terminalis (COQUILLETT) (Paryphoconus) 98
spinosus BORKENT & PICADO (Atrichopogon) 47 termophila (SPINELLI) (Mallochohelea) 30, 91
spurius WIRTH & BLANTON (Culicoides) 67 terrestris SAUNDERS (Forcipomyia) 26, 59
squamithorax CLASTRIER (Forcipomyia) 53 Tersesthes TOWNSEND 43-44
squamitibia LUTZ (Forcipomyia) 49, 51 Tetraphora PHILIPPI 48, 50
squamosa LUTZ (Forcipomyia) 55 tetrathyris WIRTH & BLANTON (Culicoides) 73
steineri SPINELLI & WIRTH (Paryphoconus) 98 tettigonaris WIRTH & CASTNER (Forcipomyia) 55
Stenoxenus COQUILLETT 32, 99, 102, 122-123, 129, thalestris MACFIE (Dasyhelea) 61
143-144, 146, 152 thersites (WILLISTON) (Atrichopogon) 47
stigmalis (COQUILLETT) (Brachypogon) 111 thomasi CLASTRIER & WIRTH (Forcipomyia) 55
stigmalis WIRTH (Culicoides) 75 thomsenae WIRTH (Stilobezzia) 87
thoracica INGRAM & MACFIE (Macrurohelea) 82 uruguayensis SPINELLI & GROGAN (Clastrieromyia) 95
Thyridomyia SAUNDERS 57 utae KNAB (Forcipomyia) 51
tibialis LANE & FORATTINI (Stilobezzia) 86 utricularis MACFIE (Atrichopogon) 47
tidwelli SPINELLI (Culicoides) 69 valleensis CLASTRIER & WIRTH (Forcipomyia) 55
tirzae BORKENT & PICADO (Atrichopogon) 47 vargasi GROGAN & WIRTH (Amerohelea) 92
todatangae WIRTH & BLANTON (Culicoides) 67 vargasi WIRTH & BLANTON (Culicoides) 64
torrens (TOWNSEND) (Leptoconops) 43, 125 varia INGRAM & MACFIE (Stilobezzia) 86
townesi LANE (Clinohelea) 88 varicolor SAUNDERS (Forcipomyia) 25, 48
townsendi KNAB (Forcipomyia) 51 variipennis COQUILLETT (Culicoides) 5
townsendi LANE (Clinohelea) 88 varipennis WIRTH & WILLIAMS (Forcipomyia) 53
transferrans ORTZ (Culicoides) 72 venezuelensis ORTZ & MIRSA (Culicoides) 29, 70-71
transversa LANE & FORATTINI (Stilobezzia) 87 venezuelensis ORTZ (Leptoconops) 44
trapidoi WIRTH & BARRETO (Culicoides) 63 venezuelensis ORTZ (Stilobezzia) 88
travassosi FORATTINI (Culicoides) 69 venustula (WILLISTON) (Bezzia) 93-94
travassosi LANE (Paryphoconus) 98 verecundus MACFIE (Culicoides) 69
travassosi LANE (Stilobezzia) 88 vernoni CLASTRIER & WIRTH (Forcipomyia) 55
Trichohelea GOETGHEBUER 58 vernoni WIRTH & BLANTON (Culicoides) 73
tricoloratus WIRTH & BLANTON (Culicoides) 74 versicolor MACFIE (Palpomyia) 97
trifidus SPINELLI & BORKENT (Culicoides) 73 villosipes (KIEFFER) (Dasyhelea) 61
trilineatus FOX (Culicoides) 75 viridans FORATTINI & RABELLO (Dasyhelea) 61
trinidadensis HOFFMAN (Culicoides) 69 volcanensis WIRTH & BLANTON (Culicoides) 70
trinidadensis MACFIE (Atrichopogon) 47 wagneri (SGUY) (Forcipomyia) 55
trinidadensis SAUNDERS (Forcipomyia) 26, 58 wallacei WIRTH & BLANTON (Culicoides) 70
tripunctata SPINELLI & WIRTH (Alluaudomyia) 77 Warmkea SAUNDERS 58
tropica (KIEFFER) (Forcipomyia) 54 warmkei WIRTH (Atrichopogon) 47
tuberculata SAUNDERS (Forcipomyia) 27, 59 weemsi WIRTH & SPINELLI (Forcipomyia) 53
tuberculatus EWEN (Atrichopogon) 25, 47 williamsi CAZORLA & SPINELLI (Stilobezzia) 88
turgidipes (INGRAM & MACFIE) (Physohelea) 90 williamsi MARINO & SPINELLI (Forcipomyia) 53
ugandae MACFIE (Pellucidomyia) 119 williamsi SPINELLI (Culicoides) 73
umbella MACFIE (Palpomyia) 97 williamsi WIRTH & WAUGH (Dasyhelea) 61
umbratilis MACFIE (Atrichopogon) 47 willistoni WIRTH & BLANTON (Culicoides) 69
uncinata (JOHANNSEN) (Parabezzia) 84 willistoni WIRTH (Forcipomyia) 55
undecimpunctatus KIEFFER (Culicoides) 67 winderi WIRTH & WAUGH (Dasyhelea) 61
unetensis PERRUOLO (Culicoides) 75 winderi WIRTH (Forcipomyia) 53
unguis WIRTH (Parabezzia) 84 wirthi (LANE) (Pellucidomyia) 89
unica BYSTRAK & WIRTH (Forcipomyia) 49 wirthi CLASTRIER & RACCURT (Parabezzia) 84
unimaculatus MACFIE (Paryphoconus) 98 wirthi LANE & FORATTINI (Stilobezzia) 88
uniradialis WIRTH & BLANTON (Culicoides) 74 wirthi LANE (Palpomyia) 31, 97
univesicula MACFIE (Forcipomyia) 58 wirthi LANE (Paryphoconus) 99
uramaensis SPINELLI & DIPPOLITO (Forcipomyia) 50 wirthi SPINELLI & FELIPPE-BAUER (Neobezzia) 91
urracaisi LANE & WIRTH (Monohelea) 83 wirthi SPINELLI & GROGAN (Macrurohelea) 82
uruguayensis (CORDERO) (Atrichopogon) 47 wirthi SPINELLI (Austrosphaeromias) 90
uruguayensis FELIPPE-BAUER & SPINELLI (Monohelea) 83 wirthi SPINELLI (Brachypogon) 79
uruguayensis RONDEROS (Culicoides) 65 wirthomyia VARGAS (Culicoides) 65
wokei BARBOSA (Culicoides) 69 yamana SPINELLI & GROGAN (Macrurohelea) 82

wokei FOX (Culicoides) 69, 75 yaracuyensis ORTZ (Culicoides) 73
wokei WIRTH & GROGAN (Heteromyia) 30, 89 yolancae BORKENT & PICADO (Atrichopogon) 47
woodfordi MACFIE (Atrichopogon) 47 youngi SPINELLI & WIRTH (Alluaudomyia) 77
woodruffi SPINELLI & GROGAN (Brachypogon) 79 youngi WIRTH & BARRETO (Culicoides) 67
woodruffi SPINELLI & WIRTH (Bezzia) 94 youngi WIRTH & GROGAN (Cacaohelea) 79, 111, 119
wygodzinskyi CAVALIERI (Forcipomyia) 50 youngi WIRTH (Forcipomyia) 50
wygodzinskyi LANE (Clinohelea) 89 yukonensis HOFFMAN (Culicoides) 104
wygodzinskyi LANE (Paryphoconus) 99 zeteki WIRTH & DOW (Forcipomyia) 57
wygodzinskyi LANE (Stilobezzia) 30, 88 zonogaster INGRAM & MACFIE (Forcipomyia) 51
xanifer WIRTH & BLANTON (Culicoides) 69 zumbadoi SPINELLI & BORKENT (Culicoides) 72
yamana MARINO & SPINELLI (Forcipomyia) 51

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Aquatic Biodiversity in Latin America ABLA

Biodiversidad Acutica en Amrica Latina Vol. 4

FOR MANY YEARS,

Neotropical Ceratopogonidae (Diptera: Insecta)

Art Borkent & Gustavo R. Spinelli

Aquatic Biodiversity in Latin America

Biodiversidad Acutica en Amrica Latina

BIODIVERSIDAD ACUTICA EN AMRICA LATINA

Lector Cientfico a los Editores:

AQUATIC BIODIVERSITY IN LATIN AMERICA

Scientific Reader to the Editors:

AQUATIC BIODIVERSITY IN LATIN AMERICA

Series Editors/Editores de la Serie:

Aquatic Biodiversity in Latin America Vol. 4

The book should be cited as follows:

Printed in Bulgaria, June 2007

Habitus of female Culicoides variipennis COQUILLETT

AQUATIC BIODIVERSITY IN LATIN AMERICA

BIODIVERSIDAD ACUTICA EN AMRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias,

Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher.

Volume 2: Ephemeroptera of South America by Eduardo Domnguez, Carlos Molineri,

Volume 3: Neotropical Simuliidae (Diptera: Insecta) by Sixto Coscarn & Cecilia L.

Volume 4: Neotropical Ceratopogonidae (Diptera: Insecta) by Art Borkent & Gustavo

Neotropical Ceratopogonidae (Diptera: Insecta)

species in this region, methods of collection and rearing ceratopogonids, phylogenetic

The Ceratopogonidae are commonly known as biting midges, no-see-ums or punkies, in

We gratefully acknowledge permission to use photographs from several colleagues

1 B References to species of Neotropical Ceratopogonidae

described as eggs, larvae and/or pupae (Table 1) 25

2C APTURE AND STUDY OF CERATOPOGONIDAE 34

4P HYLOGENETIC RELATIONSHIPS AND FOSSIL

6 C Catalog of the Ceratopogonidae of the Neotropical

Neotropical Ceratopogonidae (Table 5) 101

Key to the genera of adult Ceratopogonidae in the

10 S UBJECT INDEX 182

Table 1. References to species of Neotropical Ceratopogonidae described as eggs, larvae

Taxon Reference Stages Described

Atrichopogon bifidus EWEN & SAUNDERS, 1958 L P

Forcipomyia (Caloforcipomyia) caerulea SAUNDERS, 1957 L P

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

F. (Forcipomyia) argenteola LANE, 1947c L P

Taxon Reference Stages Described

F. (Lasiohelea) attenuata SAUNDERS, 1964 L P

F. (Lepidohelea) brasiliensis WIRTH & SPINELLI, 1993a P

F. (Metaforcipomyia) cerifera SAUNDERS, 1957 L P

F. (Phytohelea) antiguensis SAUNDERS, 1957 L P

F. (Thyridomyia) nodosa SAUNDERS, 1957 L P

F. (Trichohelea) trinidadensis SAUNDERS, 1964 P

F. (Warmkea) aeria SAUNDERS, 1957 L P

Taxon Reference Stages Described

Dasyhelea atlantis WAUGH & WIRTH, 1976 P

Culicoides annettae SPINELLI & BORKENT, 2004a P

Taxon Reference Stages Described

Taxon Reference Stages Described

Alluaudomyia bella (as splendida) THOMSEN, 1937 L P

Baeodasymyia christopheri BORKENT & CRAIG, 1999 E L P

Echinohelea lanei WIRTH, 1994b P

Taxon Reference Stages Described

Schizonyxhelea forattinii BORKENT, 2000b L P

Stilobezzia antennalis THOMSEN, 1937 L P

Clinohelea horacioi SPINELLI & DURET, 1993 P

Heteromyia clavata WIRTH & GROGAN, 1977 P