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ADULT MORPHOLOGY AND TERMINOLOGY

ABDOMEN

BODY

HEAD

TORAX

circmf, circumfilum;

comp eye, compound eye;

flgm, flagellomere;

ped, pedicel;

scp, scape;

stm bul, stemmatic bulla.

arlm, arolium;

clw, claw;

emp, empodium;

pulv, pulvillus;

spsq rg, suprasquamal ridge;

tym pit, tympanic pit;

ung plt, unguitractor plate.

cerc, cercus;

gen fk, genital fork;

gen op, genital opening;

hyp vlv, hypogynial valve;

hyprct, hypoproct;

spmth, spermatheca;

spr, spiracle;

st, sternite;

tg, tergite.

abd plaq, abdominal plaque;

acc gl, accessory gland;

acul, aculeus;

cerc, cercus;

cl op, cloacal opening;

epiprct, epiproct;

ev ovp memb, eversible ovipositor membrane;

gen fk, genital fork;

hyprct, hypoproct;

ovscp, oviscape;

rect ppl, rectal papilla;

sg, segment;

spmth, spermatheca;

st, sternite;

tae, taenia;

tg, tergite;

v rep, ventral receptacle.

cerc, cercus;

ej apod, ejaculatory apodeme;

epand, epandrium;

epand lb, epandrial lobe;

hypd, hypandrium;

hypd lb, hypandrial lobe;

hyprct, hypoproct;

pgt, postgonite;

ph, phallus;

ph gd, phallic guide;

phapod, phallapodeme;

pregt, pregonite;

sbepand scl, subepandrial sclerite;

st, sternite;

sur, surstylus;

tg, tergite.

a tnt pit, anterior tentorial pit;

ant soc, antennal socket;

cib, cibarium;

clyp, clypeus;

clyp rg, clypeal ridge;

epiphar bl, epipharyngeal blade;

hyphar, hypopharynx;

lab, labium;

lbl, labellum;

lbr, labrum;

lc, lacinia;

plp, palpus;

premnt, prementum;

psdtrch, pseudotrachea;

tnt, tentorium.

cib pmp, cibarial pump;

clyp, clypeus; epiphar, epipharynx;

fc, face;

fd can, food canal;

fr, frons;

hyphar, hypopharynx;

lab, labium;

lbl, labellum;

lbr, labrum;

md, mandible;

mx, maxilla;

ped, pedicel;

plp, palpus;

sal can, salivary canal;

sen pit, sensory pit;

tm, torma.

acc gl, accessory gland;

acroph, acrophallus;

an, anus;

bac scl, bacilliform sclerite;

cerc, cercus;

distph, distiphallus;

ej apod, ejaculatory apodeme;

ej dt, ejaculatory duct;

epand, epandrium;

epiph, epiphallus;

gls, glans;

hypd, hypandrium;

hypd arm, hypandrial arm;

hyprct, hypoproct;

pgt, postgonite;

ph, phallus;

ph gd, phallic guide;

phapod, phallapodeme;

prens, prensiseta;

pregt, pregonite;

proc, process;

rect, rectum;

spm dt, sperm duct;

spm sac, sperm sac;

st, sternite;

sur, surstylus;

syntgst, syntergosternite;

tes, testis;

tg, tergite;

v d, vas deferens.

aed tn, aedeagal tine;

cerc, cercus;

ej apod, ejaculatory apodeme;

enaed proc, endoaedeagal process;

epand, epandrium;

goncx, gonocoxite;

goncx apod, gonocoxal apodeme;

gonst, gonostylus;

hypd, hypandrium;

hyprct, hypoproct;

lat ej proc, lateral ejaculatory process;

lft cerc, left cercus;

lft epand lam, left epandrial lamella;

lft sur, left surstylus;

pm, paramere;

pm apod, parameral apodeme;

pm sh, parameral sheath;

ph, phallus;

rt cerc, right cercus;

rt epand lam, right epandrial lamella;

rt sur, right surstylus; t

g, tergite.

abd st, abdominal sternite;

abd tg, abdominal tergite;

acr s, acrostichal seta;

adv sut, adventitious suture;

a kepst s, anterior katepisternal seta;

anatg, anatergite;

anepm, anepimeron;

anepst, anepisternum;

anepst s, anepisternal seta;

a pprn s, anterior postpronotal seta;

ap sctl s, apical scutellar seta;

ar, arista;

b pprn s, basal postpronotal seta;

b sctl s, basal scutellar seta;

comp eye, compound eye;

cx, coxa; cxpl

str, coxopleural streak;

dc s, dorsocentral seta;

ds s, discal seta;

ds sctl s, discal scutellar seta;

flgm, flagellomere;

frorb plt, fronto-orbital plate;

fr s, frontal seta,;

fr vit, frontal vitta;

gn dil, genal dilation;

gn grv, genal groove;

gr amp, greater ampulla;

hlt, halter;

ial s, intra-alar seta;

ipal s, intrapostalar seta;

i vt s, inner vertical seta;

kepm, katepimeron;

kepst, katepisternum,

ktg, katatergite;

l m s, lateral marginal seta;

l sctl, lateral scutellar seta;

lun, lunule;

m cx prg, mid coxal prong;

m m s, median marginal seta;

mr, meron;

A1, A2, first and second branches of anal vein;

a1, a2, first and second anal cells;

bc, basal costal cell;

bm, basal medial cell;

bm-cu, basal medial-cubital crossvein;

br, basal radial cell;

C, costal vein;

c, costal cell;

CuA, basal part of anterior branch of cubital vein;

CuA1, CuA2, branches of anterior branch of cubital vein;

cua1 anterior cubital cell;

CuP, posterior branch of cubital vein;

cup, posterior cubital cell;

d, discal cell;

dm, discal medial cell;

dm-cu, discal medial-cubital crossvein;

h, humeral crossvein;

M, medial vein, or media;

M1, M2, M3, branches of vein M;

m1, m2, m3, medial cells;

m-m, medial crossvein;

MA, anterior branch of media;

R, radius, or radial vein;

R1, anterior branch of radius;

Rs, radial sector, branching again into R2, R3, R4, R5, or combinations of these, e.g., R2+3, or R4+5; r1, r2, r3, r4, r5, radial cells;

r-m, radial-medial crossvein;

Sc, subcosta;

sc, subcostal cell;

sc-r, subcostal-radial crossvein.

ar, arista;

clyp, clypeus;

clyp memb, clypeolabral membrane;

comp eye, compound eye;

fc, face;

fc rg, facial ridge;

flg, flagellum;

flgm, flagellomere;

frclyp memb , frontoclypeal membrane;

frgn sut, frontogenal suture;

frorb plt, fronto-orbital plate;

fr s, frontal seta;

fr vit, frontal vitta;

gn gena;

gn dil, genal dilation;

gn grv, genal groove;

gn s, genal seta;

hyps brg, hypostomal bridge;

infr s, interfrontal seta;

i vt s, inner vertical seta;

lbl, labellum;

lbr, labrum;

lun, lunule;

m ocp scl, median occipital sclerite;

oc, ocellus;

ocp, occiput;

ocp for, occipital foramen;

ocp s, occipital seta;

oc s, ocellar seta;

oc tr, ocellar triangle;

o vt s, outer vertical seta;

pafc, parafacial;

pavt s, paravertical seta;

pc orb s, proclinate orbital seta;

ped, pedicel; pgn, postgena;

plp, palpus;

pocl s, postocular seta;

poc s, postocellar seta;

premnt, prementum;

ptil fis, ptilinal fissure;

p tnt pit, posterior tentorial pit;

sbvb s, subvibrissal seta;

spc s, supracervical seta;

u orb s, upper orbital seta;

vb, vibrissa;

vrt, vertex.

a bas, anterior basalare;

abd tg, abdominal tergite;

anatg, anatergite;

anepm, anepimeron;

anepst, anepisternum;

aprn, antepronotum;

a spr, anterior spiracle;

cerv scl, cervical sclerite;

comp eye, compound eye;

cx, coxa;

flgm, flagellomere;

hlt, halter; kepm, katepimeron;

kepst, katepisternum;

ktg, katatergite; lbl, labellum;

l par sut, lateral parapsidal suture;

ltg, laterotergite;

mr, meron;

mtanepst, metanepisternum;

mtepm, metepimeron;

mtg, mediotergite;

mtkepst, metakatepisternum;

mtn, metanotum;

patg, paratergite;

p bas, posterior basalare;

ped, pedicel;

plp, palpus;

plrtrch, pleurotrochantin;

plr wg proc, pleural wing process;

pprn, postpronotum; presct, prescutum;

presct pit, prescutal pit;

presct sut, prescutal suture;

prn, pronotum;

prpl, propleuron;

prst, prosternum;

rst, rostrum;

sbal scl, subalar sclerite;

scp, scape;

sct, scutum;

sctl, scutellum;

trn sut, transverse suture;

wg, wing.

npl, notopleuron;

npl s, notopleural seta;

oc s, ocellar seta;

oc tr, ocellar triangle;

orb plt, orbital plate;

o vt s, outer vertical seta;

pafc, parafacial;

pal cal, postalar callus;

pal s, postalar seta;

pal wall, postalar wall;

pavt s, paravertical seta;

pc orb s, proclinate orbital seta;

ped, pedicel;

plp, palpus;

pocl s, postocular seta;

poc s, postocellar seta;

pprn, postpronotum;

pprn lb, postpronotal lobe;

prepm, proepimeron;

prepm s, proepimeral seta;

prepst, proepisternum;

prepst s, proepisternal seta;

presut acr s, presutural acrostichal seta;

presut sct, presutural scutum;

p spal s, posterior supra-alar seta;

p spr, posterior spiracle;

psut acr s, postsutural acrostichal seta;

psut sct, postsutural scutum;

rc orb s, reclinate orbital seta;

sbap sctl s, subapical scutellar

seta; sbsctl, subscutellum;

sbvb s, subvibrissal seta;

sct, scutum;

sctl, scutellum;

sctsctl sut, scutoscutellar suture;

spal area, supra-alar area;

spal s, supra-alar seta;

spvb s, supravibrissal seta;

trn sut, transverse suture;

vb, vibrissa;

wg b, wing base.

aed, aedeagus;

ap lb goncx, apical lobe of gonocoxite;

b lb goncx, basal lobe of gonocoxite;

cerc, cercus;

clasp, claspette;

ejac apod, ejaculatory apodeme;

epand, epandrium;

epiprct, epiproct;

goncx, gonocoxite;

gonst, gonostylus;

hypd, hypandrium;

hyprct, hypoproct;

lb epand, lobe of epandrium;

pm, paramere;

prct, proctiger;

proc hyprct, process of hypoproct;

rect, rectum;

st, sternite;

tenac, tenaculum;

tg, tergite.

Suborden NEMATOCERA (36 familias)

(=antenas en forma de hilo)

Comprende a las familias más primitivas del orden diptera. en las familias típicas, las antenas, que generalmente son filiformes y son más largas que la cabeza y tórax en conjunto, constan de dos segmentos basales y un flagelo, que generalmente está compuesto de al menos 4 y usualmente 8 segmentos libres. Solo en casos raros el flagelo tiene hasta 39 segmentos y generalmente no presenta un estilo diferenciado.

En las familias de este suborden los palpos son pendulares y están compuestos de 1 ó 5 artejos, excepto en la familia culicidae y en algunas especies de cecidomyiidae.

Las alas carecen de calypteres y las venas cua2 y a1, son divergentes.

La celda discal generalmente esta ausente. la vena rs, generalmente presenta dos ramificaciones, r2+3 y r4+5, pero cuando es triramificada, entonces es la vena r2+3 la que se ha bifurcado.

En la mayoría de las especies el cuerpo es alargado y generalmente no presenta cerdas.

La sutura pleural del tórax es recta, excepto en psychodidae.

Las larvas, muchas de las cuales son de habito acuático, presentan la cabeza excertada, bien conformada, con piezas bucales masticadoras horizontales.

Las pupas son libres, sin pupario.

Los adultos emergen a través de una apertura longitudinal o en forma de T en el aspecto dorsal.

Morphologically, the adults of Tipulomorpha can be recognized by their slender bodies with narrow wings and long legs. However, a lot of variation may occur and species with relatively more stout bodies are common, especially in the Limoniidae. The ocelli is absent in all members of the group, which also have a typical “V”- shaped transverse suture in the mesonotum. Crane flies can be small or large flies with wing lengths spanning from less than 3 mm to over 30 mm.

Infraorder Tipulomorpha Rohdendorf, 1961 (6 families)

Members of the Tipulidae typically have larger sizes as compared with the Limoniidae.

Infraorder Psychodomorpha Hennig, 1968 (5 families)

Infraorden Culicomorpha Hennig, 1948 (9 families)

Infraorder Bibionomorpha Hennig, 1954 (33 families)

Family Cecidomyiidae Newman, 1835 (761 genera, 6,296 species)

Family Keroplatidae Rondani, 1856 (90 genera, 993 species)

Family Mycetophilidae Newman, 1834 (233 genera, 4,525 species)

Family Bibionidae Fleming, 1821 (12 genera, 1,102 species)

Family Anisopodidae Knab, 1912 (24 genera, 196 species)

Family Ditomyiidae Keilin, 1919 (8 genera, 98 species)

Family Sciaridae Billberg, 1820 (92 genera, 2,455 species)

Family Scatopsidae Newman, 1834 (34 genera, 407 species)

Family Ceratopogonidae Newman, 1834 (130 genera, 5,902 species)

Family Simuliidae Newman, 1834 (35 genera, 2,121 species, 6/14)

Family Chironomidae Newman, 1834 (541 genera, 7,290 species)

Family Culicidae Meigen, 1818 (46 genera, 3,725 species)

Family Dixidae Schiner, 1868 (9 genera, 197 species)

Family Blephariceridae Loew, 1861 (39 genera, 331 species, 6/8/2)

Family Psychodidae Newman, 1834

(144 genera, 3,026 species)

Family Tanyderidae Osten Sacken, 1880 (12 genera, 55 species)

Family Limoniidae Rondani, 1856 (188 genera, 10,777 species)

Family Tipulidae Latreille, 1802 (39 genera, 4,415 species)

Family Chaoboridae Newman, 1834 (33 genera, 89 species)

Adult Dixidae are similar to some small Tipulidae but may be distinguished by their more strongly arched R2+3.

Adult Chaoboridae may be initially confused with some Chironomidae but are distinguished by their wing venation, with deeply forked R2+3 and M1+2. Their elongate R1 separates them from species of the Corethrellidae.

In Tipulidae, the terminal segment of the maxillary palps is slender and longer than the penultimate segment, and the antennae are normally 13-segmented.

In the Limoniidae, the terminal segment of the maxillary palps is as short as the others, and the antennae usually have 14 to 16 segments.

Bibionids resemble scatopsids superficially, but bibionidsn are much larger, usually have a 5-segmented palpus, and a more distal cubital fork.

Tanyderidae Osten Sacken 1880 is comprised of 39 extant species divided among 10 genera, with representatives in the Australasian (21 species), Oriental (three species), Afrotropical (one species), Palearctic (six species), Nearctic (four species) and Neotropical (three species) regions.

Adult mosquitoes may be confused at times with adult Tipulidae, Dixidae, or Chaoboridae. The presence of a long scaly proboscis (the rostrum of tipulids, when elongate (Figs. 8.104–105), is devoid of scales), and scale-covered wing veins in mosquitoes set them apart from other families.

Ditomyiidae differ from other Sciaroidea in which R2+3 is present in having Sc weak and ending free, R2+3 well over half the length of the apical part of R4+5, and the wing membrane with dense macrotrichia.

Because of the lack of a discal cell Neomesochria may be confused with some Mycetophilidae, but the Rs fork (R2+3) is at or before crossvein r-m in anisopodids; in mycetophilids the fork (R4) is distal to r-m. The latter usually have long tibial spurs and the head is smaller and tucked at least slightly beneath the thorax.

Adults are similar to some Chironomidae but may be distinguished by the combination of a lack of a medial groove on the postnotum (otherwise only in the rare podonomine Chironomidae) and a lack of a bm-cu crossvein (present in Podonominae).

Adult psychodids are easily recognized by the dense vestiture and the distinctive wing shape and venation. The hump-backed body shape is similar to some Simuliidae or Thaumaleidae. The large number of longitudinal wing veins and the reduced abdominal sternite 1 separates Psychodidae from these families.

Adults are most often confused with culicids, chaoborids or ceratopogonids. In the former two families, the wing margin is reached by 10 vein branches (at most 7 in Chironomidae), and the costa continues around the wing. Ceratopogonidae differ from all chironomids in the combination of postnotum lacking a longitudinal groove and wing without bm-cu.

Cecidomyiid adults can readily be distinguished from other Sciaroidea by the absence of tibial spurs. Not so conspicuous, but nevertheless a good discriminating character, is the fact that the costa completely surrounds the wing, usually with a slight break at its juncture with R4+5.

Adult black flies are readily recognized by their distinctive wing venation, antennal shape, and arched thorax. This combination of characters distinguishes simuliids from bibionids, empidids, phorids, and scatopsids, which they sometimes superficially resemble. The larvae, with their labral fans and single posterior proleg, and the pupae, with their thoracic respiratory organs bearing a basolateral fenestra, are unique among Diptera.

Regional Keroplatidae may be distinguished from the other five families of Sciaroidea by wing venation. Crossvein r-m is absent and R and M are in contact or briefly fused (Fig. 2); the other families all have a distinct r-m crossvein joining these two veins.

Adults are easily recognized by the simple and constant wing venation—a short C (ending at or before the tip of wing), only one short crossvein r-m near the base of wing, two strong sclerotized R-veins joining C, and the weakly sclerotized M- and Cu-forks (Fig. 5)—that is typical for all species and distinguishes the family from other Sciaroidea, including Lestremiinae (Cecidomyiidae).

Adults are distinguished from Scatopsidae and large Sciaridae by the lack of a dorsal eye bridge, eye not emarginate around the base of the antenna, and palpus with 4–5 segments. Larval bibionids are similar to some Sciaroidea but are distinguished by the presence of fleshy tubercles and sclerotized spicules.

Although adults of some species superficially resemble certain tipulids, chironomids, and other nematocerous flies, they are separated by wing venation. Adult Deuterophlebiidae have a netlike wing pattern, but differ in other features (antenna with four flagellomeres, mouthparts vestigial).

Pupae may be confused with those of deuterophlebiids, but their respiratory organs are different (common base and 3–4 tubular filaments). Although true suctorial discs are unique to larval blepharicerids, some psychodids have superficially similar friction pads.

Three dixid genera occur in the Neotropical Region: Dixella Dyar & Shannon, 1924, Mesodixa Belkin, Heinemann & Page, 1970, and Nothodixa Edwards, 1930. Dixella is found in all geographic regions and is the richest genus in Neotropics, with 26 species (Stone, 1966; Belkin et al., 1970; Peters, 1980; Chaverri & Borkent, 2007). Nothodixa is restricted in the Neotropics to southern South America (Chile and Argentina) and is comprised of four species (Stone, 1966). Mesodixa is monospecific and has been reported only from Jamaica (Belkin et al., 1970).

Mycetophilidae differ from Ditomyiidae, Keroplatidae and Diadociidae in having M and CuA connected, if at all, near the level of crossvein h (Figs. 5–36); in the latter three families crossvein bm-cu is present or the two veins come into contact at a point from one-quarter to one-half the wing length. Lygistorrhinidae, which have veins M and CuA obsolete basally, have the mouthparts tubular and longer than the head and thorax, with palpi slender, tapering and arising at the base of the mouthparts. The only New World genus of Mycetophilidae with long tubular mouthparts is Gnoriste Meigen; the palpi are near the apex of the mouthparts and M and CuA are entire. Mycetophilidae have the eyes widely separated above; Sciaridae, the sixth family of Sciaroidea present in tropical America, have the eyes in contact or nearly so above the antennae.

It is cosmopolitan with 4,091 species described in 113

genera. In the Neotropical region, there are 838 species in 14 genera (Oosterbroek, 2017). Larvae are elongated, with a distinctive retractile head capsule. Tipulids inhabit habitats 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

Tanyderidae Osten-Sacken, 1880 is a poorly known, predominantly temperate family represented by 39 extant species divided amongst 10 genera. Because of limited material in collections and the paucity of information about their natural history, the task to find novel morphological and bionomic information is of great importance for further understanding of the family.

Neotropical Tanyderidae are restricted to Chile and Argentina. Since the original discovery of Tanyderus Philippi, 1865, two additional Neotropical genera have been described, Neoderus Alexander, 1927 and Araucoderus Alexander, 1929. Despite considerable interest in the group from a phylogenetic and biogeographical standpoint, information regarding this family has remained fragmentary. Most published information on Neotropical tanyderids is from species descriptions (Philippi 1865; Alexander 1913, 1920) or geographic range extension (Alexander 1935). More recently, the immature stages of Araucoderus and Tanyderus were discovered and described (Lukashevich and Scherbakov 2014, 2016; Madriz and Courtney 2016).

Diagnosis

Tiny, dark flies, body length 0.6–3.0 mm (Fig. 1). Flagellum with 5–10 short flagellomeres (Figs. 2–3). Eye bridge incomplete. Palpus 1–segmented. Anterior spiracle situated at proepimeron (Figs. 4–5). Meron dorsally fused to pleural sclerites (Figs. 5–6). Postnotal phragma reaching abdominal segment 3. Sc incomplete, R5 reaching C close to R1, rarely close to wing apex. M3 absent. Cubital fork displaced to wing base. Anterior veins well sclerotized, posterior veins faint. Membrane and posterior veins usually bare (Figs. 11– 24). Abdominal pleural membrane with longitudinal lines. Female terminalia with T8 usually with spiracles and T10 in Scatopsinae mesally divided (“cerci” in Cook, 1981) (Fig. 9). Male sternite 7 usually ornamented (e.g., Fig. 7). Male terminalia strongly differing among genera. Sperm pump well sclerotized, connected to the terminalia (Fig. 8) only through the sperm duct, not attached to the terminalia as in anisopodids and some limoniids.

Diagnosis

Small to medium flies, body length 3.0–12.0 mm, Olbiogaster and Sylvicola with slender body and legs (Fig. 1), Neomesochria with more compact body. Antennae with 14 flagellomeres, cylindrical, tapering. Eyes dichoptic in females and male Olbiogaster, close or touching in male Sylvicola and Neomesochria. Thorax convex without distinct transverse suture; scutum and scutellum setose, bristles weakly developed. Wing pattern of pigmentation present in Sylvicola; pterostigma absent in Mycetobiinae, microtrichose, macrotrichia only in Sylvicola. M 3-branched in Olbiogaster and Sylvicola, discal cell present; in Neomesochria M 2-branched, faint distally, and discal cell absent (Fig. 2); CuA2 usually sinuous, straight in Neomesochria. Mid- and hind tibia with pubescent apical spurs; in Olbiogaster midtibial spurs of equal size, hind tibia with single spur (Fig. 3); in Sylvicola and Neomesochria spurs of mid- and hind tibiae subequal (Fig. 4); inner apex of hind tibia with comb of bristles; empodium hairy. Olbiogaster males with lateral expansions on last four abdominal tergites. Male cerci prominent in Olbiogaster as are lobes of sternite 10; gonostyli usually well developed except in Neomesochria; aedeagal complex ending internally in long coiled structure in Sylvicola. Female cerci well developed, large in Olbiogaster.

Diagnosis

Adults tiny, fragile (Fig. 1), usually between 1–4 mm long, occasionally larger. Wing veins generally reduced in number; costal vein continuous around wing, usually with a break immediately beyond insertion of R4+5. Eyes usually holoptic or nearly so; ocelli absent except in most Lestremiinae. Antenna with regular number of 10, 12, 13, or 14 flagellomeres, or irregular number from seven to usually less than 20, but as many as 65. Legs with tibial spurs absent; first tarsal segment greatly reduced and shorter than second in all but Lestremiinae.

Larvae with three instars. Head capsule tiny, conical, movable

parts reduced, mandibles internal. Tracheal system amphipneustic in first instar, peripneustic in second and third. Prothorax of third instar, usually, and second instar, rarely, with sclerotized, dermal structure termed a spatula.

Diagnosis

Body length 1.8–8.7 mm. Color yellow to brown or black, often patterned. Ocelli usually three, middle one sometimes small or absent. Antenna no longer than body, with scape, pedicel and usually 14 flagellomeres, in Cordyla with 9–13 flagellomeres. Palpus usually with basal palpifer and four palpal segments, the latter sometimes 1–3 in number. Hairs and bristles present on pronotum, scutum and scutellum; pleural sclerites and prosternum bare or with varied vestiture. Wing membrane with either or both macrotrichia and microtrichia. Venation complete in Neoempheria (Fig. 6) and many other genera, often reduced, strongly reduced in Manota (Fig. 4) and Undescribed Genus A (Fig. 8). Male and female terminalia various, but with sclerites generally as in other nematocerous families. Female with tergite 10 sometimes weak or absent; cercus articulated with last sclerotized tergite, one- or two-segmented; two unsclerotized spermathecae apparently always present.

Diagnosis

Moderate-sized, slender midges (Fig. 1) with wing length about 2–4 mm. Wing venation with R four-branched, M twobranched, Cu two-branched, and with R2+3 strongly arched

(Fig. 2).

Larvae (Fig. 3) distinctive, with elongate antenna and maxillary palpus, ventral abdominal prolegs on some segments and well-developed posterior spiracular disc.

Pupae with well-developed, open ended respiratory organ. Third leg curled under the wing sheath. Length of cephalothorax (measured from anterior margin to apex of wing sheath) about half length of remaining abdomen (posterior to apex of wing sheath). Two pointed, immovable anal processes, each with minute subapical seta.

Diagnosis (based on Chaoborus, the only genus in the Neotropical Region)

Small to moderate-sized midges (Fig. 1) wing length 1.0–2.3 mm and wing venation similar to that of Culicidae (R1 ending in C anterior to R2, R3 and R4+5; M with 2 branches) (Figs. 1, 4). Mouthparts short. Antenna with 13 flagellomeres, plumose in males.

Larvae highly distinctive, mostly transparent when alive (Fig. 3). Head capsule laterally compressed, with prehensile antennae, the bases of these abutting medially. With pair of prominent air sacs in both thorax and abdominal segment 7. Pupae with respiratory organ narrow apically, lacking obvious plastron (Fig. 2). Abdomen with pair of well-developed swimming paddles, each with two lateral and one medial rib supporting clear membrane. In living specimens, abdomen hanging straight down or slightly curved (unlike the tightly curled abdomen of Culicidae) (Fig. 2).

Diagnosis

Small to medium-sized biting midges (Fig. 1) wing length 0.4–7.0 mm. Ocelli absent. Antenna generally with 13 flagellomeres (some have fewer but always 6 or more present). Most males with plumose antennae. Most females with mandible serrate. Anepisternum divided into anterior and posterior portions by anepisternal cleft. Postnotum lacking medial longitudinal groove. Wing with 1–2 radial veins reaching anterior margin, usually with two median vein branches (posterior one may be weak) reaching wing margin. Wings overlapping each other and held flat over the abdomen in living individuals at rest (with exceptions in a few Stilobezzia). Fore- and midlegs shorter than hind leg. Tarsomere 1 of each leg equal to or longer than tarsomere 2 (with exceptions in some Forcipomyia).

Larvae with well-developed head capsule, well-developed pharyngeal complex, and no open spiracles. Pupae with well-developed, undivided respiratory organs with series of small spiracles. Third leg curled under wing sheath. Apex of the abdomen not curled under thorax. Two pointed or apically rounded anal processes.

Diagnosis

Small to medium-sized flies (Fig. 1), with wing length ranging from 4 to 7 mm. Adults slender, with elongate legs; color of most species dull brown or gray without conspicuous patterns, some species with silvery or opalescent pollinosity; mouthparts reduced or well developed, often sexually dimorphic; eyes dichoptic to subholoptic, often sexually dimorphic; dorsal ocelli present; antennae elongate, multisegmented; wing with netlike pattern of folds; legs long and thin; male terminalia dorsoflexed, female with three spermathecae, shape varied from spherical to oblong.

Larvae aquatic, cylindrical, ventrally flattened; body with six divisions, including cephalic division (fused head, thorax, and abdominal segment 1) and anal division (abdominal segments 7–10); each division usually with one lateral proleg per side and single ventral suctorial disc and paired gill tufts. Pupae aquatic, semiovoid, ventrally flattened and attached immovably to substrate; respiratory organs paired, transverse, and comprising four thin lamellae.

Diagnosis

Small to moderately large flies, wing length 2.0–15.0 mm. Thorax black, bright orange, or black and orange. Head sexually dimorphic; female dichoptic and eyes widely separated (Fig. 2); male holoptic (Figs. 1, 3). Antenna short, with 4–11 round compact flagellomeres. Antenna situated frontomedially above peristomal margin. Rostrum short or elongate. Clypeus + proboscis usually short, but sometimes longer than head. Wings hyaline to black or brown fumose; venation as in Figs. 8–12. Pterostigma often distinct (Figs. 10–12). Foretibia with either strongly developed apical spine (Fig. 4) or apical circlet of spines (Fig. 5). Males with hypopygium often dorsoflexed.

Larvae twelve-segmented, elongate, subcylindrical to slightly dorsoventrally flattened, cream to dark grey in color. Cuticle leathery, covered with microscopic sclerotized spicules and small to large fleshy tubercles (Figs. 6–7). Larvae holopneustic; posterior spiracle larger and more dorsal in position. Head capsule eucephalic and strongly sclerotized.

Diagnosis

Body length 2.3–8.8 mm.

Color yellow to brown or black, often patterned. Usually three ocelli, middle one small or rarely absent. Antenna with 14 flagellomeres, in regional Macrocerinae slender and one to five times body length in male, usually subequal to body length in female; in Keroplatinae (Keroplatini) flagellomeres compressed, often produced above and below, in regional species the lower projection at most twice as long as wide; in Keroplatinae (Orfeliini) usually cylindrical, if slightly compressed with projections not longer than wide. Palpus in Keroplatini porrect with palpifer and one or two palpal segments, the apical segment swollen; in Orfeliini and Macrocerinae with palpifer and four decumbent palpal segments. Antepronotum and postpronotum bare in Macrocerinae, setose in Keroplatinae; scutum uniformly setose or with bare longitudinal stripes; scutellum with posterior row of setae or with dorsal surface extensively setose. Scutellum sometimes with large membranous triangular basal area. Mediotergite bare. Pleural sclerites and prosternum variously haired or bare. Wing with Sc ending in C; R2+3 present, ending in R1 or C; base of M strong, weak or absent; Rs and M briefly fused at about one-third wing length. Foretibia with one apical spur, other tibiae with one or two spurs. Female cercus one-segmented in Keroplatini, two-segmented in Macrocerini.

Diagnosis

Small flies; mostly 1.0–6.0 mm in body length, rarely more than 10.0 mm, and of uniform structure (Fig. 1). Head usually ovoid or rounded, rarely with prolonged mouthparts. Compound eyes usually forming dorsal bridge in front of ocelli (Fig. 3). Antenna with rounded pedicel, shorter scape, 14 flagellomeres; flagellomeres with cylindrical basal parts (nodes), narrower necks; nodes clothed with setae about as long as width of node; short slender sensilla occur among setae. Some species with flagellomeres with long necks and long bristles, arranged in one or two roughly circular whorls (cecidomyiid-like, as in Fig. 13). Palpus usually three-segmented, in some genera with 1–2 segments. Color of most species uniformly brown or dark, rarely with yellow markings on scutum, pleural sclerites, abdominal segments, or base of antenna. Wing hyaline or brownish to blackish in larger species; rarely with dark spots or bands (Figs. 27, 31). Wing (Fig. 5) with thickened anterior veins (C, R1, R4+5), thinner medial and cubital veins, one crossvein r-m (with additional crossvein between R4+5 and M1 in Zygomma), C ending between apices of R4+5 and M1, CuA stem short, (with exception of Pseudosciara). Anterior wing veins with macrotrichia; posterior veins with or without macrotrichia. Wing membrane normally covered by microtrichia, in some genera also with macrotrichia (Figs. 24, 25, 30). Foretibia with one apical spur and modified bristle patch at tip (anteroapical depressed area); mid- and hind tibiae each usually with two subequal spurs. Tarsal claws toothed or untoothed. Abdomen cylindrical, male genitalia (hypopygium) exposed, with pair of gonocoxites, carrying articulated gonostyli (Fig. 4).

Larvae with eucephalous black head capsule and 12-segmented white and semitranslucent body. Last instar larva with one anterior and seven abdominal spiracles, abdominal segment 8 lacking spiracles.

Diagnosis

Adults small (wing length 1.5–7.5 mm), compact, humpbacked (Fig. 1). Head dichoptic in females, typically holoptic in males. Eye reddish, bluish in some species. Antenna cigarshaped, typically with 9 flagellomeres. Wing hyaline, unpatterned, with anterior veins thickened and concentrated along margin. Legs with or without bands. Claws of female of 3 types: toothless, with small subbasal tooth, or with thumblike basal lobe. Female terminalia with sternite 9 developed as slender genital fork; anal lobes (i.e., paraprocts) and hypogynial valves with diagnostic shapes and sclerotization patterns; spermatheca single, typically pigmented. Male genitalia with well-developed gonopods, each consisting of basal gonocoxite and distal gonostylus; aedeagus with ventral plate, median sclerite, parameres, and sometimes dorsal plate.

Larvae with head well-sclerotized, typically with head spots, and with pair of labral fans consisting of numerous rays endowed with microtrichia. Thorax with pair of respiratory organ histoblasts (dark in mature larva), and single ventromedial proleg. Abdomen elongate, with single terminal proleg bearing ring of minute hooks.

Pupae with respiratory organ arising from each anterolateral corner of thorax. Respiratory organs (“gills”) composed of various numbers and arrangements of slender filaments or inflated tubes; each organ with basolateral fenestra. Abdomen with various arrays of hooks and spines; segments III– VII or IV–VII with tergites and sternites separated by pleural membrane; sternites VI and VII typically divided medially by longitudinal, semi-membranous, striate area. Cocoon of three basic forms: shapeless, slipper-shaped, or boot-shaped.

Adult tanyderids are rarely collected, cryptic and believed to be short lived. Because of this, there is limited information about the natural history of Neotropical tanyderids. Of the three genera, Neoderus is the least studied, with only one account providing superficial information about its habitat (Madriz 2016). Knowledge of most aspects of its natural history is lacking partly due to sampling difficulties and uncertainty regarding the type locality. Much of the taxonomy and classification of Tanyderidae has been based on adult wing venation and colouration (Alexander 1927), with information on and use of terminalia being superficial or absent. Neoderus patagonicus (Alexander, 1913) was originally placed in Tanyderus. Alexander (1927) proposed Neoderus as a monotypic genus based on wing venation. Since then, no further information on the genus has been published, nor has any other attempt been made to recollect the species, which is confined to the fjords of Patagonia.

During a recent faunistic survey of the Ays en Region, we collected adult specimens of Neoderus and recorded detailed observations of the behaviour and habitat. The purpose of this paper is to present a diagnosis of adult Neoderus, redescription of N. patagonicus, and description of Neoderus chonos Madriz, sp. n. We also discuss evidence to clarify uncertainty about the published type locality of N. patagonicus.

Diagnosis

Adults small to medium-sized, about 1–13 mm long. Morphology diverse, some (especially marine) taxa with modifications or reductions to structures of the head, wings, legs, or terminalia. Ocelli absent; frontal tubercles present (e.g., Fig. 14) or absent. Compound eyes large, often with dorsomedial extensions (e.g., Fig. 14), but never contiguous. Antennae most commonly strongly sexually dimorphic; scape a flat ring; pedicel globose (especially enlarged in male); males with 6–15 flagellomeres, females with 4–7 or 10–14; male antennae usually with first and either last or penultimate flagellomere longer than others, and with flagellum densely plumose (Fig. 1), but exceptions occur in several genera. Mouthparts not piercing; female mandibles rarely present (no taxa with mandibles in Central America); proboscis rarely elongate; palpus usually elongate, with five (= four free) segments (Figs. 12–14), in various taxa reduced in length and/or segment number. Wing (Figs. 2–11) usually narrow (but see Fig. 5), broader in female (absent in some marine females); wings in resting position held to posterior, either flat (may be overlapping) or like sloping roof; costa reaching at most to tip of wing; wing margin usually reached by two to three branches of R, one of M, and two of Cu (but see Table 1 below). Legs (Figs. 30–49) usually elongate with forelegs longest, midlegs shortest; first tarsomeres longer than second; males at rest often hold forelegs aloft like ‘feelers’ (Fig. 1). Postnotum usually with medial longitudinal groove (Fig. 19). Abdomen usually much longer than wide, especially slender in males.

Larvae (Figs. 102–106) elongate, cylindrical (thoracic segments inflated in prepupae); with sclerotised, prognathous, usually non-retractable head capsule, three thoracic and nine abdominal segments, and usually anterior and posterior pair of prolegs bearing spines or claws; antennae various (long and retractable in Tanypodinae, Fig. 105); mouthparts directed anteriorly (“prognathous”), with mandibles operating in oblique to horizontal plane, and ventromedially with either transverse, usually toothed plate (hypostoma, Fig. 106; = mentum) or movable, toothed ligula (in larvae of Tanypodinae, Fig. 105); open spiracles usually absent (absent in all Central American species); terminal segments usually with paired procerci and four anal papillae (Fig. 102), sometimes also with short lateral and/or longer ventral tubuli (Fig. 103). Reductions occur, especially of prolegs and terminal appendages, for example in some non-aquatic larvae (e.g., Fig. 104).

Pupae (Figs. 99–101) ‘comma-shaped’ or relatively straight in lateral view, abdomen much narrower than cephalothorax, more or less flattened dorsoventrally; thorax anterolaterally often with pair of “thoracic horns” (sac-, tube-, trumpet- or tuftlike; simple, bilobed or with few to dozens of branches); antennal sheaths elongate, curving laterally over eyes; leg sheaths separate, usually at least tips of hind leg sheaths recurved under wing pads; tip of abdomen with or without apical pair of flattened anal lobes, the latter often with lateral fringe or distal setae or spines, not stiff and bare or with midrib. Various reductions from above occur, especially in non-aquatic pupae.

Diagnosis

Body length 3.0–8.0 mm (Fig. 1). Color yellow to dark brown with brown to black markings, shiny to dull. Antenna of Australosymmerus with 15 slightly to strongly compressed flagellomeres. Antenna of Rhipidita (Fig. 2) with 12–14 flagellomeres, those of males cylindrical or slightly flattened, produced below; processes narrowed below, triangular, shorter than length of flagellomere or slender, 1–6 times length of flagellomere (in the latter genus distal 1–3 flagellomeres lack ventral processes); flagellomeres of female short, cylindrical. Palpus with three segments in male, 1–3 in female. Postpronotum, propleuron, scutum near lateral margins, postsutural dorsocentral region, and scutellum with long erect setae; upper part of anepisternum with two or more shorter erect setae in Australosymmerus; mesopleuron and metathorax otherwise bare. Wing (Fig. 3) membrane with macrotrichia and microtrichia. Subcosta distinct only near base, tapering to free apex near base of Rs. R2+3 present, at least three-sevenths as long as R4+5, ending in C. Crossvein r-m in Australosymmerus present, subequal to basal section of Cu1, absent in Rhipidita due to brief fusion of Rs and M. Most tibial bristles of Australosymmerus at least twice as long as tibial diameter; those of Rhipidita not longer than tibial diameter. Australosymmerus with sternite of male normally sclerotized and with segments 8 and 9 projecting caudad; Rhipidita with sternite 7 membranous, segments 8 and 9 bent downward and forward (apparently a unique condition in nematocerous Diptera). Cercus of female with two segments.

Larvae with first eight abdominal segments each with pair of spiracles (at most first seven segments with pair of spiracles in other Sciaroidea; larvae of Lygistorrhinidae unknown).

Diagnosis

Adults slender (Fig. 1), 3–8 mm long (from anterior margin of clypeus to end of abdomen), 1–2 mm high (from upper margin of scutum to base of coxae). Head small, ovoid. Ocelli absent. Eye reniform, occupying most of side of head. Antenna with short, ringlike scape, enlarged globular pedicel, and 13-flagellomeres, usually more plumose in male. Proboscis long, slender, external part (labium) covered with scales. Thorax with patches of scales, as well as patches or rows of setae; setae usually coalesced on scutum into three paired, longitudinal rows: acrostical, dorsocentral, and supra- alar setae. Wing elongate, rounded apically, with scales along length of veins, microtrichia on membrane. Abdomen 10 segmented, segments 1–9 at least partly covered with scales in Culicinae bare in Anophelinae

Larvae (Fig. 50) somewhat dorsoventrally flattened. Head ovoid, square or broader than long. Antenna usually shorter to slightly longer than head, one segmented, arising from anterolateral angle of head. Larval eye present laterally on head in all instars, developing adult eye conspicuous only in instars 3–4 (i.e., last two instars). Labrum with elaboratelydeveloped pair of labral brushes, usually consisting of numerous closely-set hairs, for collecting food particles, or reduced in predatory forms to as few as 10 stiff hairs in Toxorhynchites. Mandible flattened, with prominent apicomedial tooth, well-developed comblike brush on anterolateral edge, whisk of long stiff hairs on inner surface for stuffing food particles into pharynx. Thorax stout, globular, of three fused segments, each with transverse row of branched setae, particularly well developed anteriorly, laterally. Abdomen 9 segmented, segment 8 with row or patch of pectinate teeth, called comb scales, sometimes accompanied by small sclerite. Posterior pair of spiracles on segment 8, flush with surface (in Anophelinae), or at apex of conical or tubular siphon (in Culicinae). Terminal segment partially or completely encircled by sclerotized band, or saddle, with longitudinal row of branched setae, called precratal setae, arising from ventral surface either on saddle or between its ventral edges. Anus, at apex of anal segment, surrounded by four conical or sausage-shaped anal papillae.

Pupae with head and thorax fused into cephalothorax, bearing one pair of respiratory organs anterodorsally. Mouthpart sheaths elongate, extending beyond base of legs. Abdomen eight segmented terminating in pair of articulated, broad, rounded, swimming paddles and genital sheaths.

Diagnosis

Small to medium-sized (body length 1–5 mm), nematocerous flies. Body covered by vestiture of hairs or scale-like setae (Figs. 1, 2). Antenna with 10–14 cylindrical, barrelshaped, pyriform or nodiform flagellomeres, with variouslyshaped translucent sensilla (ascoids). Eyes rounded (Figs. 3, 4), or reniform forming eye bridge above antennal insertions (Psychodinae) (Fig. 5, 27, 39). Males of some genera with sac-shaped structures on posterior of head (cornicula) (Fig. 13). Wings lanceolate (Phlebotominae; Fig. 7), elongate (Bruchomyiinae; Fig. 10), or oval (Trichomyiinae, Sycoracinae, Psychodinae; Figs. 6, 8, 9, 11, 12), with 9–12 longitudinal veins: Sc, R (4 or 5 veins), M (3 veins), CuA (2 veins), A (1 vein); r-m, the more constant crossvein, situated on basal third of wing; other basal crossveins may be present (Trichomyiinae, Bruchomyiinae, and Sycoracinae); with radial and medial forks; costa continuing around wing, broken just beyond base and usually with two additional breaks (Psychodinae); all veins densely haired. Male pro- and mesothorax of some genera with eversible or everted structures (patagia, tegulae) for pheromone release. Abdominal sternite 1 reduced (unsclerotized). Male genitalia inverted starting with segments 7 and 8, or only starting with segment 9 (Psychodinae); gonocoxite and gonostylus varying in form and ornamentations; aedeagus with two sclerotized lateral struts between which protrude two ejaculatory filaments (Phlebotominae) (Fig. 18) or complete or deeply divided, usually with parameres well developed and varied in form (in other subfamilies) (e. g. Figs. 30, 31). Epandrium with cerci (often called cercopods or surstyli) variously shaped, in Psychodinae with one to many stiff erect setae (tenacula, retinacula) (Fig. 32). Female genitalia with sternite 8 long and usually bilobed posteriorly, and sometimes infolded to form sclerotized plate on ventral side of genital chamber; with two weakly sclerotized spermathecae (Phlebotominae) (Fig. 22), one large spermatheca (Bruchomyiinae), two unsclerotized sac-like spermathecae (Trichomyiinae) or without spermathecae (Psychodinae).

Larvae

with head strongly sclerotized, non-retractile. Mouthparts well developed; mandible two-segmented, pharynx with filter apparatus in Psychodinae, one-segmented and without filter in remaining subfamilies. Thoracic and abdominal segments divided into pseudosegments (annuli). General habitus strongly differing among subfamilies. Phlebotomine larvae terrestrial, body elongate, thoracic segments divided into two, and abdominal segments 1–7 divided into three annuli. Abdominal segment 8 with elongate setae, segments 1–7 with ventral creeping welts. Antennae 3-segmented. Bruchomyiine larvae similar to those of Phlebotominae, but antennae 1-segmented, and abdominal segments without creeping welts. Trichomyiine larvae weakly sclerotized, segments not annulated, antenna 1-segmented. Larvae of Palearctic Sycoracinae aquatic, small, rounded, and

Classification

Blephariceridae currently are known from 27 genera and approximately 320 species, with representatives on most major continents (except Antarctica) and several continental islands (e.g., Madagascar, New Zealand, Sri Lanka). Despite this wide distribution, regional endemism is high. The subfamily Edwardsininae is restricted to the Southern Hemisphere — Australasian Region, southern South America, and Madagascar — whereas the only other subfamily, Blepharicerinae, occurs in both hemispheres.

The Neotropical fauna contains five described genera, including one Edwardsininae (Edwardsina Alexander) and four Blepharicerinae: Aposonalco, Kelloggina Williston, Limonicola Lutz, and Paltostoma.

Three of these genera have relatively restricted distributions: Edwardsina confined to the southern Andes (Chile and Argentina), Aposonalco endemic to central Mexico, and Kelloggina found mostly in Brazil. The remaining genera are relatively widespread, as follows: Limonicola widespread in northern South America and possibly extends into southern Central America, and Paltostoma occurs throughout Central America, Andean South America, and the Caribbean. Although common, Paltostoma is a poorly defined genus containing many species that do not fit easily in other genera (Hogue & Bedoya-Ortiz, 1989). Furthermore, the immature stages of Paltostoma show striking similarities with those of Limonicola. Unfortunately, adults of many species are known only from pharate specimens (dissected from pupae), and certain life stages remain undescribed for other species.

Taxonomía

Genus Erioptera Meigen

Genus Molophilus Curtis

Subgenus Molophilus Curtis

Genus Teucholabis Osten Sacken

Subgenus Teucholabis Osten Sacken

Genus Epiphragma Osten Sacken

Genus Hexatoma Latreille

Subgenus Eriocera Macquart

Genus Dicranomyia Stephens

Subgenus Dicranomyia Stephens

Genus Geranomyia Haliday

Genus Helius Lepeletier & Serville, 1828

Genus Neolimonia Alexander

Genus Rhipidia Meigen

Subgenus Rhipidia Meigen

Genus Toxorhina Loew

Subgenus Ceratocheilus Wesche

Suborder BRACHYCERA Macquart, 1834 (2 subordinate groups) Sequential

Clade CYCLORRHAPHA Brauer, 1863

(2 subordinate groups) Sequential

Comprende familias de moscas con adultos que poseen una lunula frontal y generalmente tambien una sutura frontal, que permite la salida del ptilinum al momento de la emergencia del adulto del pupario.

La vena r solo presenta tres ramificaciones: r1, r2+3 y r4+5. celdas m1 y m3 ausentes. la vena m3 esta ausente o se une a la vena m-m para formar la vena dm-cu que se une a la vena cua1 para formar la celda dm, denominada tambien “nueva celda discal” (dc), generalmente presente en todos los cyclorrhapha.

Empodia nunca son pulviliiformes. larvas: forman pupario con la exubia del ultimo estadio larval quedando dentro la pupa (coartata) y adultos emergen a traves de un abertura circular en el extremo cefalico del pupario.

Comprende las familias mas evolucionadas del orden diptera.

Presentan antenas cortas, generalmente trisegmentadas, provistas de una arista o de un estilo diferenciado. En algunas familias primitivas, el tercer artejo antenal presenta una anillacion mas o menos definida.

En las familias de este orden los palpos son cortos, uni o bisegmentados, prorrectos.

Alas con calipteres desarrollados en mayor o menor grado o asuentes.

La celda cup es mas angosta o cerrada hacia el margen del ala; en muchas especies es muy corta y en algunas esta ausente.

La celda discal (d) generalmente esta presente en las familias mas primitivas y es reeemplazada por la celda dm en las formas mas evolucionadas, solo en algunas familias esta ausente.

La vena rs en la mayoria de las familias presenta dos ramificaciones, r 2 + 3 y r 4+5; pero cuando es triramaficada, entonces es la vena r4+5 la que se ha bifurcado.

El cuerpo generalmente es mas robusto que en los nematocera y la sutura pleural del torax es bisinuada, angulada excepto en acroceridae.

Este suborden incluye 101 familias, que conformaban la serie brachycera del suborden orhtorrhapha y el sub orden cyclorrhapha de autores mas anitiguos.

Bruce et al. (1954) al tratar el suborden brachycera, agrupan a estas familias en dos divisiones, orthorrhapha y cyclorrhapha.

Infraorder ASCHIZA Becher, 1882 (2 subordinate groups, 7 familias)

Infraorder SCHIZOPHORA Becher, 1882 (2 subordinate groups)

Comprende aquellas familias de cyclorrhapha en que los adultos no poseen una sutura frontal.

La vena cua2 generalmente es larga y se extiende hasta el margen del ala o se une con la vena a1 cerca del apice de esta.

Comprende todas las familias de cyclorrhapha que presentan una sutura frontal a traves de la cual es empujado el ptilinum cuando los adultos emergen del pupario.

Antenas: trisegmentadas y casi siempre provistas de arista dorsal

La vena rs solo presenta dos ramificaciones.

Las familias incluidas en esta serie estan agrupadas en dos secciones: acalyptratae y calyptratae.

Clade Orthorrhapha Brauer, 1863 (7 subordinate groups)

Clade Archischiza

(1 family)

Clade Muscaria Enderlein, 1936

(2 subordinate groups)

Parvorder ACALYPTRATAE Macquart, 1835 (9 subordinate groups)

Comprende las familias más primitivas del suborden brachycera, las cuales NO presentan lunula frontal ni sutura frontal.

En algunas familias, las antenas presentan el flagelo anillado y aparenta constar de cinco o más segmentos, asemejandose en algo a los nematocera mientras que en otras el primer flagelomero no es anillado y termina en un estilo diferenciado o presenta una arista.

Pueden ser reconocidos por los palpos cortos, uni o bisegmentados, que se proyectan hacia delante, así como por la empodia que son pulvilliformes, setiformes o asuentes.

En la gran mayoria de familias, excepto empididae y dolichopodidae, la vena rs es triramificada por la bifurcacion de la vena r4+5.

La celda discal casi siempre está presente y la celda cup es más angosta hacia el margen del ala o está cerrada cerca de este.

Comprende aquellas familias de schizophora en que los calypteres siempre son pequeños o aun vestigiales.

Son moscas pequeñas. pedicelo de las antenas no presenta una sutura longitudinal en el aspecto dorsal, o cuando presente es incompleta. la vena sc frecuentemente es vestigial o incompleta, bien preservada en algunas familias.

La vena r1 es corta a muy corta. torax: sin sutura transversal completa en el dorso del mesoscutum.

Callo postalar generalmente ausente.

Los espiraculos abdominales

Generalmente estan ubicadas en las conjuntivas.

Superfamily Phoroidea Curtis, 1833 (5 families)

Superfamily Syrphoidea Latreille, 1802 (2 families)

Superfamily Tephritoidea Newman, 1834 (10 families)

Superfamily Tanypezoidea Rondani, 1856 (7 families)

Superfamily Sphaeroceroidea Macquart, 1835 (5 families)

Superfamily Opomyzoidea Fallén, 1820 (17 families)

Superfamily Nerioidea Westwood, 1840 (3 families)

Superfamily Sciomyzoidea Fallén, 1820 (11 families)

Superfamily Lauxanioidea Macquart, 1835 (3 families)

Superfamily Xylophagoidea Fallén, 1810 (5 families)

Superfamily Tabanoidea Latreille, 1802 (5 families)

Superfamily Carnoidea Newman, 1834 (7 families)

Superfamily Ephydroidea Zetterstedt, 1837 (7 families)

Superfamily Asiloidea Latreille, 1802 (11 families)

Superfamily Rhagionoidea Latreille, 1802 (6 families)

Superfamily Stratiomyoidea Latreille, 1802 (4 families)

Family Xylophagidae Fallén, 1810 (15 genera, 145 species)

Family Pelecorhynchidae Enderlein, 1922 (2 genera, 49 species)

Family Athericidae Nowicki, 1873 (12 genera, 133 species)

Family Stratiomyidae Latreille, 1802 (385 genera, 2,690 species)

Family Rhagionidae Latreille, 1892 (47 genera, 756 species,

Family Panthophthalmidae Bigot, 1886 (2 genera, 20 species)

Family Syrphidae Latreille, 1802

(209 genera, 6,107 species)

Family Pipunculidae Walker, 1834

(22 genera, 1,428 species)

Family Phoridae Curtis, 1833

(302 genera, 4,202 species)

Family Ephydridae Zetterstedt, 1837

(128 genera, 1,994 species)

Family Conopidae Latreille, 1802

(52 genera, 831 species)

Family Chloropidae Rondani, 1856

(194 genera, 2,885 species)

Family Drosophilidae Rondani, 1856

(76 genera, 4,017 species)

Family Cryptochetidae Brues & Melander, 1932

(3 genera, 34 species)

Family Agromyzidae Fallén, 1823

(41 genera, 3,017 species)

Family Lauxaniidae Macquart, 1835

(168 genera, 1,900 species)

Family Micropezidae Blanchard, 1840

(52 genera, 583 species)

Family Neriidae Westwood, 1840

(19 genera, 112 species)

Family Therevidae Newman, 1834 (128 genera, 1,143 species)

Family Nemestrinidae Griffith & Pidgeon, 1832 (26 genera, 300 species)

Family Mydidae Latreille, 1809 (66 genera, 498 species)

Family Asilidae Latreille, 1802 (555 genera, 7,531 species)

Family Tabanidae Latreille, 1802 (156 genera, 4,434 species)

Family Tephritidae Newman, 1834

(492 genera, 4,716 species)

Family Ulidiidae Macquart, 1835

(110 genera, 678 species,

Family Richardiidae Loew, 1868

(34 genera, 178 species,

Family Lonchaeidae Rondani, 1856

(10 genera, 504 species)

Family Tanypezidae Rondani, 1856

(5 genera, 68 species,

Family Diopsidae Bilberg, 1820

(14 genera, 194 species)

Family Psilidae Macquart, 1835

(13 genera, 322 species)

Family Anthomyzidae Czerny, 1903

(22 genera, 95 species)

Family Sphaeroceridae Macquart, 1835

(137 genera, 1,571 species)

Family Sciomyzidae Fallén, 1820

(66 genera, 618 species)

Nemestrinidae are distinctive flies that generally will not be misidentified. They are most likely to be confused with Bombyliidae that have a similar general appearance. Bombyliidae

have a setiform empodium and lack the “diagonal”

vein that is characteristic of nemestrinids.

Athericidae are easily recognized by the presence of the postspiracular scale on the metathorax, shared only with Tabanidae, from which they can be separated immediately by their modified antenna with slender stylus and their more delicate habitus. Athericids resemble Rhagionidae closely in general form, but rhagionids have no postspiracular scale and usually have a more tapered abdomen.

Syrphid flies are easily recognized by a combination of large basal cells (cells r, bm, and cup) with a closed apical cell (cell r4+5) (Fig. 51). A long spurious vein between the radial and medial sectors is a useful diagnostic character, but is not found in all species, and shorter spurious veins are found in some Conopidae.

Central American Mydidae are readily recognized by their general habitus, distinctive antenna with elongate antennal flagellum with apical club (except Rhaphiomidas), and characteristic wing venation with apical portions of veins R5 and M1 parallel to each other, M1 ending well anterior to wing apex. Mydids are most likely to be confused with Asilidae. However, asilids have different wing venation and mouthparts modified for predation. Mydids are also similar to Apiocera Westwood (Apioceridae), but Apiocera has a large, two-segmented palpus, and the genus is not known south of Mexico in Central America.

The wing venation of phorids separates them from most other families, although some species of primitive Diptera with long antennae, such as Simuliidae, Scatopsidae, and Cecidomyiidae, are superficially similar. Wingless females can be confused with some Sphaeroceridae, but lack a ptilinal suture and have the pedicel of the antenna largely hidden.

The family Tabanidae in Central America is most similar to the family Athericidae, but can be distinguished from athericids by the following features: antennal flagellum composed of 5–8 (rarely 4) flagellomeres forming a complex that is much longer than the pedicel (except in Silvius, subgenus Assipala) and without an arista-like stylus (Figs. 6–13); wing vein R2+3 ending well beyond R1 at the costa; veins R4 and R5 strongly divergent beyond the fork; fork originating well beyond distal margin of cell d (Fig. 5).

Rhagionids are recognized by their bulbous clypeus and may be separated from other families that have this character (Tabanidae and Athericidae) by their lack of a scalelike elevation immediately posterior to the posterior thoracic spiracle.

Pipunculids can be differentiated from syrphids by the large compound eyes that occupy most of their globose or semiglobose head, lack of spurious vein, and open r4+5 cell. A few bombyliids such as Metacosmus Coquillett resemble pipunculids when encountered in the field, but these roundheaded bombyliids can be easily differentiated by their antennae that lack the dorsal arista found in pipunculids.

Adults (Fig. 1) small to large flies (body length 2.0– 13.0 mm), mostly yellow to brown, sometimes black. Head with antenna porrect, pedicel usually elongate; arista bare to plumose; ocellar setae present in most species; inner and outer vertical setae developed; postocellar setae parallel or slightly divergent. Face slightly to strongly concave, uniformly sclerotized; vibrissa absent; clypeus small, withdrawn. Thorax with zero or one proepisternal seta; zero or one presutural supra- alar; zero, one, or two postalar; zero or one postpronotal; most with two notopleural; usually two pairs of scutellar setae. Wing elongate, completely hyaline (Fig. 7) to patterned (Figs. 6, 8); C and Sc complete; C unbroken; A1 + CuA2 reaching margin. Legs slender; some or all tibiae with preapical dorsal seta. Male terminalia symmetrical to asymmetrical; tergite 6 usually absent; right spiracles 6 and 7 mostly in tergite; epandrium well developed, symmetrical; usually with anterior and posterior surstylus, sometimes anterior one vestigial or absent; cercus well developed, setose. Female sternites 6, 7, and 8 separate or fused; two spermathecae present.

Medium-sized (body length 2–14 mm, most commonly

4–7 mm long) flies with moderately elongate body (Fig. 1);

some species that mimic ants with elongate or somewhat petiolate

abdomen, while some species associated with cacti with

body short and stout, resembling bugs (Heteroptera). Coloration

yellow to brown or black with pattern of microtrichose

spots or bands, often with metallic blue or green sheen. Head

variously shaped; inner and outer vertical, ocellar, divergent

postocellar and orbital (one or two, anterior one shortest)

setae present; frontal setae absent (except in Chaetopsis

and most Aciuroides). Frons sometimes pitted or wrinkled.

Clypeus moderately large. Proboscis with moderately developed

prementum. Thorax usually longer than broad. Wings

with a pattern of dark spots or bands, rarely completely hyaline;

Sc vein complete, C with distinct humeral break, subcostal

break indistinct. Legs moderately developed, usually

without rows of spines or strong setae; tibiae without dorsal

setulae or setae. Females, like in all other Tephritoidea, with

telescopic ovipositor, whereas males with long phallus, hidden

in membranous pocket on ventral side of abdomen and

bearing no glans at apex.

Although Stratiomyidae are one of the most morphologically diverse groups of Diptera, they are easy to recognize. Their distinctive wing venation, with the radial veins crowded anteriorly and the small discal cell, are characteristic. They could possibly be confused with Xylomyidae, but this family has two tibial spurs on the midleg and one or two on the hind leg. The nematocerous family Anisopodidae has wing venation somewhat similar to stratiomyids, but these flies have more than two palpal segments and cell cup is broadly open at the wing margin. Syrphids superficially resemble some stratiomyids, but syrphid wing venation is different in having the venation less crowded toward the costal margin and having a much larger cell dm.

Xylophagidae are easily differentiated from other lower Brachycera with a pulvilliform empodium by their relatively unmodified wing venation and lack of a scalelike elevation behind the metathoracic spiracle. They key out near Vermileonidae, but Neotropical xylophagids do not have the apical part of the flagellum modified into a stylus. Xylophagids most closely resemble Xylomyidae in general appearance, but xylomyids lack an apical spur on the foretibia. Rachicerus, the only known genus of xylophagids from Central America, is the only genus of lower Brachycera with a pectinate antennal flagellum with more than eight flagellomeres (Figs. 5–6).

Conopid flies are easily recognized among higher flies by a combination of characters: ptilinal suture present, lack of distinct bristles (except Stylogaster), cell r4+5 frequently petiolate; cell cup usually large (small in Stylogaster), often with short spurious vein between radial and medial fields, antenna and proboscis elongate, usually greatly so. The only group that conopids may be confused with are syrphids, but the well-developed ptilinal suture will separate these two families. Many conopids and syrphids are sorted as hymenopterans, but obviously differ in the number of wings and other characters.

Moderately small (body length 3.0–6.0 mm), stout, relatively setose, shiny black to metallic blue or green flies, with wings usually hyaline, sometimes partly yellowish or fumose (Fig. 1).

Body stout, abdomen broad and flat. Frons narrower in male than in female; setulose, with one reclinate orbital seta.

Ocellar and inner and outer vertical setae strong (Fig. 2). Postocellar setae divergent, weak. Without vibrissa, but one or more subvibrissal setae sometimes vibrissa-like. Lunule large, exposed, bare or setulose. Face broad, depressed, usually without strong carina.

Eye large, pilose or bare. First flagellomere short and porrect to long and decumbent; arista bare, pubescent or plumose. Thorax with one postpronotal, two notopleural, one presutural, and one postsutural supraalar, two postsutural dorsocentral, two prescutellar acrostichal, often one weak intra-alar, and two postalar setae. Scutum strongly arched, setulose. Scutellum with two large setae and sometimes additional setulae on margin or disk. Proepisternum with one seta, proepimeron with one to many setulae. Anepisternum with numerous setae, at least posteriorly, often smaller anteriorly. Katepisternum with one to three dorsal setae and numerous setulae. Anepimeron usually bare, sometimes with one to several setulae medially. Meron bare. Legs mostly dark, tarsi sometimes partially or entirely yellow. Wing (Figs. 22–23) somewhat triangular, tapered, with anal lobe and alula well developed. Venation with crossveins bm-cu, r-m, and dm-cu and vein CuA2 present. Costa extended to vein M, with costagial and humeral weakenings and subcostal break. Subcosta complete, apex free from R1. Vein R1 nonsetulose dorsally. Crossvein r-m near or proximal to midlength of cell dm. Vein A1+CuA2 sometimes reaching wing margin as fold. Dorsal calypter well developed, with whitish to brownish fringe of hairs, those at calyptral fold sometimes longer and darker than others (Neosilba, some Dasiops). Halter dark brown. Abdomen broad and flat. Male with epandrium and hypandrium symmetrical. Phallus short and unsegmented (Dasiops; Figs. 14–15); or elongate, bisegmented, and ornamented (other genera; Figs. 16–19). Surstylus usually with short, stout prensisetae (Figs. 14–15). Pregonite (=gonopod of McAlpine, 1987) with base fused to hypandrium, with tiny setula on medial surface near apex; postgonite reduced and semimembranous to strongly developed and toothed. Female with sclerotized telescopic ovipositor (Figs. 20–21). Seventh tergite and sternite fused to form oviscape. Eighth tergite and sternite comprising four elongate rods, closely associated with or occasionally fused

to usually oval or pointed apical sclerite formed by fused epiproct, hypoproct, and cerci. Three spermathecae present (McAlpine, 1987).

Larvae (Fig. 24) typical muscomorph maggots, relatively long and slender, cylindrical, tapered anteriorly, and with spiculose intersegmental rings and creeping welts. Mandible without secondary teeth,

Medium-sized to large flies (body length 5–15 mm), with long, slender legs (Fig. 1).

Head usually longer than high. Frons with one to five fronto-orbital bristles, ocellar bristles absent, vertical bristles sometimes reduced, vibrissa present in some species. Upper face deeply divided medially and forming conspicuous antennal bases in Neriinae (Fig. 2: ant bas). Antennae porrect; pedicel with elongate to triangular projection on inner side; scape and/or pedicel elongated in some species; arista apical to dorsoapical, bare or pubescent. Thorax with relatively short and stout bristles in Central American species; katatergite protuberant. Foreleg with elongated coxa, at least forefemur usually with stout, spinelike ventral bristles, mid- and hind tibia with ventroapical bristle, foretibia with rows of spine- or tubercle-like bristles in males of some species. Wing elongate, with R4+5 and M strongly convergent; subcostal break usually absent, additional crossveins present in one Central American species. Male terminalia elongate, flexed below abdomen. Female abdominal segment 7 developed as conspicuous oviscape.

Neriidae can be distinguished from other long-legged flies with apically narrowed wing cell r4+5 (Micropezidae and Tanypezidae) by the porrect antennae with dorsoapical or apical arista, the apicomedial projection of the pedicel, the peculiar development of the upper face, and the presence of ventral femoral spines (at least on forefemur).

Greatly varied in size and color, Neotropical species ranging from just slightly more than 1 mm in body length (some Cladochaeta, Diathoneura) to 6–7 mm (some Drosophila); from yellow to brown, black, mottled, and banded, or metallic colors. Eyes often red, but can be brownish or gray. Antennal arista usually plumose (Figs. 1, 2, 3, 9, 10), sometimes with just ventral branches lacking, occasionally with all branches of arista lacking. Frons with three pairs of fronto-orbital setae: the anterior-most pair proclinate, then two pairs ofreclinate setae, referred to as anterior and posterior reclinates (Figs. 4–8). Anterior reclinates often small, sometimes reduced or lost (Figs. 2, 3). Drosophilidae are superficially similar to the smaller, yellowish Lauxaniidae, some of which have a plumose arista (but usually with all fronto-orbital setae being reclinate). The inside surface of the forefemur of Drosophilidae lacks ctenidial spines, which many species in the closely related families of Ephydroidea, such as Camillidae and Curtonotidae, possess. Ephydridae commonly have a plumose arista, but almost always just with dorsal rays (and they further have a bulging face).

Medium to large (body length 5–17 mm), distinctively long-legged flies (Fig. 1).

Wing veins R4+5 and M1+2 convergent towards wing tip. Ocellar bristles and vibrissae absent. Most Neotropical species with distinctive vertical, fanlike row of bristles on katepisternum. Females with tergite and sternite 7 fused to form rigid oviscape housing ovipositor. Males of Micropezinae and Eurybatinae with surstyli; Taeniapterinae without. Males of most species with prominent forked process on sternite 5.

Larvae of relatively few species known, but known species of genera that occur in the Neotropical Region with two dorsally pointed posterior respiratory processes bearing spiracular discs, each with three or four slits (numerous pores in one Nearctic species, and spiracles vestigial in one Australian species). Mandibles of known Neotropical species lack teeth (the mandible is ventrally toothed in at least one Australian species). Anterior spiracles usually fan-shaped, but sometimes reduced or divided into two lobes. Larvae of genera that occur in Central America were described by Albuquerque (1972), Steyskal (1964), Müller (1957), and Sabrosky (1942).

Neriidae, the long-legged flies most similar to Micropezidae, have an apical arista in contrast with the dorsobasal arista of Micropezidae. Neriids can also usually be distinguished from micropezids by the presence of vibrissae and spinelike femoral bristles. Tanypezidae are superficially similar to micropezids, but are easily distinguished by the presence of strong anepisternal bristles and the absence of katepisternal bristles.

Minute to medium-sized flies (wing length 0.9–6.5 mm, usually 2–3 mm).

Often yellow and/or black, brown, or gray, few have some metallic greenish, bluish, or coppery coloration (genus Japanagromyza and Melanagromyza). Head (Figs. 1–6) with first flagellomere small, round, or sometimes enlarged, elongate, or with anterodorsal projection or spine and with well-developed dorsobasal arista. Fronto-orbital and ocellar bristles strong; postocellar bristles divergent; fronto-orbital setulae in one or more rows, usually reclinate (Fig. 2), but sometimes proclinate (Fig. 3), upright or absent; vibrissa well developed, some males (genus Ophiomyia) with vibrissal fasciculus (Fig. 4). Scutum with two to five dorsocentral bristles present, but many genera in Phytomyzinae with three postsutural and one presutural dorsocentral bristles; acrostichals present (in up to 10 rows) or absent. Costal break present near the end of R1, humeral break absent; Sc incomplete distally (Fig. 9), or reaching C as linear fold, independently of R1 (Fig. 10), or Sc complete and fusing with R1 before reaching costa (Fig. 11); CuA2 and A1 present, forming small cell cup (Fig. 11); A1 not reaching wing margin. Abdomen tapered, composed of six visible segments. Females with abdominal segment 7 modified into large conical nonretractile oviscape (Fig. 12).

Some female Tephritidae have an oviscape similar to that in Agromyzidae, but their subcostal vein is abruptly bent forward at an angle of 90˚ (not so in Agromyzidae), and they frequently have patterned wings (rare in Agromyzidae). Chloropidae usually have much reduced chaetotaxy on the head and thorax, and wing vein A1 and cell cup are absent. In Clusiidae, the subcostal vein is more pronounced and complete, the subcostal break is sometimes absent, and many species have a triangular extension on the outer side of the pedicel. Members of the family Odiniidae have the anepisternum bare (always with some setae in Agromyzidae).

Adults minute to medium-sized flies (body length 0.6–11.0 mm), usually dull and dark colored (Fig. 1), often invested with whitish, grayish, or brownish microtomentum; but unusually diverse in body structure, vestiture, and ornamentation; female usually larger than male. Head, including subcranial cavity, large; one to three, rarely up to five lateroclinate or reclinate/proclinate fronto-orbital setae; pseudopostocellar setae divergent or lacking. Arista pectinate on dorsum only or bare to pubescent; face flat to conspicuously protruding. Scutum usually bearing numerous acrostichal setulae, one to five dorsocentral setae, two to three supra-alar setae, one postalar seta; anepisternum setulose. Wing usually well developed, hyaline but sometimes maculate; C with humeral and subcostal breaks; Sc incomplete; vein R1 merging with C before middle of wing; cells bm and dm confluent, crossvein bm-cu absent; cell cup absent. Preapical dorsal seta lacking on fore- and hind tibiae. Males commonly with five exposed tergites; female with six to seven; abdominal spiracles 1–5 located in tergites, 6th in pleural membrane near genital capsule. Male terminalia (Fig. 58) symmetrical. Female usually with two spermathecae, rudimentary and probably nonfunctional; ventral receptacle (Fig. 59) heavily sclerotized, functioning in place of spermathecae. Immature stages mostly aquatic or associated with aquatic habitats.

Small to moderately large (body length 2–11 mm) flies; compact bodied (Fig. 1) or elongate; varied in color patterns, often with marks, spots, bands, or reticulating patterns.

HEAD with postocellar setae convergent to cruciate, rarely absent; ocellar setae proclinate, divergent to parallel, small and hairlike to strong; orbital plate with two fronto-orbital setae (rarely zero to one), usually reclinate, but anterior seta sometimes slightly (Fig. 23) to strongly (Fig. 27) inclinate; lunule small, narrowly exposed and bare; face various, from slightly concave or flat to convex or bulging; subgena usually thin and inconspicuous, but occasionally enlarged and bulging; vibrissa absent. Scutum setulose, not hairy, occasionally nearly bare; with two to four dorsocentral setae (rarely only one), either posteriorly located (0+2, 0+3) (Figs. 33, 34) or evenly distributed with one presutural (1+3, 1+2) (Fig. 35); prescutellar acrostichal seta usually present, sometimes with additional acrostichal setae in row (Fig. 35); scutellum with two pairs of setae; anepisternum setulose, with one strongseta near posterior margin; katepisternum setulose, with one to two setae dorsally. Wing with C unbroken, weakened beyond R4+5 and ending at M1, with stout black spinules on anterodorsal surface ending slightly beyond apex of R2+3 (Fig. 44), or continued to R4+5 in Homoneurinae (Fig. 43), although fine hairs often continue around wing beyond spinules; Sc complete, separated from R1 (Figs. 43–52); cell cup small with rounded apex; A1 usually short (long in Eurychoromyiinae), not reaching wing margin; membrane various, from hyaline to darkened crossveins to intricately patterned, irrorated, or banded. Legs each with preapical dorsal tibial seta (except in some Eurychoromyiinae). Abdomen short and stout to elongate and tapering; segment 7 in female usually normal (Fig. 55), but occasionally forming complete sclerotized ring; sternite 8 often variously modified (e.g., Figs. 53, 54). Some genera display sharp reductions or losses of setae.

Minute to small (body length 1.0–5.0 mm, rarely larger) flies (Fig. 1). Shining black to dull gray or yellow, often with patterns of black and yellow. Antenna with first flagellomere usually round, arista dorsal, bare or pubescent. Cephalic chaetotaxy reduced: ocellar, postocellar, inner, and outer vertical setae usually present, fronto-orbital setae usually reduced, only slightly longer than surrounding setulae, occasionally with one to two longer setae present, frontal setulae usually uniformly distributed, except on surface of frontal triangle; distinct frontal triangle usually present, occupying half or more of frons; thoracic chaetotaxy usually reduced (less so in Apotropina), one to two postpronotal, one anterior, one to two posterior notopleural, one to two postsutural dorsocentral setae present, acrostichal setulae in several rows, two to three pairs of scutellar setae present; propleuron with complete, sharp vertical carina, prothoracic spiracle round, wing with subcostal break (secondarily lost in some species of Pseudogaurax), sc incomplete, cell cup and CuA2 absent, kink in CuA1 usually present basal to dm-cu. Legs usually without outstanding bristles, but apicoventral hind tibial spur (actually a spurlike seta) present in some genera; hind tibia often with elongate or oval tibial organ. Male postabdomen symmetrical, genitalia simple. Female abdomen telescoping, cerci long, spermathecae unsclerotized.

The combination of wing venation (missing cup and CuA2, kink in CuA1), vertical propleural carina, reduced frontoorbital setae (in most species), and prominent frontal triangle (in most species) distinguishes chloropids from other acalyptrate flies. Some shiny black species of Milichiidae and some yellow and black Agromyzidae and Clusiidae are externally similar but can be distinguished by the number and orientation of fronto-orbital bristles and by differences in wing venation. Ephydridae with similar wing venation generally have a projecting face and longer fronto-orbital bristles.

Diagnosis

Bristleless, slender flies approximately 4–12 mm in length (Fig. 1). Body color generally drab: gray, brown or black, frequently with some yellow or orange. Some with distinctive, silvery, golden, green, or blue hairs on thorax. Eyes of male holoptic or narrowly separated, eyes of female widely separated. Antenna with small scape and pedicel and first flagellomere round or conical, bearing elongate terminal arista-like stylus (Fig. 2). Adult with convex, bulbous clypeus. Labella fleshy; palpus one-segmented (extralimital taxa may have palpus two-segmented). Legs elongate and delicate; tibial spurs 0:2:2 or 0:2:1. Wings patterned or hyaline (Fig. 3). Abdomen tapered posteriorly; female ovipositor telescopic, cercus two-segmented (Fig. 4); three spermathecae spherical or elongate, with or without sclerotization. Male terminalia with simple epandrium; epandrial sclerite wider than long; subepandrial sclerite present or absent; hypandrium fused entirely to gonocoxites (Chrysopilus) or hypandrium separated from gonocoxites by complete or incomplete suture (Rhagio, Undescribed Genus A); gonostylus simple; aedeagal tines absent (Fig. 5).

The Cryptochetidae are a small family of tiny flies (generally 2 to 4 mm long).

Some twenty to thirty species are known. Generally they are metallic blue black, stoutly built, with the head broad and high and with clear wings.[1] Like other species in the superfamily Lonchaeoidea, the Cryptochetidae have antennae with a cleft in the second segment. Unlike practically all Schizophora however, they lack an arista, or if they do have one, it is too small to distinguish with any confidence. The family name refers to this unusual distinction; "Cryptochetidae" literally means "those with hidden bristles". The adult flies also are unusual among insects in that they have only a single pair of abdominal spiracles — this is not a serious physiological challenge in such small insects.

Again in resemblance to other Lonchaeoidea, the Cryptochetidae do not have more than one proclinate orbital bristle on each side. The frons is densely setulose. The costa has a break at the end of the subcosta. The sixth abdominal sternite in the male is symmetrical and it has an 8th tergite. In the female the seventh tergite and sternite are fused, and the eighth segment is elongated.

The larvae are of biological and economic interest, being endoparasitoids of coccids. In Cryptochetum iceryae, which parasitizes Icerya, there are four larval instars. The first instar is sac-like and lacks both trophi and tracheae but at the caudal end it bears a pair of finger-like processes. The caudal end of the digestive tract is closed. During subsequent instars the caudal processes grow longer and become filamentous; in the final instar they are much longer than the whole body. These filaments probably are respiratory organs. Only a few species of Cryptochetidae have been described, and most of those occur in tropical countries. At one time they were allocated to the Agromyzidae but now are regarded as a separate family.[2]

Diagnosis

Medium sized flies (body length 7–10 mm), often with patterned wings (Fig. 1). Eyes nearly to fully holoptic in males, widely separated in females. Antenna with scape and pedicel short, flagellum with first flagellomere small, ovoid or kidney shaped, with long, arista-like stylus often inserted dorsally. Clypeus convex, bulbous (Figs. 2, 3). Mouthparts sometimes elongate, adapted for blood-feeding, otherwise short, labella fleshy; palpus two-segmented, basal segment small, ovoid, apical segment elongate. Thorax unmodified, scutellum evenly rounded apically. Posterior spiracle with scalelike elevation immediately behind it. Legs simple, sometimes elongate and delicate; tibial spurs 0–2–2, occasionally 1–2–2. Wings moderately long, often patterned. Vein R2+3 ending in C very near or at end of R1; cell cup elongate, closed slightly before wing margin. Abdomen about as wide as thorax, only slightly tapered posteriorly, apex somewhat blunt. Larvae with distinctive morphology featuring abdominal prolegs and other fleshy processes (Fig. 4).

Diagnosis

Medium-sized flies (body length 4–16 mm; larger in non-Central American forms), body variously colored, short and compact, often with conspicuous, dense pilosity; wings elongate, ranging from hyaline to dark brown (Fig. 1). Eyes holoptic to separate in males, usually separate in females; ranging from bare to densely pilose. Antennae small to minute, scape longer than short pedicel; flagellum with first flagellomere short, conical, followed by arista-like stylus composed of three flagellomeres. Face flat to moderately convex, sometimes with grooves near eye margin for reception of palpi (Fig. 2). Mouthparts varied, ranging from vestigial and difficult to observe, to short with labella fleshy, to elongate with reduced labella; palpus usually twosegmented, slender, sometimes reflexed dorsally and housed in groove at margin of face, or occasionally vestigial. Thorax unmodified; scutellum short, evenly rounded apically, often with submarginal groove. Legs simple; tibiae without apical spurs; tarsi with empodium pulvilliform. Wings usually longer than body. Venation complex, with so-called “diagonal” vein formed of variously fused veins (elements of Rs, R4+5, r-m, M1, M2, M3, CuA1) (Fig. 3); cell cup elongate, open at wing margin in Central American species. Abdomen about as wide as thorax, short and rounded to only slightly elongate.Female cerci one-segmented, short to elongate, forming ovipositor (Fig. 1). Female with only two spermathecae.

Larvae with later instars grublike (Fig. 4).

Diagnosis

Medium-sized flies (body length 5–12 mm) in Central America, wings clear to infuscated, without obvious pattern (Fig. 1). Eyes widely separated in both sexes. Antenna with scape and pedicel usually short; flagellum composed of 7–8 or more than 15 flagellomeres (Figs. 5–6), usually at least slightly pectinate ventrally. Clypeus more or less flat, not bulbous. Mouthparts unmodified; palpus two-segmented, basal segment small, apical segment expanded apically. Thorax unmodified, scutellum evenly rounded apically. Legs simple; tibial spurs 1–2–2. Wings (Figs. 3–4) moderately long, anal region sometimes reduced. Vein R2+3 ending in C near end of R1; alula reduced or essentially absent. Abdomen about as wide as thorax or slightly narrower, only slightly tapered posteriorly in males, females with apical segments moderately elongate, strongly narrowed posteriorly. Larvae with distinctive morphology featuring strongly sclerotized, cone-shaped head capsule and last abdominal segment with posterodorsal sclerotized plate bearing pair of processes (Fig. 2).

Diagnosis

Relatively large to huge flies (body length 11–60 mm), elongate in general form (Fig. 1), color various, often wasplike in general appearance. Eyes separate in both sexes, bare. Vertex flat to usually sunken; with three ocelli in primitive taxa, but in most species ocellar triangle highly modified with lateral ocelli nearly linear, appearing absent. Antenna with scape more than twice length of pedicel (scape shorter in Rhaphiomidas); flagellum slightly to greatly elongate, with basal stalk and apical thickened club, but flagellomeres not well delineated (Figs. 2, 3) (in Rhaphiomidas flagellum does not have stalklike base and is spindle shaped; Fig. 4). Mouthparts various, short to elongate, labellum fleshy but occasionally reduced; palpus one segmented, small to greatly reduced and difficult to see. Thorax unmodified, scutellum usually small, rectangular; marginal areas of scutum and scutellum without macrosetae (except in Rhaphiomidas). Legs simple, usually with some macrosetae; hind femur sometimes thickened, with short, stout toothlike ventral setae; tibiae without spurs, hind tibia often slightly arcuate, with ventral carina prolonged into toothlike process at apex of tibia (Fig. 5). Wings various, transparent to strongly pigmented, lacking microtrichia. Branches of M peculiarly curved forward, more or less parallel to posterior wing margin, apical portions of veins R5 and M1 parallel to each other with M1 ending well before wing apex; M2 usually fused apically with M1 or CuA1 (Figs. 8, 9), but when free (Rhaphiomidas), ending anterior to wing tip (Fig. 10). Abdomen elongate, stout, cylindrical to slightly tapered, usually more flattened and larger in female; female with blunt apex, often with conspicuous acanthophorites with spines (Fig. 7). Larva (Fig. 11) elongate, external portion of head capsule sclerotized, posterior spiracle at anterior margin of penultimate abdominal segment.

Diagnosis

Small to large flies (body length 2–28 mm), highly varied in shape and coloration, wings hyaline to variously infuscated or patterned. Some species mimic wasps and bees, particularly in flight. Eyes various, males often holoptic (Fig. 5). Antenna with scape and pedicel usually short, flagellum with 5–8 flagellomeres, highly varied in shape (Figs. 7–21), sometimes with apical stylus or arista-like stylus (e.g., Figs. 13, 18). Face usually flat or evenly rounded, occasionally tuberculate or conically produced (Figs. 2, 3). Mouthparts typically short with fleshy labellum (Fig. 4), lateral area of labellum sometimes sclerotized, occasionally narrowed (Fig. 3). Palpus usually two-segmented, occasionally one-segmented, sometimes more or less concealed in subcranial cavity. Thorax unmodified, scutellum short, unarmed, with pair of apical spines of various size (Figs. 28, 30–32), or less commonly with 4 or more marginal spines (Fig. 29). Legs usually simple, occasionally hind legs elongate; fore- and hind tibiae without apical spurs (Fig. 26), midtibia usually without spurs but occasionally with one tiny spur (Fig. 25). Wings well developed, hyaline, infuscated, or occasionally with distinct pattern (Figs. 36–44); radial veins characteristically crowded toward anterior margin; R5 or R4+5 ending well before wing apex; cell m3 open; M3 usually present and long but occasionally reduced or absent (Figs. 36–37, 44); discal cell characteristically reduced in size and usually in anterior half of wing; cell cup elongate, closed. Abdomen highly varied in shape, ranging from short and rounded to elongate (Figs. 22–23, 27), sometimes clavate (Fig. 24). Male with phallic complex trifid in the vast majority of Central American taxa, occasionally bifid, rarely simple and tubular. Female cercus usually two-segmented, occasionally one-segmented, not strongly modified. Female almost always with 3 spermathecae, rarely with only 2, ducts moderately elongate and usually with distinct hingelike bend (e.g., Ururahy-Rodrigues & Pujol-Luz, 2000: figs. 2, 5). Larvae with distinctive morphology featuring shagreened appearance with cuticular warts (Figs. 45–46). Larval pupation within last larval skin.

Diagnosis

Slender to moderately thick-bodied (Fig. 1); body length, excluding antennae, 3–15 mm. Background color light yellow to black, wholly or partly covered with pubescence, and often including bare regions displaying ground color (e.g., Fig. 10). Macrosetae usually prominent, black to pale. Eyes usually holoptic in males (Fig. 8), dichoptic in females (Figs. 7, 9). Wing venation constant (Figs. 16–17): R4 elongate, sinuous, divergent from R5, with cell r4 encompassing the wing apex; cell bm truncate distally and with four corners from which arise four separate veins (M1+2, M3, CuA1, and CuA2); discal (d) cell elongate, with three veins extending from apex (M1, M2, M3); crossvein m-cu present, so base of cell m3 truncate; cell br extends beyond apex of cell bm; r-m attached to basal half of discal cell; cell cup closed near wing margin. Abdomen with 8 well developed pregenital segments, often with fine silvery or bronzy pubescence covering dorsum of male, often reduced or lacking to display ground color in female. Female terminalia with large conspicuous sternite 8 (Figs. 18–19); acanthophorite macrosetae on tergite 10 absent (Fig. 18) or present posterodorsally and anterolaterally (Fig. 19). Internally, female with 2 (in Therevinae and Xestomyzinae) or 3 (in the remaining therevids) thinly sclerotized spermathecae and a central spermathecal sac (lacking in Phycinae).

Larvae long and slender, with posterior spiracles situated laterally on apparent fourth segment from end (Fig. 20, right); head capsule is complete and permanently exerted (Fig. 20, left) with flexible, spatulate, posteriorly articulated metacephalic rod. Antennae minute and peglike (Fig. 21). Pair of heavy, elongate tentorial arms present within head capsule (Fig. 21). Each thoracic segment bears pair of lateral setae (Fig. 20, left). Terminal abdominal segment bears pair of retractable prolegs (Fig. 20, right).

Pupae (Fig. 22) with head segment bearing pair of antennal sheaths, median labral sheath and proboscial sheath ventrally. Thorax bearing pair of wing sheaths, each with spinelike wing process. Leg sheaths and wing sheaths visible ventrally. Each abdominal segment bearing ring of spines in posterior half, and abdomen terminates in pair of caudal spines.

Diagnosis

Adults small to large flies (body length 3–60 mm for all species, up to 40 mm long in Central America); shape long and slender to robust and beelike; of nearly any color (though rarely green), but most often gray or dark brown; body often bristly, sometimes nearly bare or densely long-pilose. Eyes large and bulging, separated dorsally by depressed vertex; sometimes closely convergent above, but never holoptic; rarely vertex nearly level with eye margin. Face with row or clump of bristles (“mystax”)—may be limited to subcranial margin or cover entire face. Mouthparts adapted for piercing, with labium formed into rigid proboscis, enclosing knifelike hypopharynx; mandibles absent; maxillary palpus one- or two-segmented. First flagellomere of antenna varied in shape, although usually elongate (may be several times longer than scape and pedicel together), with one to two additional apical flagellomeres; second flagellomere often styluslike (rod-shaped, pointed) or arista-like (slender, bristlelike), sometimes with broad apex, which may have apical concavity, sometimes short and collarlike; third flagellomere, if evident, usually either small and spinelike, or elongate and stylus- or arista-like; apical flagellomeres sometimes apparently absent or fused with first flagellomere (latter structure may then have apical or dorsal pit with minute, inset, spinelike peg; second flagellomere together with reduced third are often simply termed “stylus” in key to the genera. Wings present, though sometimes narrowed at base (reduced to long-stemmed paddle in one genus) and adults capable of flight; venation may be unmodified, with all peripheral cells open, or cells r1, r5 and m3 may be closed (cell cup is usually closed); R vein always four-branched, may have spur vein near base of R4 (which may be connected to R2+3 or not); sometimes one or more M veins shortened or absent, resulting in reduced number of cells posteriorly. Legs relatively stout, with anterior and middle pair provided with long spinelike bristles adapted for catching and holding prey; hind legs varied; pulvilli and empodium sometimes reduced in size or absent. Unusual internal characteristics include presence of large venom glands in thorax of all species, as far as is known; absence of crop (oesophageal diverticulum) in some groups. Spermathecae varied in shape, but usually three present; only two present in some, or median one highly modified.

Diagnosis

Small to large (6–25 mm) stout-bodied flies (Fig. 1). Eyes holoptic in male, separated by frons in female. Frons often bearing one or more calli (Fig. 4). Antennal flagellum with larger first flagellomere and 4–8 apical flagellomeres. Subcallus usually inconspicuous but sometimes protuberant, bare and shiny. Maxillary palpus two-segmented (Figs. 18, 19), apical segment long and curved ventrally in female, shorter and more rotund in male. Mandible and maxilla of female styliform and adapted for piercing in most species. Thorax with notopleural lobe prominent. Legs with paired apical spurs on midtibia, present or absent on hind tibia. Tarsi with pulvilliform empodium. Wing venation consistent throughout family (Fig. 5). Veins R4 and R5 enclose wing apex. Basal radial (br) and basal median (bm) cells and discal (d) cell large (Fig. 6), cell cup usually closed near wing margin. Wing membrane often with variety of distinctive patterns (Figs. 21–30). Terminalia of male with gonocoxite fused with hypandrium, and with gonostylus single or partially divided. Aedeagus and associated pair of slender filamentous recurved aedeagal tines enclosed in sheath derived from parameres. Epandrium entire or divided, tergite 10 absent. Cercus flattened. Female usually with tergite 10 divided, sternite 8 shield shaped and cercus one-segmented.

Families superficially similar to lauxaniids (e.g., some Drosophilidae) can be readily separated from them by the following lauxaniid characteristics: two fronto-orbital setae (rarely zero to one), both usually reclinate, but anterior seta inclinate in some genera; vibrissa absent (although subvibrissal setae can be enlarged near oral margin in some genera, such as Hypagoga, Trigonometopus, and Oncodometopus); ocellar setae proclinate; postocellar setae convergent; anepisternum with at least one strong seta on posterior margin; C unbroken; Sc complete, separated from R1; presence of costal spinules ending slightly beyond R2+3 (ending at R4+5 in Homoneurinae, with only one genus in the Neotropical Region, but common in the Nearctic). From the most similar family, the closely related Chamaemyiidae, they can be separated on the basis of the prescutellum being absent, the presence of preapical dorsal tibial setae, and the presence of distinct black costal spinules; some genera of chamaemyiids share the presence of reclinate fronto-orbital setae and an anepisternal seta.

Cryptochetids are easily recognized by their short, stout bodies, metallic color, and lack of arista.

Eggs spherical, oval, or capsular in shape, sometimes tapered (Figs. 136–138). Musso (1981) recognized three classes of eggs: pigmented (in Laphriinae); unpigmented (Asilinae); and eggs covered with sand grains (found in Antipalus varipes (Meigen), also Asilinae), which correlated with a ‘sand pouch’ found in the female (analogous to many Bombyliidae). Sand-covered eggs (in sandy “cocoons”) have since been reported for Dasypogon diadema (Fabricius), but this species of Dasypogoninae apparently does not have the sand pouch configuration found in A. varipes (Geller-Grimm, 1998b). Eggs with chorion ornamented or not. Larvae elongate, with relatively robust cylindrical or tapered shape, usually white or cream in color (Fig. 141). Head (Figs. 139–140) exerted, reduced in size; maxilla broad, triangular, dorsoventrally flattened; mandible small (strongly reduced or absent in Leptogastrinae). Thorax with one or more pairs of long setae on each segment (conspicuous in early stages, reduced thereafter); spiracles of mesoand metathorax greatly reduced, evidently non-functional. Abdominal segments 1–7 with small spiracle laterally; with one to four pairs of contractile warts; segment 8 usually short, with large, conspicuous spiracle posteriorly at dorsal edge; apparent segment 9 tapered, with four pairs of terminal bristles (two anteriorly, two posteriorly); anus slitlike; apical segment may terminate in short, blunt spine or process, sclerotized plate, or keel-like ridge.

Pupae mobile (not enclosed in skin of last larval instar). Head with row of prominent hooklike spines (antennal processes) in two groups: anterior process usually large; posterior group smaller, varying from three to five confluent hooks. Abdomen with seven or eight rows of short, stout spines of varied shapes; apex with one or more pairs of prominent spines or processes (Fig. 142).

Therevids are easily recognized by their uniform wing venation. Most Scenopinidae have an unbranched M vein (M1) curving towards R5 and joining or nearly joining it near the apex, while in therevids the wing tip is usually encompassed by cell r4 (branched R4 and R5), and the M veins (M1, M2, M3) are distinct, originating on the discal cell. The only exceptions are in the scenopinid genera Alloxytropus Bezzi, Caenotus Cole and Prorates Melander, but these usually still lack vein M3 (except for one species of Caenotus). The related families Apsilocephalidae (New World members are Nearctic only) and Evocoidae (Chile) differ in having a more delicate and thin body form with elongated legs, having wing veins R4 and R5 parallel, and having the distal flagellar segments longer than flagellomere 1. Evocoidae also differs in having wing vein R4 ending at the wing apex so cell r4 is below the wing apex, vein M3 is absent, the cell cup is open at the wing margin, and the anal lobe is very reduced to absent. Apsilocephalidae differs in having wing vein R5 ending at or above the wing apex so cell r4 is above the wing apex. Apioceridae and Mydidae, although often found in similar habitats, differ from therevids by branches of the M vein, which in the former two families are curved distinctly forward to the distal margin, with at least vein M1 ending anterior to the wing apex; in addition therevids do not have clubbed antennae as in most Mydidae. In comparison with Asilidae, therevids never have the typical asilid mouthparts and thickened legs adapted for predation, and therevids lack a mystax and the vertex is not deeply excavated. Although few bombyliids are superficially similar to therevids, those that are similar differ in wing venation, with the cell bm (when present) being pointed distally with three corners giving rise to three separate veins (M1+2, M3+CuA1, and CuA2) and the pointed base of cell m3 (when present), while in therevids cell bm is truncate apically, with four corners giving rise to four separate veins (M1+2, M3, CuA1, and CuA2) and cell m3 is truncate. Therevid females also have a novel internal reproductive structure associated with the spermathecae known as the spermathecal sac (Winterton et al., 1999a), although a similar structure is known in certain other asiloids, including Tongamya Stuckenberg (Mydidae) (Irwin & Wiegmann, 2001), Prorates (Scenopinidae) (Metz et al., 2002), Apsilocephalidae, and Evocoidae.

Larvae of Therevidae, and of their sister-family Scenopinidae, are readily separated from other flies based on the apparent secondary segmentation of the larval abdominal segments, giving the appearance of 17 segments (Woodley, 1989). These families are separated from each other by the condition of the metacephalic rod, being spatulate posteriorly in therevids (Fig. 21), rather than slender and parallel sided in scenopinids. Hauser & Irwin (2003) provided an overview of pupal morphology including details and figures for one species

Parvorder CALYPTRATAE Robineau-Desvoidy, 1830 (3 subordinate groups)

Comprende aquellas familias de schizophora en que los calypteres estan desarrollados.

Son moscas medianas a grandes.

Pedicelo de las antenas con una sutura longitudinal en el aspecto dorsal. la vena sc completa en todas las familias.

La vena r1 es corta o llega mas alla de la mitad del ala.

Tórax: con sutura transversal completa en el dorso del mesoscutum.

Callo postalar generalmente presente.

Celda r5 de lados paralelos o haciendose mas angosta hacia el margen del ala.

Superfamily Oestroidea Leach, 1815 (7 families)

Superfamily Muscoidea Latreille, 1802 (4 families)

Superfamily Hippoboscoidea Samouelle, 1819 (2 families)

Family Scathophagidae Robineau-Desvoidy, 1830 (57 genera, 419 species,

Family Anthomyiidae Robineau-Desvoidy, 1830 (53 genera, 1,941 species,

Family Oestridae Leach, 1815 (30 genera, 176 species,

Family Sarcophagidae Macquart, 1834 (173 genera, 3,094 species,

Family Tachinidae Robineau-Desvoidy, 1830 (1,597 genera, 9,626 species,

Family Fanniidae Schnabl & Dziedzicki, 1911 (4 genera, 359 species)

Family Glossinidae Theobald, 1903 (1 genus, 25 species)

Family Muscidae Latreille, 1802 (187 genera, 5,218 species,

Family Hippoboscidae Samouelle, 1819 (68 genera, 782 species)

Family Calliphoridae Brauer & Bergenstamm, 1889 (97 genera, 1,525 species,

Genus Hirmoneuropsis Bequaert, 1932

Genus Hyrmophlaeba Rondani, 1863

Moscas de tamaño mediano a grande (longitud corporal de 5 a 15 mm) (Fig. 1), generalmente brillantes, de color verde metálico, azul o violeta, al menos en el abdomen.

Frentes de los machos más estrechos y ojos más grandes que los de las hembras de la misma especie y, con pocas excepciones, carecen de cerdas orbitales proclinadas (Fig. 2); queetotaxia por lo demás similar en ambos sexos. Cerda frontal más baja que surge al nivel de la base de la antena (Fig. 3). Hembras de Mesembrinellinae con un solo par de cerdas cruzadas erectas en la vitta frontal anterior al triángulo ocelar, entre las cerdas frontales. Gena con dilatación genal bien desarrollada. Vibrissa suele surgir por encima del nivel del margen facial inferior, acompañada de cuatro o más cerdas subvibrisales cortas en Chrysomyinae, Calliphorinae y Melanomyinae, o por sólo una o dos cerdas grandes en Mesembrinellinae. La cresta facial generalmente se eriza más de la mitad de la base de la antena. Pedicelo con una sola cerda alargada entre unas pocas setas más cortas. Primer flagelómero similar en longitud en ambos sexos, o ligeramente más largo en la hembra. Arista largo plumoso dorsal y ventralmente. Palpus clavate, de longitud reducida en Cochliomyia. Notopleuron con dos setas. Pospronoto con tres cerdas principales en fila transversal, a veces precedidas por una cuarta cerda más pequeña; o con la cerda media desplazada anteriormente, en Mesembrinellinae; raramente, faltan cerdas internas. Meron con fila de cerdas bien desarrolladas.

Ala con curvatura en la vena M.

The single feature that distinguishes all Tachinidae from most members of the other families is the presence of an evenly convex subscutellum. A few calliphorids, especially some members of the genus Mesembrinella Giglio-Tos, also have a convex subscutellum, but have an enlarged posterior spiracle that extends anteriorly nearly to the posterodorsal corner of the katepimeron almost obliterating the katepimeron. Central American Rhinophoridae have a weakly convex subscutellum but lack the apex of vein M, as the vein ends in the membrane where the bend otherwise occurs. A few acalyptrates also have a moderately developed subscutellum but lack meral bristles. When alive, most tachinids hold their wings at a wider angle to the body than do sarcophagids and calliphorids, and most are also more active, seldom staying in one place for more than a few seconds. Tachinids are widely believed to be more bristly than members of the other families, but this is not a reliable criterion.

Los califóridos comparten con Sarcophagidae y casi todos los Tachinidae la presencia de una curva en la vena M, combinada con una fila de cerdas merales bien desarrolladas. Algunos Muscidae de la subfamilia Muscinae también tienen una curvatura en la vena M e incluso pueden ser de color verde metálico (Neomyia Walker) o azul (Morellia Robineau-Desvoidy) y, por lo tanto, se confunden con frecuencia con moscardones, pero carecen de cerdas merales. La mayoría de los califóridos, especialmente las especies comunes, se distinguen fácilmente de casi todos los Sarcophagidae y Tachinidae por su color azul o verde metálico brillante, especialmente en el abdomen. Algunos Tachinidae (Chlorohystricia Townsend, Chrysotachina Brauer & Bergenstamm) y Sarcophagidae (algunos subgéneros de Lepidodexia Brauer & Bergenstamm) también tienen el abdomen verde o azul metálico y, por lo tanto, podrían identificarse erróneamente como califóridos, pero los taquínidos tienen un gran subcutelo convexo, mientras que cuatro sarcófagidos metálicos tienen cerdas notopleurales. Los callifóridos no metálicos, incluidos algunos Mesembrinella y todos los Angioneura, son superficialmente similares a algunos múscidos.

Small to large flies (body length 3–25 mm), usually bristly

(Fig. 1), extremely varied in shape and proportions. Most

genera of usual calyptrate shape, but some slender and wasplike

in appearance. Frons of male either narrower than that of

female of same species and usually lacking proclinate orbital

bristles, or about as wide as that of female and usually also

with proclinate orbital bristles; first flagellomere of latter

type usually much longer in male than in female, in some

species bifurcate, trifurcate, or multibranched (fissicorn).

Arista usually bare, or short to long plumose in most Dexiinae.

Eye usually with ommatrichia, which may be long and

dense or so short and sparse as to make the eye appear bare;

ommatrichia usually longer in male than in female and often

longer dorsally than ventrally. Ocelli and ocellar bristles usually

present, the latter sometimes directed laterally (lateroclinate)

or posteriorly (reclinate). Upper orbital bristles usually

present, anterior one(s) reclinate, posteriormost sometimes

lateroclinate. Occipital setae usually long and fine, delineated

laterally by evenly spaced vertical row of stouter postoccipital

bristles, either extending ventrally to meet lower

cranial margin (Figs. 14, 30) or curving forward on genal

dilation (a setose strip or patch between eye and lower cranial

margin; Figs. 20, 27, 28), that may also have one or

more large bristles (Fig. 4). Vibrissa usually well developed,

usually accompanied by one or more smaller bristles both

below (on subvibrissal ridge) and above it (on facial ridge).

Face concave or convex, often protruding between and/or

below vibrissae. Facial ridge, especially in species with long

antennae, usually with row of erect bristles. Proboscis usually

short and straight, with padlike labella, but may be long

Glossina sp.

Euryomma peregrinum, wing

Fannia canicularis, wing

Vein R1 bare or setose above; A1 reaching wing margin.

M1 not strongly curved or angled forward on distal part; vein A1 usually traceable to margin of wing. Upper and lower calypteres usually forming subequal, modest-sized lobes, but exceptionally, lower calypter strongly reduced.

The family Scathophagidae is similar to Anthomyiidae and Muscidae. The Neotropical species of Scathophagidae may be distinguished from species of the other two families by two characters: only one katepisternal bristle present, and posterior margin of the lower calypter straight rather than slightly to strongly convex. The following characters together also separate most species from those of the other two families:

Vein A1 is present and usually extended halfway to wing margin, not reaching completely to wing margin as in most Scathophagidae and Anthomyiidae.

The curvature of vein Sc separates most Muscidae from Fanniidae: Sc lies close to R1 over most of its length in the majority of muscids, while it diverges from R1 at a point near its base in fanniids and some members of the muscid tribe Azeliini.

La vena A1 está presente y generalmente se extiende hasta la mitad del margen del ala, sin llegar completamente al margen del ala como en la mayoría de Scathophagidae y Anthomyiidae.

La curvatura de la vena Sc separa a la mayoría de Muscidae de Fanniidae: Sc se encuentra cerca de R1 en la mayor parte de su longitud en la mayoría de los muscids, mientras que diverge de R1 en un punto cercano a su base en fanniids y algunos miembros de la tribu muscid Azeliini.

Adult anthomyiids are 2–12 mm in length and can be recognized by their general color, usually brown, gray or black, never metallic; wings usually hyaline, rarely spotted; legs vary from yellow to black; lower calyptra welldeveloped; a pair of interfrontal setae is present; and posterior tarsus on the ventral surface has a noteworthy bristle at base. Most species have fine setae below the apex of the scutellum, except those in the genera Coenosopsia and Fucellia. Wing vein CuA+1 usually reaches the margin of the wing (Huckett 1987; Michelsen 2010), except in the genus Coenosopsia

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Topic

The nematoceran family Psychodidae (moth flies or drain flies) are small (<2 mm) true flies (Diptera) with short, hairy bodies and wings giving them a “furry” moth-like appearance. The adults have long antennae and the wings are leaf-shaped, either slender or broad, with the most elementary wing venation of any Diptera, having little more than a series of parallel veins without crossveins.

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Topic

Taxonomic Key

Classification

The position of Psychodidae in the phylogenetic system of the Diptera is unclear. Oosterbroek & Courtney (1995) presented summaries of classification of nematocerous flies, in which either an infraorder Psychodomorpha (with Deuterophlebiidae, Blephariceridae, Tanyderidae, Ptychopteridae, Nymphomyiidae and Psychodidae; Hennig, 1973), or a superfamily Psychodoidea (with Trichoceridae, Scatopsidae, Anisopodidae, Perissommatidae, Canthyloscelidae, and Psychodidae; Wood & Borkent, 1989) was recognized. The analysis of Oosterbroek & Courtney (1995) indicated that Psychodidae is the sister group of (Tipulidae + Trichoceridae) + (Anisopodidae + Brachycera). Equally unclear is the ranking of the different subgroups within the family. The description of a Sycorax-like psychodid from South America and its inclusion into the subfamily Sycoracinae by Duckhouse (1972) renewed discussion about the phylogenetic relations with other subfamilies of Psychodidae (sensu lato), and about the ranking of various taxa, especially of Phlebotominae. The latter group has been reviewed several times and has been treated either as a tribe within the Psychodidae (Fairchild, 1955), as a subfamily, or even raised to family rank because of the blood-sucking mouthparts of the females (Theodor, 1958; Rohdendorf, 1974; Abonnenc & Leger, 1976; Lewis et al., 1977; Dallai et al., 1984). Among the various proposals of the past four decades for a phylogenetic system of subfamilies within the Psychodidae sensu lato, Hennig’s (1972) is still most widely accepted.

At present six subfamilies are recognized, five of which occur in Central and South America: Phlebotominae, Bruchomyiinae, Sycoracinae, Trichomyiinae and Psychodinae. Horaiellinae (with its sole genus Horaiella Tonnoir, with 4 species) is restricted to the Himalayas, China and Thailand (Curler et al., 2006).

The systematics of the Phlebotominae are controversial. The present subdivision of the Phlebotominae into three Old World (Chinius Leng, Phlebotomus Rondani & Berté, Sergentomyia França & Parrot), and four New World genera (Brumptomyia, Lutzomyia, Warileya, and more recently Edentomyia) has been considered as stable by some (Young & Duncan, 1994), and premature by others (Dujardin et al., 1999; Galati et al., 2003). The generic composition of the subfamily Sycoracinae is debatable. Sycorax is probably confined to the Old World (Palearctic, Afrotropical, and Oriental Regions) and includes more than a dozen species at present. The Brazilian species described by Barretto (1956), and the Chilean species described by Tonnoir (1929) were revised by Duckhouse (1972), who proposed two genera (Aposycorax and Parasycorax) based mainly on the great differences of the genitalia. Young (1979) described 4 species of Sycorax from Colombia, different from Duckhouse’s Neotropical genera, with some characteristics that probably will group them in other taxa in the near future.

Within the Psychodinae, tribal classification, including the number of tribes and genera included, as well as the phylogenetic relationships among them, is highly unstable. In contrast, the recognition of genera seems less problematic. There are numerous features of the head, antenna, thoracic appendages, genitalia, and larvae that can be used for recognition, so that most genera appear to be good assemblages of closely related species. Species are mainly described based on males, but several descriptions are based on females only. The association of males and females is sometimes problematic.

Psychoda Latreille.

This species-rich genus with polyphagous larvae has several species that are widespread, with pantropical, or even cosmopolitan distributions. Some species probably are spread occasionally by human transport with vegetables, soil, or water. Many species are synanthropic with larvae and adults often found in human biotopes (e.g. houses, toilets, sewage treatment facilities, mushroom cultures). Life cycles are short, one to several weeks. There are proposals to split this genus into about 12 genera (Ježek, 1984; Ježek & Harten, 2005), but Bravo et al. (2006) considered them as subgenera. A total of 43 species are recorded from Central America (Bravo et al., 2006), of which 29 Costa Rican species were figured and keyed by Quate (1996, including those recorded as Tinearia Schellenberg).

Lutzomyia França.

Because the hematophagous species have been extensively studied for their medical importance, this is the most diverse Neotropical genus, with more than 300 species described, and with a large number of subgenera (more than 20) and species groups. Some of the subgenera have been treated as genera in recent literature. Young & Duncan (1994) provided a comprehensive survey of Central and South American Lutzomyia. There are about 97 species recorded in Central America.

Topic

Biology of Phlebotomine

Los flebótomos (Diptera: Psychodidade: Phlebotominae) son dípteros de importancia en salud pública debido a que son transmisores de diferentes patógenos, como los virus Phlebovirus, Vesiculovirus, Orbivirus (Hertig 1942; Tesh 1988; Comer y Tesh 1991; Killick- Kendrick 1990; Young y Duncan 1994); la bacteria Bartonella bacilliformis causante de la bartonelosis o enfermedad de Carrión, endémica en regiones por encima de los 1000 m en la cordillera de los Andes de Perú, Ecuador y Colombia (Hertig 1942; Young y Duncan 1994); protozoarios como Leishmania spp., causante de leishmaniasis con más de 20 especies presentes en 17 de los 35 países de las Américas, desde el sureste de Estados Unidos hasta el sur de Argentina (OPS 2018), que afectan una gran variedad de hospedantes mamíferos, incluyendo humanos (Lane 1993; Young y Duncan 1994; Akhoundi et al. 2016). En las Américas se han reportado más de 500 especies de flebótomos de los cuales 30 son vectores de leishmaniasis (Killick-Kendrick et al. 1990; Kato et al. 2010); de los tres géneros de flebótomos registrados para el Nuevo Mundo (América), Lutzomyia França, Brumptomyia França y Parrot, y Warileya Hertig (Lewis et al. 1977), sólo el género Lutzomyia es de importancia médica. Las hembras de estos Psychodidae son hematófagas y se alimentan de una gran variedad de vertebrados mamíferos (Young y Arias 1991; Killick-Kendrick 1999; Alexander 2000; Alvar 2001). Además, presentan actividad preferentemente en lugares oscuros como cuevas, agujeros de árboles, madrigueras, abrigaderos de animales domésticos y grietas en paredes de viviendas, los cuales son usados como sitios de reposo y/o criaderos.

En Perú, se han registrado 162 especies de flebótomos (Cáceres et al. 2000), con presencia en 22 de los 24 departamentos de Perú (Moo et al. 2017); con Lu. flaviscutellata (Mangabeira, 1942), Lu. yulli yulli (Young y Porter, 1972), Lu. tejadai (Galati y Cáceres, 1990), Lu. pescei (Hertig, 1943), Lu. ayacuchensis (Cáceres y Galati, 1988), Lu. peruensis (Shannon, 1929), Lu. verrucarum (Townsend, 1913) y Lu. ubiquitalis (Mangabeira, 1942), implicados en la trasmisión de las especies de Leishmania: Le. amazonensis, Le. braziliensis, Le. peruviana, Le. lainsoni y Le. guyanensis (SisLeish - OPS/OMS 2017).

En estas últimas décadas, se ha demostrado la emergencia y reemergencia de ciertos patógenos, resultado de las alteraciones ecológicas de un hábitat o modificaciones globales como el cambio climático (Lindsay y Birley 1996; Kovats et al. 2001; Reiter 2001; Molyneux 2003). Estas alteraciones han desencadenado epizootias y epidemias de enfermedades debido a la modificación en las interacciones de especies de organismos que cohabitan un medio, favoreciendo la colonización vectorial en un entorno modificado “antropizado” (Lainson y Rangel 2005; Shaw 2007). Las adaptaciones de flebótomos con capacidad vectorial a nuevos nichos ecológicos han originado nuevas áreas de transmisión de leishmaniasis (Fonseca da Silva et al. 2018).

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