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Chapter 16 - Order Diptera

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 Chapter 16

Order Diptera
Lı́via Maria Fusari
Depto. de Hidrobiologia, Universidade Federal de São Carlos, São Carlos, São Paulo, Brazil

Galileu P.S. Dantas

Coordenação de Biodiversidade, Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil

Luiz Carlos Pinho

Departamento de Ecologia e Zoologia, Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina, Brazil

Chapter Outline Muscidae 610

Introduction 607 Scathophagidae 610
Tipulidae 608 Sarcophagidae 610
Limoniidae 608 Sciomyzidae 610
Tanyderidae 608 Limitations 610
Ptychopteridae 608 Terminology and Morphology 610
Scatopsidae 609 Material Preparation and Preservation 612
Athericidae 609 Preservation 612
Rhagionidae 609 Slide Preparation 613
Tabanidae 609 Keys to Larvae of Aquatic and Semiaquatic Diptera 619
Pelecorhynchidae 609 Diptera: Suborders 619
Empididae 609 Diptera: “Nematocera”: Families 619
Dolichopodidae 609 Diptera: Brachycera: Families 620
Lonchopteridae 610 References 621
Phoridae 610

INTRODUCTION Diptera is not restricted to the considerable number of

Diptera representatives (flies, mosquitoes, and associated)
are holometabolous insects, distinguished from others by
the presence of a single pair of functional wings in adults.
This characteristic gives the order its name: “two” (di-)
and “wings” (ptera). Anterior wings are prominent, and
the posterior wings, usually found in other insects, in
dipterans are differentiated into small structures known as
halteres. These halteres have balancing functions during
flight.
This order, comprising over 153,000 species distributed
species, but it also includes a wide variety of adaptations,
which reflect the large morphologic and ecologic diversity.
Groups with aquatic members are found in a wide range of
environments, both natural and artificial (e.g., rivers,
streams, ponds, phytotelmata, and temporary pools), as
well as in marine habitats known for harboring few insect
groups.
Adults of most mosquitoes, black flies, biting midges,
deer flies, horse flies, and some other aquatic families feed
on the blood of humans, livestock, and wild animals. These
ORDER DIPTERA

dipterans are vectors for several diseases, such as malaria,

in approximately 180 families (Brown et al., 2009),
currently forms one of the most diverse group of insects. In
the Neotropical region, there are over 31,000 species
recognized in 118 families (Amorim, 2009b); however,
these numbers are already obsolete. The diversity of
yellow fever, dengue fever, chikungunya fever, and leish-
maniasis. Midges and other aquatic flies are of general
importance to freshwater ecosystems, often being a major
fish food and an important part of bottom-dwelling
communities.

Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00029-9 607

Copyright Ó 2018 Elsevier Inc. All rights reserved.
 608
Among aquatic insect immatures, dipteran larvae are
easily recognizable through the complete absence of arti-
culated thoracic appendages (legs). Apod larvae (those
lacking legs) also occur in some other orders, such as
Siphonaptera, and some Hymenoptera and Coleoptera, but
hardly any species of such groups inhabit aquatic or
semiaquatic environments. Immature stages of aquatic flies
can be found in virtually every type of aquatic habitat.
Among Diptera families recorded for the Neotropical
region, 31 show larvae inhabiting a wide variety of aquatic
environments. They are found on or within the substrate, on
or in aquatic plants, swimming or floating in open water, or
associated with the airewater interface.
Diptera have been divided into “Nematocera” (“lower”
Diptera) and Brachycera (“higher” Diptera). Brachycera is
monophyletic, but the lower Diptera is paraphyletic, lack-
ing any unifying characters (synapomorphies). This implies
that Brachycera originated within lower Diptera. In this
chapter, the lower Diptera larvae are considered in the
following suborders, Culicomorpha, Blephariceromorpha,
Tipulimorpha, Ptychopteromorpha, and Psychodomorpha,
as suggested in Carvalho et al. (2012).
There are 31 aquatic or semiaquatic Diptera families in
the Neotropical region, 14 of them are treated in detail in
this book; see Chapters 16.1e16.14, listed subsequently:
Chapter 16.1 - Ceratopogonidae
Chapter 16.2 - Chironomidae
Chapter 16.3 - Simuliidae
Chapter 16.4 - Thaumaleidae
Chapter 16.5 - Culicidae
Chapter 16.6 - Dixidae
Chapter 16.7 - Corethrellidae
Chapter 16.8 - Chaoboridae
Chapter 16.9 - Blephariceridae
Chapter 16.10 - Psychodidae
Chapter 16.11 - Stratiomyidae
Chapter 16.12 - Syrphidae
Chapter 16.13 - Periscelididae
Chapter 16.14 - Ephydridae
Characteristics of the other 17 families, for which there is
very scarce information on the aquatic or semiaquatic
immature forms, are summarized next.
Tipulidae
It is cosmopolitan with 4,091 species described in 113
genera. In the Neotropical region, there are 838 species in
ORDER DIPTERA
14 genera (Oosterbroek, 2017). Larvae are elongated, with
a distinctive retractile head capsule. Tipulids inhabit habi-
tats that range from strictly aquatic or accumulated water in
plants (phytotelmata), to completely terrestrial, as in
decomposing vegetable material, wood, or humid soil near
streams (Alexander & Alexander, 1970). Aquatic species
Thorp and Covich’s Freshwater Invertebrates
are shredders of fallen leaves and inhabit a large variety of
environments, from fast current and very oxygenated
streams, to temporary lakes and ponds, where they gener-
ally colonize the areas with vegetal detritus.
Limoniidae
It is the largest family in Tipulomorpha, comprising more
than 10,000 species described worldwide. In the Neotrop-
ical region, 2,801 species are recognized in 65 genera
(Oosterbroek, 2017). Larvae are hemicephalic, with two
lateral and one dorsal, very well sclerotized sclerites.
Mandibles of detritivore species are robust, with widened
teeth, while predaceous species show slender and falciform
mandibles. Larvae are metapneustic, rarely apneustic, and
the last terminal segment bears disc-shaped spiracles.
These are covered with hydrophobic secretions and are
surrounded by five lobular projections. Species of this
family are aquatic or semiaquatic and colonize habitats that
go from deposit areas to vegetal detritus in the shores of
rivers and lentic environments, and hygropetric surfaces.
Tanyderidae
Species in this family (primitive crane-flies) are mid-sized
to large nematocerans, resembling Tipulidae. It is
comprised of 38 extant species divided among 10 genera,
and three species have a Neotropical distribution (Madriz &
Courtney, 2016). Larvae of five genera are known: they
occur in two types of habitats, the hyporheic zone of cobble
and sand bottom streams, or in the outer layers of
submerged rotting wood (Exner & Craig, 1976; Krzemi nski
& Judd, 1997). They are found in unpolluted mountainous
streams.
Ptychopteridae
This family is composed of 78 species described world-
wide, recorded in three genera: Bittacomorpha, Bittaco-
morphella, and Ptychoptera (Pape & Thompson, 2013); but
in the Neotropical region, the family has been recorded by
Hancock et al. (2006), after the discovery of adults of
Ptychoptera from rain forest habitat in southeastern
Mexico, and recently by Fasbender & Courtney (2017) with
a record of Bittacomorpha from Costa Rica. The oviposi-
tion occurs in the limit between still water and sediment
where their larvae and pupae live. Long respiratory tubes
are used to breathe atmospheric oxygen. The specificity of
their habitat is likely to be high, and habitats may be
restricted (Wagner et al., 2008). Larvae are associated with
saturated mud or soils near water (Brindle, 1962). They are
considerably caudally elongated, presenting a superficial
resemblance to the long-tailed larvae of the Eristalinae
(Syrphidae). To cope with respiration after pupation, the
Chapter | 16 Order Diptera
right thoracic breathing horn is greatly elongated, up to
twice the length of the pupa, whereas the other horn is
atrophied.
Scatopsidae
This family is cosmopolitan, with 400 described species in
36 genera (Amorim, 2016). There are 67 named scatopsid
species in the Neotropical region, but more than 130
undescribed species are known (Amorim, 2009c).
Scatopsid larvae live in decaying plant or animal material,
humid wood holes, and excrement (Cook, 1967; Amorim,
2009a). Few larvae are aquatic living under the surface of a
thin water film among waterlogged dead tree leaves, while
few others are dendrolimnobiontic (e.g., inhabit phyto-
telmata) (Wagner et al., 2008). Pupae are sometimes found
involved in the exuvia of the last larval instar (Amorim,
2009a).
Athericidae
Worldwide, there are 123 recorded species in nine genera.
The 27 Neotropical species are described in four genera:
Atherix, Dasyomma, Suragina, and Xeritha. Neotropical
species has no larvae described formally; however,
immature stages of Athericidae from Brazil were recently
collected in streams, and in other regions, immature stages,
so far as known, are strictly aquatic. Larvae are usually
found in rocky or gravelly riffle areas (Woodley, 2009) and
are predaceous, feeding on larvae of other aquatic insects
(Rozkosny & Nagatomi, 1997).
Rhagionidae
The family have about 720 species in 22 genera worldwide
(Kerr, 2009). Most representatives of this family are
terrestrial, with a single aquatic genus, Chrysopilus (Kerr,
2009; Oscoz et. al. 2011). In the Neotropical region, 70
species were reported in Chrysopilus (Santos, 2008).
Larvae of Neotropical species are unknown, but larvae of
other regions are recorded for lake and river shores, but also
in areas where rock substrates are covered with a thin layer
of water, or between aquatic bryophytes. They can also
develop on humid soils and are predators that feed on
aquatic invertebrates (Oscoz et. al., 2011).
Tabanidae
This family has global distribution, with approximately
4,500 species distributed in 230 genera; in the Neotropical
region, there are over 1,200 species described (Carvalho
et al., 2012). Most females are hematophagous and have
mammals, reptiles, and even birds as hosts (Ferreira et al.,
2002; Oliveira et al., 2002). Larvae are fusiform
609
(Chrysopsinae, Tabaninae) or anteriorly and posteriorly
widened (Pangoniinae). Their coloration varies from white,
green, to dark brown. They have 11 segments, a completely
retractile head capsule, and partially retractile respiratory
siphon on the last abdominal segment. They are known to
be predatory and to practice cannibalism (Ferreira &
Rafael, 2006), but some species are detritivores. Many
species, while in the larval phase, inhabit humid soil, and
they are found in fresh water, saline swamps, and in the
banks of rivers and ponds. Only a few species are restricted
to sand and gravel (Pechuman & Teskey, 1981). Many
larvae inhabit specialized niches, such as holes in trees,
decomposing trunks, bromeliads, or submerged aquatic
vegetation.
Pelecorhynchidae
This family is composed of 49 species distributed in two
genera and has worldwide distribution. In the Neotropics,
nine species of Pelecorhynchus are reported (Llanos et al.,
2015), with distribution exclusive to Chile. Immature
stages of Pelecorhynchidae from Chile were recently
known, and only some immatures of the Pelecorhynchus
from Australian species are known (Mackerras & Fuller,
1942; Llanos et al., 2015). Larvae were collected in humid
margins of swampy areas, where they feed on earthworms
and possibly other soft-bodied animals (Mackerras & Ful-
ler, 1942). They move on to dryer levels to pupate. Larvae
and pupae are similar to those of Tabanidae (Mackerras &
Fuller, 1942).
Empididae
Empididae are cosmopolitan, with about 3,051 species
worldwide in 68 genera (Pape & Thompson, 2013). In the
Neotropical region there are approximately 500 species in
32 genera. Adults are mostly predaceous, and some species
feed on pollen and nectar. Known larvae are predaceous,
living in aquatic and terrestrial environments. Various
species that have aquatic larvae prey on simuliid pupae
(Hamada, 1993).
Dolichopodidae
This family is greatly diverse in tropical areas. There
are 7,600 known species worldwide (Yang et al., 2006;
ORDER DIPTERA
Grichanov, 2014), of which 1,200 are Neotropical
(Capellari, 2016). Adults prey on other smaller insects, and
they are often observed on the dorsal face of foliage, trunks,
stones, etc. Larvae are predominantly predaceous, and
some are phytophagous. Larvae are found in mud,
decomposing wood, under bark, and soil litter.
 610
Lonchopteridae
About 58 species are known worldwide, all of them cited
mostly in a single genus (Carvalho et al., 2012),
Lonchoptera. An aquatic developmental stage is known for
Lonchoptera nigrociliata Duda and L. lutea Panzer
(Wagner et al., 2008; Vaillant & Rozkosný, 2002). Only
L. furcata (Fallén) was recorded in the Neotropics (Amorim
et al., 2002). The biology of species in this family is not
well known. Larvae live in a wet microenvironment of
decaying organic matter, among dead leaves, in plant
debris, under stones on shores of water courses, or they are
semiaquatic. They are saprophagous, microphagous, or
presumably, mycetophagous.
Phoridae
Phoridae is a cosmopolitan family with about 4,087 species
in 271 genera (Pape & Thompson, 2013); there are about
225 genera reported for the Neotropical region (Amorim
et al., 2002). They are considered to be one of the groups
with the most diverse biology among all insects, being
found in several different types of environments, such as
decomposing bodies, rodent and bird nests, flowers, fungi,
wasp, ants, termite nests, and in caves; and with the greatest
diversity of larval habits, but it is the least studied of the
large families of flies due to identification difficulties.
There are many genera that include species with larvae that
are parasitoid of other insects, spiders, and myriapods, and
others that are parasites or commensals in ant or termite
nests. True aquatic larvae are found only in phytotelmata
(Disney, 2004).
Muscidae
The family is cosmopolitan with about 5,190 species (Pape
& Thompson, 2013) and 846 Neotropical species (Carvalho
et al., 2005). The phylogeny of the family has been inves-
tigated by Carvalho (1989), Couri & Carvalho (2003), and
Haseyama et al. (2015). Adults have extremely diverse
habits, which include some predaceous species. Adults in
most species feed on decomposing animal or vegetable
matter (Carvalho & Couri, 2002). Muscid larvae are
coprophagous, saprophagous, or carnivorous (Skidmore,
1985). Larvae of aquatic muscid species can be found in
ORDER DIPTERA
running and standing waters in streams and ponds. They are
generally collected in decaying organic matter and algal
mats, presumably where they find their prey. A few species
of Muscidae are truly aquatic and prey primarily on other
Diptera. The aquatic larvae of this group are not well
studied.
Thorp and Covich’s Freshwater Invertebrates
Scathophagidae
The family has about 404 species worldwide in 57 genera
(Pape & Thompson, 2013). About six species were recor-
ded in Neotropical region, being divided in three genera,
Cordilura, Scathophaga, and Scatogera (Vockeroth, 2010;
Wolff et al., 2016). Of these, only Scathophaga include
truly aquatic larvae, but their biology and ecology are still
poorly described (Oscoz et. al., 2011). Females enter the
waters of streams and ovoposit their eggs on the surface of
the eggs masses of other invertebrates such as Trichoptera
and other Diptera. The hatched larvae feed on the eggs and
surrounding jelly of the host (Purcell et al., 2008).
Sarcophagidae
The family has more than 3,000 described species (Pape
et al., 2011), distributed almost worldwide, but the greatest
diversity is found in the tropical regions. About 750 species
are known to occur in the Neotropical region, most found in
tropical or warmer temperate areas. In aquatic environ-
ments, some larvae may be found mining bases of pitcher
plants (Courtney & Merritt, 2008).
Sciomyzidae
It is cosmopolitan and has approximately 600 species
described in 67 genera. In the Neotropical region, there are
74 species in 23 genera (Amorim et al., 2002). Larvae of
most species are primarily parasitoid or predators of
aquatic or terrestrial mollusks, including a few species
of operculate mollusks. The feeding habits of larvae make
it possible for them to be used in the biologic control of
vectors of parasitic diseases, such as fascioliasis and
schistosomiasis (Carvalho et al., 2012).
LIMITATIONS
Most systematic studies of Neotropical Diptera have
focused on adult morphology, so the knowledge of aquatic
stages of many families is fragmentary. Based on this fact,
we included in the key to Neotropical families those aquatic
larvae that are reported elsewhere, even though the larval
habitats of the species in this region are not currently
known (e.g., Pelecorhynchidae and Lonchopteridae).
TERMINOLOGY AND MORPHOLOGY
Larval dipterans are characterized by a highly diverse body
shape that is fusiform (Tabanidae, Dolichopodidae),
cylindrical (Chironomidae, Empididae), or sometimes
anteriorly tapered (Muscidae, Phoridae) or strongly
dorsoventrally flattened (Stratiomyidae). Others, such as
Culicidae, Chaoboridae, and Corethrellidae, have an
Chapter | 16 Order Diptera
enlarged thorax, while Simuliidae have an enlarged
abdomen (Teskey, 1981). The most common body
segmentation pattern is that with 12 segments, 3 of which
are thoracic, and 9 abdominal. There are numerous varia-
tions to this basic plan. In Brachycera, for example, there
are frequently only 11 obvious segments. In Blepharicer-
idae, in which the body is divided in six parts, the first one
is correspondent to the entire thorax and the first abdominal
segment. Similarly, pseudo-segmentation also occurs, as
observed in Psychodidae, in which each segment has two or
three subdivisions (Teskey, 1981; Courtney & Merritt,
2008). The cephalic head can be complete, sclerotized and
exposed, with biting and chewing mouthparts, or reduced in
many levels, partially or entirely inserted in the thorax, with
mouthparts modified for stinging and scraping (Teskey,
1981; Guimarães & Amorim, 2006). Three states of head
capsule reduction illustrate these patterns, as describe next.
Eucephalic larvae. These exhibit a totally exposed,
well-developed, and sclerotized cephalic capsule (except
for Tipulidae, in which the head is inserted in the thorax and
sometimes reduced, e.g., Fig. 16.2 C; and Blephariceridae,
that show reduction in the dorsal region of the cephalic
capsule, e.g., Fig. 16.3 A), with mandibles operating in a
horizontal or oblique plan (Figs. 16.2 A, 4 A, G, J).
Hemicephalic larvae. These have a relatively reduced
head that is posteriorly incomplete and partially retracted in
the thorax, with mandibles or mouth hooks in the shape of
sickles, operating in a vertical plan (Figs. 16.6 A, I, 7 A, D).
Acephalic larvae. These show additional reduction and
almost total retraction of cephalic capsule in the thorax. The
cephalic thorax is modified in an internalized cephalophar-
yngean skeleton, of which only the vertical, sickle-shaped
mandibles are exposed (Figs. 16.8 A, C, 9 E, M).
FIGURE 16.1 Position of respiratory apertures (spiracles) in Diptera larva: (A) peripneustic; (B) amphipneustic; (C) metapneustic; (D) apneustic.
611
True thoracic appendages (articulated legs) are entirely
absent in all Diptera larvae. Locomotion is conducted by
pseudopods, various fleshy projections, typically bearing
hairs or spines, present in anterior and posterior regions of
the larva (Figs. 16.5 A, C, D, F), or through creeping welts,
normally in greater number than pseudopods, and present
in ventral intersegmental regions (Figs. 16.5 E, H). Such
protrusions work as supporting points that, together with
peristaltic motions of the body, allow for larval locomotion.
Mandibles can aid this process, by becoming additional
anchoring points (Teskey, 1981). Larvae show a variety of
respiratory adaptations. Respiration may be directly from
the atmosphere (e.g., Dixidae, most Culicidae), from plant
tissues (e.g., some Culicidae), or from the surrounding
water. According with the number and position of respi-
ratory apertures (spiracles), aquatic dipteran larvae can be
the following (Fig. 16.1):
Peripneustic. Loss of the posterior thoracic spiracles
yields the peripneustic system, which is characteristic of
the larvae (Fig. 16.1 A). An example is Scatopsidae.
Amphipneustic. These Diptera are characterized by
spiracles on the first thoracic segment and also in the pos-
terior body area (Fig. 16.1 B). Examples are Psychodidae,
Syrphidae, and some Muscidae.
Metapneustic. Such dipterans have spiracles limited to
the posterior body area (Fig. 16.1 C). Examples include
Culicidae, Tabanidae, and certain Muscidae.
Apneustic. Functional spiracles are absent in these
Diptera. In this case, gas exchanges occur throughout the
entire tegument or in dedicated areas in the tegument,
commonly named branchiae (Fig. 16.1 D). Examples are
Chironomidae, Simuliidae, and Blephariceridae. In several
groups the spiracles are located at the end of a retractile
ORDER DIPTERA
 612 Thorp and Covich’s Freshwater Invertebrates

FIGURE 16.2 Tipulidae larva: (A) habitus, lateral view; head capsule retracted within thorax; (B) Posterior end of larva; (C) spiracular disk, in posterior
view. Limoniidae larva (D) larval habitus; (E) head capsule fully exposed; (F) head capsule, dorsal view. Abbreviations: ag, anal gills; ant, antenna; md,
mandible; mxp, maxillary palp; scl, sclerites; sp, spiracle; spl, spiracular lobes.

respiratory siphon (respiratory tube or breathing tube).
Examples are Ptychopteridae, Ephydridae, and Syrphidae.
Further information on the morphology of Diptera larvae is
provided by Teskey (1981) and Guimarães & Amorim
(2006).
ORDER DIPTERA
study goal. Alcohol-preserved immatures must be accom-
modated in tightly sealed jars, so that volatilization is
reduced. It is frequently impossible to sort immatures while
in the field; therefore biologic material can be fixed in 96%
ethyl alcohol, along with the substrate in which it has been
collected. After sorting, immatures should be preserved in
80% ethyl alcohol. Depending on the purpose of the study,

MATERIAL PREPARATION AND other fixative solutions may be necessary. For example, to
PRESERVATION do chromosomal studies (e.g., on Chironomidae or Simu-
liidae), organisms must be fixed directly in Carnoy solution
Preservation (3:1; absolute ethyl alcohol and glacial acetic acid).
After sampling, immatures have to be fixed and preserved Molecular analyses require that organisms are fixed
using the appropriate method for each taxonomic group and directly in ethylic alcohol. In both cases, fixative solutions
Chapter | 16 Order Diptera 613

FIGURE 16.3 Blephariceridae larva: (AeB) ventral view. Psychodidae larva: (C) habitus, lateral view; habitus in (D) ventral view; (E) dorsal view;
(F) habitus, lateral view. Scatopsidae larva: (G) habitus, dorsal view. Ptychopteridae larva: (H) habitus, lateral view. Tanyderidae larva: (I) habitus, lateral
view. Abbreviations: asp, anterior spiracle; d. plt, dorsal plate; psp, posterior spiracle; pro, proleg; suc. d., suctorial disc.

must be changed at least three times, and material must be
stored in low-temperature ( 20 C) and light-free
environments.
Slide Preparation
The preparation of permanent or semipermanent slides
requires that immature parts be separated (e.g., cephalic
capsule and abdomen). For the observation on optical mi-
croscopes, muscles can be dissolved, which results in
clarification. This process is used particularly for perma-
period of 1e2 h (always following the process, ensuring
that the medium does not evaporate completely). Heating
time used for clarification will depend on the material’s size
and degree of sclerotization. Rinse the specimen or struc-
ture in water and transfer to an alcohol series (ethyl alcohol
70%, 80%, 90%, and absolute). The material should remain
soaked in each alcohol concentration for 15e20 min. to
ensure material dehydration.
Hot potassium hydroxide (KOH) 10% (or 5%)
solution. Heat insect or desired structure in KOH on a hot
plate, for a period of 1e2 h (always following the process,
ORDER DIPTERA

nent slide mounting.
There are various methods for the clarification of
specimens or selected structures before slide preparation.
Three techniques are introduced as follows:
Lactic acid 85%. Heat insect or desired structure in
lactic acid in a porcelain crucible, using a hot plate, for a
ensuring the structure is not overly transparent). Heating
time used for clarification will depend on the material’s size
and degree of sclerotization. Rinse the specimen or struc-
ture in water, neutralize for 15 min in acetic acid 10% or
15%, and rinse in water again. Transfer the material to an
alcohol series (ethyl alcohol 70%, 80%, 90%, and
 614 Thorp and Covich’s Freshwater Invertebrates
ORDER DIPTERA

FIGURE 16.4 Culicidae larva: (A) habitus, dorsal view; (B) anal region, lateral view; (C) head, thorax, and six abdominal segments, in dorsal view.
Chaoboridae larva: (D) habitus, lateral view; head and thorax in (E) lateral view and (F) dorsal view. Corethrellidae larva (G) habitus, dorsal view;
(H) head, dorsal view. Dixidae larval habitus in (I) lateral view and (J) ventral view; (K) terminal segments, dorsal view. Abbreviations: ant, antenna; pro,
proleg; psl, postspiracular lobe.
Chapter | 16 Order Diptera 615

FIGURE 16.5 Ceratopogonidae larva: (AeC) habitus, lateral view. Chironomidae larva: (DeE) habitus, lateral view. Simuliidae larva habitus,
(F) lateral view and (G) dorsal view. Thaumaleidae larva: head and thorax in (H) dorsal view and (I) in lateral view; (J) habitus, lateral view. Abbre-
viations: asp, anterior spiracle; lf, fans labral; pro, proleg; psp, posterior spiracle.
ORDER DIPTERA

absolute). The material should remain soaked in each
alcohol concentration for 15e20 min to ensure material
dehydration.
Cold KOH 10% solution. Immerse specimen or desired
structure in KOH for w12e24 h (it is important to follow
the process, to ensure the structure will not become overly
transparent). Immersion time used for clarification will
depend on the material’s size and degree of sclerotization.
Following this process, the same washing protocol used for
heated 10% KOH should be used. Sometimes, it is necessary
to remove with dissection needles the undissolved muscles
from clarified specimens or separated structures.
 616 Thorp and Covich’s Freshwater Invertebrates
ORDER DIPTERA

FIGURE 16.6 Tabanidae larva: habitus, (A) dorsal view, (B) lateral view; (C) respiratory siphon and spine; (D) lateral view of head capsule;
(E) respiratory siphon. Pelecorhynchidae larva: (F) habitus, lateral view; (G) habitus; (H) head capsule, ventral view. Athericidae larva: (I) habitus, lateral
view; (J) habitus, dorsal view; (K) posterior region of abdomen, dorsal view. Rhagionidae larva: (L) habitus, lateral view, with detailed posterior segments.
Abbreviations: br, mandibular bristles; cl, clypeus; cr. w., creeping welt; md, mandible; pro, proleg; sb, stigmatal bar; sm, submentum; tr, tentorial rod.
Chapter | 16 Order Diptera 617

FIGURE 16.7 Stratiomyidae larva: habitus, (A) dorsal view; (B) lateral view. Empididae larva: (C) head capsule, ventral view; (DeF) habitus, lateral
view. Dolichopodidae larva: (GeH) habitus, lateral view. Abbreviations: asp, anterior spiracle; cl, clypeus; cr. w., creeping welt; md, mandible; mr,
metacephalic rod; pro, proleg; psp, posterior spiracle; tr, tentorial rod.

ORDER DIPTERA

FIGURE 16.8 Syrphidae larva: habitus, (A) lateral view, (B) dorsal view. Phoridae larva: (C) habitus, dorsal view; (D) posterior spiracles. Lon-
chopteridae larva, habitus, in dorsal view. Abbreviations: psp, posterior spiracle; res. tube, respiratory tube.
 618 Thorp and Covich’s Freshwater Invertebrates

FIGURE 16.9 Sciomyzidae larva: (A) head capsule, lateral view; (B) habitus, lateral view. Sarcophagidae larva: (C) habitus, lateral view; (D) posterior
ORDER DIPTERA

spiracles. Ephydridae larva: habitus, (E) lateral view; (F) dorsal view; (GeH) habitus, lateral view, with detail of respiratory tube. Muscidae larva: (IeJ)
habitus, lateral view. Scatophagidae larva: (K) typical bicornuate anterior spiracle. Periscelididae larva: (L) habitus, dorsal view, with detail of anterior
spiracles. Abbreviations: res. tube, respiratory tube; ven. arch, ventral arch.
Chapter | 16 Order Diptera 619

For permanent slides, the most common preparation
mediums are Euparal and Canada Balsam. Euparal is
soluble in alcohol and, therefore, specimens can be trans-
ferred directly into this product after dehydration. For
preparation in Canada Balsam, it is necessary that the
dehydrated structure undergo creosote or eugenol treatment
before specimens are mounted into slides. Following
preparation, slides must be incubated at 50 C for drying.
For semipermanent slides, the most frequently used
medium for this type of preparation is Hoyer (solution of
chloral hydrate, distilled water, glycerin, and gum arabic).
Chironomidae larvae are normally mounted using this
protocol (Trivinho-Strixino & Strixino, 2011; see Chapter
16.2 on Chironomidae).
Instant analysis with temporary slides in optical mi-
croscopes can be performed by mounting the specimen in
an excavated slide with glycerin or a 1:1 mixture of glyc-
erin and alcohol, and then covering this gel with a glass
slide. Subsequently, specimen is stored in a microtube with
80% alcohol.

KEYS TO LARVAE OF AQUATIC AND SEMIAQUATIC DIPTERA

The following keys to suborders and families pertain only to taxa recorded in the Neotropical region. These keys are modified from Pinho (2008) and
Pepinelli & Pinho (2014). Those keys were, in turn, based on Courtney & Merritt (2008) and have been adjusted according to groups found in the
Neotropical region, both in the description of steps and in the associated illustrations.

Diptera: Suborders
1 Mandibles moving against one another in a horizontal or oblique plane (Figs. 16.2 EeF); head capsule usually complete and fully exposed,
except retracted and reduced in Tipulidae ................................................................................................................ “Nematocera” [p. 619]
1’ Mandibles or mouth hooks moving parallel to one another in a vertical plane (Figs. 16.6 D, H, 7 C); head capsule variously reduced
posteriorly, partially or almost completely retracted within thorax .............................................................................. Brachycera [p. 620]

Diptera: “Nematocera”: Families

1 Head capsule partially to fully retracted within thorax (Figs. 16.2 A, D) (when the specimen is observed alive, you can check the exposed
head capsule), although sometimes head capsule consisting of elongated sclerites (Fig. 16.2 F), posterior spiracles usually bordered by 1e3
pairs of short lobes that are often fringed with short to very long hairs (Figs. 16.2 BeD)......................................................................... 2
1’ Head capsule fully exposed, usually without longitudinal incisions dorsolaterally (Figs. 16.3 C, 4 A), except reduced dorsally in
Blephariceridae (Fig. 16.3 A), posterior spiracles usually without bordering fringed lobes ....................................................................... 3
2(1) Spiracular disk bordered by six (rarely eight) usually subconical lobes, usually two dorsal, two dorsolateral, two ventral spiracles
(Figs. 16.2 BeC); lobes sometimes short and blunt .........................................................................................................................Tipulidae
2’ Spiracular disk, if present, bordered by five (rarely seven) or fewer lobes, usually one dorsomedial, two lateral, two ventral spiracles; lobes
of variable shape..............................................................................................................................................................................Limoniidae
3(1) Head with distinct constrictive separation from thorax (Figs. 16.4 A, G, J); suctorial discs absent ............................................................ 4
3’ Head not distinctly separated from thorax (Fig. 16.3 A); body divided into six major divisions, the first comprising fused head, thorax and
first abdominal segment; each division with a ventromedial suctorial disc (Figs. 16.3 AeB).......Blephariceridae (Chapter 16.9, p. 759)
4(3) Respiratory system not as follows .................................................................................................................................................................. 5
4’ Respiratory system peripneustic (Fig. 16.1 A), with eight pairs of abdominal spiracles; posterior spiracles usually conspicuously larger than
other abdominal spiracles (Fig. 16.3 G) ........................................................................................................................................ Scatopsidae
5(4) Abdomen not as follows.................................................................................................................................................................................. 6
5’ Abdomen ending in a slender, respiratory siphon that when fully exerted is nearly as long as body (Fig. 16.3 H) ............. Ptychopteridae
6(5) Thoracic segments fused and indistinctly differentiated, forming a single segment that is conspicuously wider than abdominal segments
(Figs. 16.4 C, E, G); thoracic and abdominal segments with prominent, lateral flabelliform setal tufts, and/or terminal segment with a setal
flabellum (Figs. 16.4 B, D) ............................................................................................................................................................................. 7
6’ Thoracic segments usually individually distinguishable, thorax about as wide as abdomen, rarely abdomen wider (Fig. 16.4 J); setae on
thoracic and abdominal segments not tufted and terminal segment without setal flabellum........................................................................ 9
7(6) Antennae prehensile (raptorial), with long apical setae (Figs. 16.4 E, H); mouth brushes absent; respiratory siphon absent .................... 8
7’ Antennae not prehensile and with only short apical setae (Figs. 16.4 A, C); prominent oral brushes present on either side of labrum;
respiratory siphon present (Figs. 16.4 AeB) or absent...............................................................................Culicidae (Chapter 16.5, p. 723)
ORDER DIPTERA

8(7) Antennae inserted far apart; head laterally without a transverse setal row; terminal segment with longitudinal flabelliform setal row
(Fig. 16.4 E); thoracic air sacs present (Fig. 16.4 D) or absent............................................................Chaoboridae (Chapter 16.8, p. 755)
8’ Antennae inserted close together; head laterally with a transverse row of spiniform setae (Fig. 16.4 H); terminal segment ventrally with a
tuft of long setae; thoracic air sacs absent (Fig. 16.4 G) ................................................................... Corethrellidae (Chapter 16.7, p. 751)
9(6) Abdominal segments I & II without prolegs; abdomen posteriorly lacking fringed postspiracular lobes ................................................. 10
9’ Prolegs present on abdominal segments I and usually II (Figs. 16.4 IeJ); abdomen posteriorly with two flattened dorsolateral postspiracular
lobes with setose margins (Fig. 16.4 k) ..........................................................................................................Dixidae (Chapter 16.6, p. 747)
 620 Thorp and Covich’s Freshwater Invertebrates

10(9) Prothorax with prolegs (Figs. 16.5 A, D, F) ................................................................................................................................................. 11

10’ Prothorax lacking prolegs (Figs. 16.3 F, I, 5 B) ........................................................................................................................................... 14
11(10) Head capsule lacking labral fans (Fig. 16.5 D, H, I); posterior abdominal segments not markedly broad; terminal ring of hooks absent; anal
prolegs bearing crochets may be present ...................................................................................................................................................... 12
11’ Head capsule usually with a pair of conspicuous labral fans (Fig. 16.5 FeG); abdominal segments 5e8 broad, posterior segment ending in
a ring of hooks............................................................................................................................................Simuliidae (Chapter 16.3, p. 701)
2(11) Respiratory system apneustic (Fig. 16.1 D); prothoracic or anal prolegs usually paired, even if distinction is only a slight separation of the
apical spines (Fig. 16.5 D) ............................................................................................................................................................................ 13
12’ Respiratory system amphipneustic (Fig. 16.1 B); prothorax with a pair of short respiratory tubes dorsolaterally (Fig. 16.5 I); eighth
abdominal segment with a transverse respiratory opening near hind margin, flanked by a pair of sclerotized digitiform processes, bearing
several longer dark setae apically (Fig. 16.5 J); prothoracic and anal prolegs unpaired .................. Thaumaleidae (Chapter 16.4, p. 719)
13(12) All body segments dorsally with prominent tubercles (elevated fleshy processes) and/or setae (Figs. 16.5 A, C)
........................................................................................................................................... Ceratopogonidae (in part) (Chapter 16.1, p. 621)
13’ Body segments lacking prominent dorsal tubercles and setae (Fig. 16.5 D, E) ................................ Chironomidae (Chapter 16.2, p. 661)
14(10) Posterior abdominal segments without long filamentous processes; without prolegs (Figs. 16.3 F, 5 B), or at most, only a single anal proleg
present ............................................................................................................................................................................................................ 15
14’ Posterior abdominal (8) segment laterally with long filamentous processes (Fig. 16.3 I), abdominal segment 9 with pairs of such processes
arising dorsolaterally and from near apex of two elongate cylindrical posteroventrally projecting prolegs.............................. Tanyderidae
15(14) Thoracic and abdominal segments with three dorsal sclerotized plates (Figs. 16.3 C, E); remainder of integument with numerous dark spots
that together with the dorsal plates impart a greyish brown coloration; respiratory system amphipneustic (Fig. 16.1 B); posterior spiracles
usually at apex of a relatively short, conical respiratory siphon; sometimes with ventral suctorial discs (Fig. 16.3 D)
................................................................................................................................................................Psychodidae (Chapter 16.10, p. 765)
15’ Dorsal sclerotized plates absent; integument smooth, shiny, and creamy white, lacking all surface features except a few setae that may be
noticeable at terminal segment tip (Fig. 16.5 B); respiratory siphon absent; ventral suctorial discs absent
........................................................................................................................................... Ceratopogonidae (in part) (Chapter 16.1, p. 621)

Diptera: Brachycera: Families

1 Sclerotized portions of head capsule exposed externally (Figs. 16.6 A, 7A, D), if greatly reduced, with slender tentorial and metacephalic
rods prominent internally (Figs. 16.6 H, 7 C) ................................................................................................................................................ 2
1’ External sclerotized portions of head capsule absent; head reduced to an internal cephalopharyngeal skeleton, with externally exposed
bucal hooks (Figs. 16.9 AeB) ........................................................................................................................................................................ 8
2(1) Larva usually not conspicuously depressed nor with a toughened integument bearing a network of facets; head capsule capable of extensive
independent movement; without distinctive eye prominences ....................................................................................................................... 3
2’ Larva tightly flattened, integument toughened and leathery from calcium deposits that are evident as numerous small reticulately arranged
facets (Figs. 16.7 AeB); head capsule capable of only sight independent movement; usually with distinctive lateral eye prom-
inences ................................................................................................................................................Stratiomyidae (Chapter 16.11, p. 771)
3(2) Head capsule well developed dorsally (Figs. 16.6 A, J), closed ventrally by a submental plate (Figs. 16.6 D, H); tentorial rods solidly fused
with head capsule internally; each side of clypeus above and near base of each mandible usually a brush of backwardly curved bristles
present .............................................................................................................................................................................................................. 4
3’ Head capsule reduced to a pair of slender metacephalic rods; these and tentorial rods flexibly articulate with anterior cephalic sclerites
(Fig. 16.7 C); submental plate and brushes of bristles above mandibles absent ........................................................................................... 7
4(3) Posterior spiracles greatly reduced or situated within a small terminal cavity (Fig. 16.6 F); integument without striations or extensive
covering of pubescence; prolegs, if present, limited to one to two ventral pairs on each abdominal segment............................................ 5
4’ Posterior spiracles present, opening within slits on either side of a vertically linear stigmatal bar (Fig. 16.6 E) or a retractile, laterally
compressed spine (Fig. 16.6 C); body integument with longitudinal striations, except in some species where integument is covered by short,
velvety pubescence; first seven abdominal segments girdled by three or four pairs of fleshy creeping welts (setae-bearing swellings) or
prolegs, these being the only projections from the segments (Figs. 16.6 AeB).............................................................................Tabanidae
5(4) Larva cylindrical, lacking tubercles and prolegs, at most with ventral creeping welts (Figs. 16.6 F, L) ..................................................... 6
5’ Larvae with slender lateral and dorsolateral tubercles of progressive size on abdominal segments 1e7; terminal segment with two longer
tubercles, fringed with setae; all abdominal segments ventrally with pair of elongate prolegs, each proleg with semicircle of subapical and
apical claws (Figs. 16.6 IeJ).......................................................................................................................................................... Athericidae
6(5) Labrum and apically on maxilla laterally with stout spines (Fig. 16.6 H); integument smooth, shiny, segmentation beadlike; respiratory
system apneustic (Figs. 16.6 FeG).......................................................................................................................................Pelecorhynchidae
6’ Labrum and maxilla without stout spines; first seven abdominal segments with ventral creeping welts, segmentation not as previous;
respiratory system metapneustic or amphipneustic (Fig. 16.6 L) ................................................................................................Rhagionidae
ORDER DIPTERA

7(3) Larva usually apneustic, terminal segment with one to four rounded lobes bearing apical setae, and abdominal segments bearing paired
prolegs with apical crochets (Figs. 16.7 D, F); if metapneustic, then terminal segment with only a single lobe below spiracles, and
abdominal segments with ventral creeping welts (Fig. 16.7 E); metacephalic rods slender posteriorly (Fig. 16.7 C)................ Empididae
Chapter | 16 Order Diptera 621

7’ Larva metapneustic; posterior spiracles situated at the base of upper two of four smooth primary lobes of terminal segment (Figs. 16.7
GeH); transverse ventral creeping welts present on abdominal segments; metacephalic rods expanded posteriorly..........Dolichopodidae
8(1) Body not distinctly flattened; all segments tergal plates smooth at margins; first two thoracic and terminal abdominal segments without
slender processes ............................................................................................................................................................................................. 9
8’ Body distinctly flattened; all segments tergal plates striated at margins; first two thoracic and terminal abdominal with slender processes
(Fig. 16.8 E).............................................................................................................................................................................. Lonchopteridae
9(8) Posterior spiracular plates always distinctly separated whether mounted on a telescopic respiratory tube or not .................................... 10
9’ Posterior spiracular plates fused or very closely approximated, usually on apex of a telescopic respiratory tube (Figs. 16.8 AeB)
................................................................................................................................................................... Syrphidae (Chapter 16.12, p. 779)
10(9) Body often dorsoventrally flattened and bearing a series of spicules or tubercles (Figs. 16.8 C, 9 L) ...................................................... 11
10’ Body often cylindrical without spicules or tubercles ................................................................................................................................... 12
11(10) Anterior spiracles simple, each with one to several sensile spiracular openings arrayed peripherally at apex of short tubular or conical
projection; tentoropharygeal and hypopharyngeal sclerites finely constructed and fused together; body often dorsoventrally flattened and
bearing a series of spicules or tubercles (Fig. 16.8 C); posterior spiracles with openings arranged in two pairs placed one behind the other
(Fig. 16.8 D) ........................................................................................................................................................................................Phoridae
11’ Anterior spiracles short, bulbiform with four to five small bulbous papillae; tentoropharygeal and hypopharyngeal sclerites often more
strongly constructed than previous and distinctly separated (Fig. 16.9 L) ....................................... Periscelididae (Chapter 16.13, p. 785)
12(10) Body segments not covered with short, fine setae; if present, a respiratory siphon present, or each spiracle situated on a short tubular
posterior projection (Figs. 16.9 C, E); cephalopharyngeal skeleton lacking a ventral arch........................................................................ 13
12’ Body segments often extensively covered with short, fine setae; abdomen often somewhat tapered posteriorly, its apex with tubercles
surrounding posterior spiracles that are only slightly elevated (Fig. 16.9 B); cephalopharyngeal skeleton with a sclerotized ventral arch
(Fig. 16.9 A) ................................................................................................................................................................................. Sciomyzidae
13(12) Posterior spiracular often well exposed, not localized on cavity; one or more of the spiracular slits nearly horizontal........................... 14
13’ Posterior spiracular often on lowest point in depression on terminal segment (Fig. 16.9 D); spiracular slits inclined
subvertically................................................................................................................................................................................Sarcophagidae
14(13) Abdomen rather truncate posteriorly, with short respiratory tube or basally fused conical spiracular prominences, each bearing a dorsal
tubercle; integument with setae only on intersegmental areas (Fig. 16.9 I); setaceous tubercles, if present, restricted to posterior end
........................................................................................................................................................................................................................ 15
14’ Abdomen somewhat tapered posteriorly, usually ending in an elongate retractile respiratory tube that lacks dorsal tubercles (Figs. 16.9
EeH); integument of posterior abdominal segments covered with setae or spinules, or with setaceous tubercles on some segments
(Fig. 16.9 F)............................................................................................................................................Ephydridae (Chapter 16.14, p. 789)
15(14) Posterior spiracular plate without lobes, spines, or ridges along margins (Figs. 16.9 IeJ) .............................................................Muscidae
15’ Posterior spiracular plate with one or more bordering lobes or spines, or with a sharp ridge along dorsal margin (Fig. 16.9 K)
...................................................................................................................................................................................................Scathophagidae

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