Chapter 16.
Family Corethrellidae
André Pereira Amaral
Departamento de Ecologia e Zoologia, Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina, Brazil
Heron Huerta
Instituto de Diagnóstico y Referencia Epidemiológicos, Laboratorio de Entomologı´a, Ciudad de Me´xico, Me´xico
Luiz Carlos Pinho
Departamento de Ecologia e Zoologia, Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina, Brazil
Chapter Outline
Introduction
Limitations
Terminology and Morphology
INTRODUCTION
The family Corethrellidae is considered the sister taxon of
Culicidae (mosquitoes) þ Chaoboridae (phantom-midges),
which together form the superfamily Culicoidea. Some
synapomorphies supporting this group include larval
thoracic segments fused in a single distinct region and
larval spiracle elevated on a siphon structure. The early
development of adult eye in larvae and movable paddles in
pupae are characters not shared by Corethrellidae that
support the relationships between the superfamily members
(Wood & Borkent, 1989).
Corethrellidae is a monobasic family with only the
genus Corethrella Coquillet. Its members are known as
frog-biting midges, due to its female’s habit of feeding on
anuran blood after first locating the male hosts by eaves-
dropping on their mating calls. The group distribution
ranges from 50 N to 50 S of latitude, although most of the
species are found between 30 N and 30 S (Robert &
Borkent, 2014). The distribution and diversity of habitats
occupied by the aquatic stages of Corethrella is coincident
to those of their anuran hosts, such as ponds, phytotelmata,
swamps, and the lentic parts of streams (Borkent, 2008). A
total of 108 extant and nine fossil species have been
recorded, but the actual diversity of the family is expected
to be much higher, as only a small part of its distribution
has been sampled (Borkent, 2008; Yu et al., 2013; Amaral
& Pinho, 2015; Baranov et al., 2016; Caldart et al., 2016).
Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00036-6
Copyright Ó 2018 Elsevier Inc. All rights reserved.
751 Material Preparation and Preservation 753
752 References 753
752
Larvae of Corethrella are ambush predators and feed on
a variety of aquatic invertebrates, including Culicidae and
Chironomidae larvae, microcrustaceans, rotifers, and
nematodes (Borkent, 2008). They are thought to alter the
structure of communities, influencing the abundance and
development of prey species (Alto et al., 2005; Griswold &
Lounibos, 2005). Most of the time, they are found sub-
merged, holding on the substrate by their siphon, but can
also be found at the surface, lying horizontally on the
meniscus (it is not well known how this behavior varies
between species). Pupae have almost no motility and
generally stay at the surface, although some may prefer
hanging vertically or at about 45 angle (Borkent, 2008).
According to Borkent (2008), the overall length of devel-
opment stages, based on specimens reared in laboratory is
2e6 days for egg hatching, 11e37 days at larval stage and
3e8 days as pupa.
There are currently 73 valid species from the
Neotropical region, 19 of which were described by Lane
FAMILY CORETHRELLIDAE
(1953) and Lane & Aitken (1956), who had been the
main contributors to the taxonomy of the family in
Neotropics until Borkent’s publication in 2008. The latter
summarized essentially all the knowledge about the
taxonomy, phylogeny, biology, zoogeography, and
evolution of the family at that time, while adding a vast
amount of new information and analyses. That work
encouraged novel research in the ecology and systematics
751
 752 Thorp and Covich’s Freshwater Invertebrates

of Corethrellidae (Silva & Bernal, 2013; Amaral &
Pinho, 2015; Baranov et al., 2016; Caldart et al., 2016);
although, according to the author, there is still a lot to
discover.
LIMITATIONS
The only available keys to immature stages of Corethrella
were provided by McKeever & French (1991), comprising
only larvae of four Nearctic species, and Amaral & Pinho
(2015), for pupae of the Neotropical peruviana species
group. Of the 73 valid extant Neotropical species, 28 were
redescribed and 42 new ones were first described by
Borkent (2008); however, those descriptions were focused
FAMILY CORETHRELLIDAE
mainly on morphology of adult females and some males,
with a few incomplete descriptions of larvae and pupae that
were not sufficient to create a comprehensive taxonomic
key for the group. Descriptions from previous authors also
do not give enough attention to immatures, resulting in 18
species that cannot be identified. Hence, a key will not be
provided here, because the small number of taxa that would
be included, compared to the diversity in the region, would
probably lead to misidentifications.
TERMINOLOGY AND MORPHOLOGY
Larva (Figs. 16.7.1 A, B) shows unique posterolateral rows
of spines on each side of head capsule (Borkent, 2008). The

FIGURE 16.7.1 (A) Corethrella sp. larval habitus, dorsal view; (B) Corethrella borkenti Amaral & Pinho head capsule, ventral view. Dorsal view of
pupal exuviae: (C) Corethrella alticola Lane; (D) Corethrella infuscata Lane; and (E) C. borkenti.
Chapter | 16.7 Family Corethrellidae
shape of larval head capsule ranges from somewhat round
(Fig. 16.4. H) to anteriorly elongate (Fig. 16.7.1 A), with an
anterolateral groove in which the antennae, apically inser-
ted, is kept. Mandibles and mentum are well developed,
with bold teeth. The thorax is large, with three fused seg-
ments and first instar have a ventral row of hooks. The
abdomen is elongate, with 10 segments and a short but well
developed siphon on eighth segment. Chaetotaxy is
described in details in Belkin (1962).
The pupa body (Figs. 16.7.1 C, E) is oriented in one
plane and has a broad cephalothorax, compared to the
narrower and posteriorly tapered abdomen. Total length of
pupa is about 2.5 mm. Cephalothorax bears dorsolateral
respiratory organs, which may be funicular (Figs. 16.7.1 D,
E) or expanded and flattened (Fig. 16.7.1 C). Most species
possess campaniform sensilla on the metathorax (except
those from peruviana species group), which are large
spherical pits derived from a modified sensillum
(Figs. 16.7.1 D, E). The abdomen is dorsoventrally com-
pressed and slightly to strongly tapering distally, ending in
a pair of tapering terminal processes, which may hold either
a fused spine (Fig. 16.7.1 C) or a socketed seta (Fig. 16.7.1
E). The sensilla are variable and some species may have
conspicuously long lateral and/or mesal setae, as well as
other shorter ones distributed on dorsum and ventrum.
Some species have multiple lateral, marginal setae on the
abdomen (Borkent, 2012).
MATERIAL PREPARATION AND
PRESERVATION
Preparation and preservation techniques are detailed in
Borkent & Spinelli (2007) and Belkin (1962). Immatures
and adults can be preserved temporarily in 70%e80%
ethanol inside a freezer to avoid bleaching, and should be
slide-mounted or pinned (adults) for permanent preservation.
For exuviae, the method we use for slide mounting
consists in transferring the specimen from the 70% ethanol
to absolute ethanol for about 30 min, transfer it to a small
drop of Euparal (dorsal side up for pupa, ventral side up for
larva) with a small piece of glass shard on each side, to keep
the cover slip from crushing the skin. It is common to
separate the respiratory organs of pupae and place them by
the side of the body, under the same coverslip. Whole
specimens must be cleared in advance, using KOH (10%).
753
REFERENCES
Alto, B.W., M.W. Griswold & L.P. Lounibos. 2005. Habitat complexity
and sex-dependent predation of mosquito larvae in containers.
Oecologia 146: 300e310.
Amaral, A.P. & L.C. Pinho. 2015. New species and records of frog-biting
midges from southern Brazil (Diptera: Corethrellidae). Zootaxa 3946:
274e284.
Baranov, V.A., G.M. Kvifte & E.E. Perkovsky. 2016. Two new species of
fossil Corethrella Coquillett from Late Eocene Rovno amber, with a
species-level phylogeny for the family based on morphological traits
(Diptera: Corethrellidae). Systematic Entomology 41: 1e11.
Belkin, J.N. 1962. The Mosquitoes of the South Pacific (Diptera, Culi-
cidae). Volume 1. University of California Press, Berkeley and Los
Angeles, CA.
Borkent, A. 2008. The frog-biting midges of the world (Corethrellidae:
Diptera). Zootaxa 1804: 1e456.
Borkent, A. 2012. The pupae of Culicomorpha e Morphology and a New
Phylogenetic Tree. Zootaxa 3396: 1e98.
Borkent, A. & G.R. Spinelli. 2007. Neotropical Ceratopogonidae
(Diptera: Insecta). Pages 1e198 in: J. Adis, J.R. Arias, G. Rueda-
Delgado and K.M. Wantzen (eds.), Aquatic Biodiversity in Latin
America (ABLA). Volume 4. Pensoft, Sofia-Moscow.
Caldart, V.M., M.B. Santos, S. Iop, L.C. Pinho & S.Z. Cechin. 2016.
Hematophagous flies attracted to frog calls in a preserved seasonal
forest of the austral Neotropics, with the description of a new species of
Corethrella (Diptera: Corethrellidae). Zoological Science 33: 527e536.
Griswold, M.W. & L.P. Lounibos. 2005. Competitive outcomes of aquatic
container Diptera depend on predation andresource levels. Annals of
the Entomological Society of America 98: 673e681.
Lane, J. 1953. Neotropical Culicidae. Volume 1. University of São Paulo.
Lane, J. & T.H.G. Aitken. 1956. Chaoborinae from Trinidad, B.W.I.
(Diptera, Culicidae). Annals of the Entomological Society of
America 49: 530e543.
McKeever, S. & F.E. French. 1991. Corethrella (Diptera: Corethrellidae)
of North America north of Mexico: distribution and morphology of
immature stages. Annals of the Entomological Society of America
84: 522e530.
Robert V. & A. Borkent. 2014. First record of Corethrellidae (Diptera),
the frog-biting midges. Malagasy Nature 8: 49e53.
Silva, P.D. & X.E. Bernal. 2013. First report of the mating behavior of a
species of frog-biting midge (Diptera: Corethrellidae). Florida
Entomologist 96: 1522e1529.
Wood, D.M. & A. Borkent. 1989. Phylogeny and classification of the
Nematocera. Chapter 114, Pages 1333e1370 in: J. F McAlpine and
D. M. Wood (eds.). Manual of Nearctic Diptera. Volume 3. Agri-
culture Canada Monograph 32.
Yu, Y.-X., E.-J. Huang & J.-Q. Zhang. 2013. [A new species of
Corethrellidae in China Corethrella feipengi sp. nov. (Diptera:
Corethrelliidae).] Acta Parasitologica et Medica Entomologica Sinica
FAMILY CORETHRELLIDAE
20: 261e264 (in Chinese, English summary).