Table Of ContentRevision of the Neotropical Family Inbiomyiidae
(Diptera, Schizophora)
Matthias Buck^ and S. A. Marshall^
ABSTRACT. The Central and South American family Inbiomyiidae with its single genus Inbiomyia
Buck is revised, including descriptions of 10 new species: 1. acmophallus Buck sp. nov. (Colombia), I.
anemosyris Buck sp. nov. (Colombia, Peru), I. matamata Buck sp. nov. (Venezuela, Colombia), I.
pterygionBuck sp. nov. (Bolivia), 7. regina Bucksp. nov. (French Guiana), 7. exulBuck sp. nov. (Costa
Rica, Ecuador), 7. empheres Buck sp. nov. (Ecuador), 7. anodonta Buck sp. nov. (Colombia), 7.
scoliostylus Buck sp. nov. (Costa Rica), and 7. zeugodonta Buck sp. nov. (Colombia, Erench Guiana).
Threespeciesknownfromsinglefemalesarebrieflydiagnosedbutnotnamed.Akeytothe 11 described
species ofInbiomyia is provided. The species-level phylogeneticrelationships are analyzed on the basis
ofamatrixof35 morphologicalcharacters. Eourspeciesgroupsarerecognized: the7. anemosyrisgroup
(7.acmophallus,7. regina,7.pterygion,7. matamata,7.anemosyris),the7. exulgroup(7. exul,Inbiomyia
species 1), the 7. mcalpineorum group (7. empheres, I. mcalpineorum, Inbiomyia species 2), and the 7.
scoliostylus group (7. anodonta, 7. scoliostylus, 7. zeugodonta, Inbiomyia species 3).
INTRODUCTION tropical part of the Neotropical region with the
apparent exception of the Caribbean. Inbiomyii-
The family Inbiomyiidae, recently erected for the dae occur in primary tropical forests (rain and
newly described Neotropical acalyptrate genus cloud forest) from sea level to 2000 m, with the
Inbiomyia Buck, 2006, is the fourth family of highestnumberofspeciesfound atlowelevations.
Cyclorrhapha to be described on the basis of The morphology, biology, family relationships,
a newly discovered group of flies in the past and history of discovery of Inbiomyiidae were
50 years (i.e., excluding families erected through discussedin detail byBuck (2006). Thefamilywas
taxonomic “splitting”). None of the other 3 described with the single included species Inbio-
families, Xenasteiidae Flardy, 1980, Neminidae myia mcalpineorum Buck, 2006; we here add
D.K. McAlpine, 1983 (originallyasa subfamilyof another 10 new species to the family and consider
Aulacigastridae), andMarginidaeD.K. McAlpine, the distribution and phylogeny ofthe species.
1991, occurintheNewWorld. Inbiomyiidae is in
fact the only valid family of Cyclorrhapha that MATERIALS AND METHODS
has ever been described from the New World on
the basis of a newly discovered genus. Other MORPHOLOGY AND TERMINOLOGY
relatively recently erected families were based on
genera described decades earlier, the most recent The morphological terminology largely follows J.E.
eMxcaAmlppliense,be1i9n7g0T(ebraasteodmyoznidTaeeraCtoolmlyeszsaaMnadllDo.cKh., iMnceAdlpoinnes(p1e9c8i1mae)n.sPocslteaarbeddomiinnalhofteat1u0re%s wKerOeHexaamn-d
subsequently neutralized in glacial acetic acid. Eggs
1933), and Syringogastridae do Prado, 1969 were obtained by dissection from cleared abdomens of
(based on Syringogaster Cresson, 1912). gravidfemales (museumspecimens). Dissectedparts are
Inbiomyiidae are characterized by several high- kept in glycerine in a microvial with the specimen.
ly unusual and unique morphological features of
the head, male and female genitalia, and egg. PHOTOGRAPHY
Buck (2006) placed Inbiomyiidae as the sister
group of the monotypic Australasian family The habitus photograph (Eig. 1) was taken with
Australimyzidae, within the superfamily Carnoi- a Microptics Digital Lab XLT imaging system using
dea. The larval biology of Inbiomyia remains a Canon EOS 1 Ds camera and Microptics ML-1000
unknown, but the unusual egg morphology flash fiber optic illumination system. Each image was
assembled from a series of photographs (with different
suggests a specialized biology. So far, species of focal planes) using the computer freeware CombineZ
Inbiomyiidae have been found in 8 Central and (Hadley,2005). Wingandeggphotographs (Eigs. 9, 10,
South American countries from Guatemala in the 13) were taken with a Nikon Coolpix 4500 digital
north to Bolivia and French Guiana in the south. camera mounted on a Zeiss compound microscope.
The actual range probably includes the entire
PHYLOGENETIC ANALYSIS
1-2. Department ofEnvironmental Biology, Univer-
sity of Guelph, Guelph, Ontario, Canada NIG 2W1. Parsimony analysis of the character matrix was
^Email: [email protected] performedwithPAUP"' 4.Ob10forWindows (Swofford,
Contributions in Science, Number 512, pp. 1-30
Natural History Museum ofLos Angeles County, 2006
2 Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae
Figure 1 Inbiomyia exul Buck sp. nov., female (Corcovado National Park, Puntarenas Province, Costa Rica).
mm
Scale 0.2
2001) usingthe “branchand bound” algorithm. Branch CBFC ColeccionBolivianadeFauna,LaPaz, Bolivia
support was calculated with PAUP'"'' according to CNCI Canadian National Collection of Insects,
Bremer (1994). The Bremer value for a certain branch Ottawa, Ontario, Canada
indicates the minimum number of additional steps DEBU Department of Environmental Biology, Uni-
(compared to the most parsimonious tree[s]) that is versity of Guelph, Guelph, Ontario, Canada
requiredtogenerateatree (ortrees) inwhichthebranch INBC Institute Nacional de Biodiversidad (INBio),
in question is unresolved. High Bremervalues therefore Santo Domingo de Heredia, Costa Rica
indicate strong branch support. LACM Natural History Museum of Los Angeles
County, Los Angeles, California, U.S.A.
ABBREVIATIONS UNCB Museo de Historia Natural, Institute de
Ciencias Naturales, Universidad Nacional de
Costal ratios: e.g., Cs2/Cs3 = length ofcostal sector 2/ Colombia, Same Ee de Bogota, Colombia
length of costal sector 3. Costal sector 2 is measured
between the apices ofR] and R2+3, Cs3 between apicMes
ofR2+3andR4+5,andCs4betweenapicesofR4+5and SYSTEMATICS
(Fig. 10).
ABBREVIATIONS OE DEPOSITORIES Family Inbiomyiidae
Inbiomyiidae Buck, 2006: 379 (type and only
AMNH American Museum of Natural History, New included genus: Inbiomyia Buck, 2006).
York, New York, U.S.A. Genus Inbiomyia Buck.
Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae 3
Figure 2 Species-level phylogeny ofInbiomyia. Explanations: Character numbers are given above the circles; states
for multistate characters below the circles. Characters whose polarity was established by the FIG-FOG method (see
text) are italicized on the tree. Bremer support values are shown in rectangles beside each node
Inbiomyia Buck, 2006: 379 (type species: L setulae in 2, often short rows. Pleuron without
mcalpineorum Buck, 2006, by original desig- hairs or bristles except on katepisternum. Legs
nation). slender. Forefemur with anteroventral, pos-
teroventral and posterodorsal series of bristles,
DESCRIPTION. A detailed description was femora otherwise devoid of bristles. Male fore-
provided elsewhere (Buck, 2006). The most tibia slightly clavate in some species (Fig. 7),
important morphological features are summa- usually with row of enlarged, more or less erect
rized below. ventral bristles on apical half (Fig. 6), tibiae
Small, mostly dark, acalyptrate flies (Fig. 1); otherwise devoid of bristles (including ventroapi-
body length 1.3-1.6 mm. cal bristle of mid tibia). Wing (Figs. 9, 10) with
Head (Fig. 3). Very short, wider than thorax, incomplete subcosta andvery shortRi; costa with
M
with protuberant eyes. Head chaetotaxy reduced, humeral and subcostal breaks, reaching
including single long, inclinate orbital bristle, (weaker in last sector), with small, well-spaced
inner and outer vertical bristles, long vibrissa, spinules on second and base of third sector. Cell
shorter subvibrissal and several genal bristles, and cup with convex outer margin; anal vein very
strong ventral preapical bristle on palpus. Ocellar strong, fading away before wing margin. Halter
plate very large and shining, occupying most of with darkened knob.
frons. Ptilinum extremely reduced. Face with Preabdomen. With 4 large tergites (syntergite
arcuate, ridgelike internal thickening at or above 1+2, tergites 3 and 4, male syntergite 5+6 or
lower margin, here called “lower facial ridge” female tergite 5).
(Fig. 5: fr); lower margin deeply emarginate. Eye Male Terminalia. Sternites 5, 6, and 7 fused,
short-pubescent. Antenna short with small scape separate from short, transverse sternite 8 (Buck,
and pedicel, and hemispherical first flagellomere 2006: figs. 4A-C). A small ring-shaped sclerite
bearing extremely long, pubescent, dorsoapical (nonfunctionalremnantofleftspiracle 7?) present
arista. Proboscis with small, largely separate dorsally between syntergite 5+6 and synsternite
labella pointing in nearly opposite directions. 5+6+7. Epandriumsaddle-shaped, anteroventrally
Thorax. Strongly arched, humpbackedwiththe with a pair of slender processes on each side
following complement of bristles: 3 dorsocentrals (Fig. 15: vp) which are either fused to or
(one presutural), 2 supra-alar (pre- and postsu- articulated with the epandrium, posteroventrally
tural), postalar, small postpronotal (sometimes with a pair of lobes (fused cerci?) behind each
hairlike), 2 notopleurals (posterior one inserted surstylus (Fig. 16; vl). Perianal field (Fig. 16: pf)
much higher than anterior one), (l-)2 scutellars usually closed ventrally by subanal plate (divided
(anterior one usually hairlike, sometimes absent), medially by suture). Surstylus (Fig. 15: ss) usually
and dorsoproclinate katepisternal. Acrostichal with slender basal and dilated distal portion.
—
4 Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae
Figures3-8 Inbiomyiaheadsandmaleforetibiae. 3.1. mcalpineorumBuck,head,frontal (slightlymodifiedfromfig.
3A in Buck, 2006). 4. I. matamata Buck sp. nov., head capsule, frontal. 5. 7. acmophallus Buck sp. nov., ditto. 6. 7.
exul Buck sp. nov., ditto. 7—. 7. regina Buck sp.—nov., left mal—e foretibia. 8. 7. anemosyris B—uck sp. nov., ditto. Scale
0.1 mm. Abbreviat—ions: as antennal—socket, cl clypeus, fcl f—rontoclypeal membrane, fr lower facial ridge, or
orbitalbristle,ors orbitalsetulae,vte outerverticalbristle,vti innerverticalbristle.Drawings:A. Cormier(3) and
M. Musial (4-8)
usually bearingconspicuous bristles. Hypandrium part nonstriate with small spicules and/or micro-
Y-shaped with slender anterior apodeme (Fig. 15: trichia. Ejaculatory apodeme small and incon-
ha). Postgonites (Fig. 16: pg) well developed spicuous, disklike (Fig. 21).
(absent in 1 species), with stout bristles. Phalla- Female Terminalia. Ovipositor broadly trun-
podeme rodlike, separate from hypandrium cate, nontelescopic; tergite 7 narrowed or divided
(Fig. 16: pa). Phallus clearly divided into basi- medially, sternite 7 comparatively broad and
and distiphallus (Fig. 20: bp, dp). Basiphallus long, laterally almost reaching corresponding
sclerotized, with simple, anchor- or shieldlike tergite (narrowly fused to tergite in some species);
epiphallus. Distiphallus short and simple, com- all sclerites beyondsegment 8 completelyreduced,
pletely membranous; surface in part striate, in cerci absent. Intersegmental membrane behind
Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae 5
Figures 9-13 Inbiomyiawings andeggs. 9. I. extilBuck sp. nov., rightwing. 10. /. anemosyris Bucksp. nov., ditto.
11./. acmophallusBucksp. nov.,egg, dorsal (firstpublishedinBuck,2006: fig. 5C). 12. L zeugodontaBucksp. nov.,
mm
d0i.t0t5o.m1m3.(Ie.ggesx)u.lABbubrcekvisapt.iononvs.:,Cdsit—toc,os(tpahlosteocgtroar,phmpof—clmeiacrreodpyalne.d Dsrlaiwdei-nmgosu:ntAe.dCsopremciiemren)(1.1,Sc1a2l)e 0.2 (wings),
sternite 7 pouchlike, with a field of conspicuous, across the genus. Size large compared to abdo-
posteriorlydirected (wheninvaginated), serrate or men, shape elliptical (length 0.39-0.48 mm,
crenulate scalelike cuticular outgrowths. Sternite width 0.21-0.27 mm, length/width ratio 1.67-
8 hardly discernable, completely membranous 1.86), strongly flattened dorsoventrally (thickness
and medially divided. Two moderately sclero- estimated at ca. 0.05 mm). Ventral surface mem-
tized, tire-shaped, oval, or egg-shaped spermathe- branous, unornamented; dorsal surface thicker,
caewith striate surface. Spermathecal ducts short, coarsely granulose and unpigmented, with polyg-
joining genital chamber separately; duct insertion onal, reticulated pattern. Micropyle on small
lateral to main axis of spermatheca. elevation.
Egg (Figs. 11-13). Mature eggs were obtained DIAGNOSIS. Inbiomyia (Inbiomyiidae) can be
from 5 species (/. acmophallus sp. nov., /. identified using the key to Neotropical Diptera
pterygion sp. nov., I. regina sp. nov., I. exul sp. families by Buck et al. (in press). The following
nov.,I. zeugodonta sp. nov.) andareveryuniform characters are diagnostic: small, acalyptrate flies
6 Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae
Table 1 Character state matrix for described species of Inbiomyia. Australimyzidae, the sister group of
Inbiomyiidae, servedas outgroupforthe analysis.Putativeground-plancharacterstatesofInbiomyiidae (“ancestor”)
were in part determined after the analysis using the FIG-FOG method (see text).
Character numbers
1 2 3
Species 1234567890 1234567890 1234567890 12345
Australimyzidae 0000000000 on 0700077700 70007
(ancestor 0000000000 0700000000 0700000700 00000)
1. anemosyris 1111001010 0000011000 2002000011 00012
1. matamata 1011111010 0000011000 2002000001 00011
I. pterygion 1111111010 0000011000 2002002011 00101
I. acmophallus 0010101000 1000011101 2001012001 00110
I. regina 1011111010 1000211101 2001012001 00110
I. exul 1111000110 1010000022 0110001100 10000
I. mcalpineorum 0000100101 0111000000 1001100000 00000
I. empheres 0000100111 0111000000 1001100001 00000
I. anodonta 0070000001 0111010001 2102000001 07777
0000000000 0111100011 1120000101 lioio
I. scoliostylus
I. zeugodonta 0000000001 0111100011 1120000100 11010
with (1) very shortened head (frons) and pro- SPECIES-LEVELPHYLOGENYOEINBIOMYIA
truding eyes, (2) single pair of inclinate orbital
bristles, (3) well-developed vibrissae, (4) no Characters Used in Phylogenetic Analysis
ocellar or postvertical bristles, (5) extremely long
Characters were polarized based on the putative
and nearly apically inserted arista, (6) strongly outgroup Australimyzidae (Buck, 2006). No
divergent and largely free labella, (7) aset(ul)ose
additional outgroups were taken into consider-
afonrepriesdtuecrendums,ub(c8o)stcao,mpalnedte(9w)icnogstvaenwaittihonhuemxecreaplt ation because the sister group of Inbiomyiidae +
and subcostal breaks. In previously published Australimyzidae is uncertain (Carnidae are the
putative sister group of Inbiomyiidae + Australi-
keys, Inbiomyia runs to Aulacigastridae (e.g.,J.F.
McAlpine, 1981b) or does not key out properly myzidae, butsupportforthisrelationship isweak;
see Buck, 2006). Inbiomyiidae and Australimyzi-
(e.g., Hennig, 1973: conflict at couplet 138).
BIOLOGY. There are no directobservationson dae have diverged considerably, especiallyinmale
and female postabdominal features. Fourteen of
the biology of Inbiomyia. Adults are apparently
microbial grazers as can be concluded from the the 35 characters (Table 1), mostly male genitalic
characters, could therefore not be polarized
substantial amounts of fungal, algal (including
through outgroup comparison. Eleven of these
diatoms), and probably bacterial material found
in the guts ofdissected specimens. Nearly 80% of characters could be polarized a posteriori (i.e.,
after the parsimony analysis; see “ancestor” in
the currently known material of Inbiomyia {ca.
220 specimens in total) was collected in Malaise Toaubtlgero1u)pu”si(n“gFtIhGe-“FfOuGn”ct)ioanpaplrionagcrhou(pW-afturnocutsioannadl
traps, another 16% in pan traps placed in Wheeler, 1981). The polarity of 3 characters (12,
decaying vegetation (mostly decaying foliage of
22, 28) remains ambiguous.
fallen trees). The junior author observed an adult In the following list of characters, apomorphic
of an unidentified species on green leaves of states are coded with “1” or “2” and plesio-
a recently fallen tree at Kara Avis (Heredia, Costa morphic states with “0” (except characters 12,
Rica). The presence of adults on decaying plant
22, 28). Character 21 is ordered. Polarities of
matter suggests a phytosaprophagous larval bi- characters denoted by asterisks ('•') were estab-
ology (larvae feeding on the microbial flora ofthe
lished a posteriori.
substrate). However, the comparatively large egg
size and low number of mature eggs per female
Head, Thorax, and Legs
(1-4 in dissected specimens) is unusual for species
developing in a nutrient-rich, rapidly decaying,
ephemeral food source. Most Diptera that exploit 1 Fa—cial sclerotization below l—ower facial ridge:
this type of substrates are r-strategists and pro- 0 largely absent (Fig. 5); 1 present (Fig. 6)
duce large numbers of relatively small eggs. The 2 Ocellar hairs: 0—present; 1—-absent
unusual egg morphology, which is reminiscent of 3 Inner vertical bristle, length: —0-—more than
macrotype Tachinidae eggs, could be indicative of halfaslongasorbital bristle; 1 -atmosthalf
a more specialized, possibly parasitic, lifestyle. as long as orbital bristle
—————— —————
Contributions in Science, Number 512 25 Buck and Marshall: Revision ofInbiomyiidae 7
4 Dorsocentral hairs between postsutur—al and Postgon—ite, stout, peglike bristl—es of apical
prescut—ellar dorsocentral bristles: 0 pres- row: 0 less than 8 (Fig. 14); 1 more than
ent; 1 absent 8 (Fig. 61)
5 Male foretibia, row of ventral bristles: 0 26"' Postgonite, stout—apical bristle and slender
absent or weakly developed (Fig. 8); 1 pr—eapical hair: 0 small or absent (Fig. 24);
strongly developed (Fig—. 7) 1 enlarged (Fig. 14—)
6 Male foretibia shape: 0 simple (Fig. 8); 1 IT" Epiphallus shape: 0 slender —(Fig. 29); 1
slightly clavate (Fig. 7) broad throughout (Fig. 60); 2 apically ex-
panded (Fig. 19) —
28 Epiphallus, dorsal surface: 0 smooth; 1
Male Terminalia striate (Eig. 57) (polar—ity unknown)
29 Distiphallus shape: 0 simple (Fig. 20); 1
7 S0-y—ntergite 5-f-6, ventrallobe belowspiracle 6: twisted (Fig. 44) —
0 absent; 1—-present (Fig. 17: tl) 30 Distiphallu—s vestiture: 0 microtrichose
8 Ventral portion of synsternite 5+6+7 {- (Fig. 60); 1 bare —
0st—ernaibtseen5t),(Fipgo.st1e7r)i;or1—prporceessesnto(fFigr.ig5h4t:spirdve): 31'‘' D1i—stsimphoaoltlhus surface: 0 spinulose (Fig. 29);
=
9 Lateral portion of synsternite 5+—6-1-7 (
sternite 6), posterior process: 0 absent
(Fig. 17); 1-—present (Fig. 27: prl) Female Terminalia
10 Epandrium, dorsolateral anterior apodemes:
—absent (Fig. —15); 1—present (Fig. 62—: ap) 32 Female tergite 7: 0—entire; 1—medially
!!" Perianal field: 0 elliptical (Fig. 41); 1 not divided (Fig. 81) —
12 eSlulbiaptniaclal p(lFaigt.e—:160)—desclerotized or weakly 33 pFiegmmaelnetesdt;ern1i—tepa7r,tipailglymepnatlaetion: 0 evenly
uscnlkernootwizne)d; 1 strongly sclerotized (polarity 34 Female tergite 8,—anterior apodeme: 0
13 Ab1rr—toiacdu;la1t—ionnaorfroewpa(nBdurciku,m/2h00y6p:an0fdirg—.iu4mE:) 0— 35'‘‘ oaSvbpaselernm(taFti(ghFsei.gc.a3:78,0)03;—8)1;ti2rp—er-eseshgeagnp-tesd(hFai(pgF.iegd.81(2)F2i)g;. 213)
14 Ventral process of epand—rium: fused to
epandrium (Eig. 15); 1 articulated with
epandrium (Eig. 62) RESULTS OF PHYLOGENETIC ANALYSIS
15'‘‘ Ventral process of epandrium, apex: 0
—
slender (E—ig. 18); 1 broadly truncate The analysis yielded a single most parsimonious
(Eig. 87); 2 bilobate (Eig. 49) tree (Eig. 2; tree length 71, consistency index =
16"‘ Ventral process of epandrium, shape: 0 0.61, retention index = 0.72, rescaled consistency
curved medially (Fig. 18); 1—not curved index = 0.44). Four species groups are recog-
medially (Fig. 66) nized: the I. anemosyris group (/. acmophallus, I.
17"' Ventral process ofepandrium, small point at regina^ 1. pterygion, I. matamata, I. anemosyris),
or near apex: 0—-absent (Fig. 66); 1—pres- the /. mcalpineorum group (/. empheres, 1.
ent (Fig. 18) mcalpineorum), the I. exul group (I. exul), and
18''' Ventral process of epandrium, stout blunt the I. scoliostylus group (/. anodonta, I. scolio-
bristlesoninnersurface: 0—absent (Fig. 28); stylus, I. zeugodonta). The latter 3 groups each
-present (Fig. 18) — include 1 unnamed species (see species accounts
19'‘‘ Posteroventral —lobe of epandrium: 0 large below), which were not included in the analysis
(Fig. 16: vl); 1 small (Fig. 76); 2—-absent because they are known from females only. Most
(Fig. 57) — species groups are well defined and show moder-
20"', Surstylus basal portion:—0 more or less atetohighBremersupport. The exceptionisthe1.
parallel-sid—ed (Fig. 25); 1 widening distally scoliostylusgroup, which iswell supported onlyif
(Fig. 15); 2 -tapering distal—ly (Fig. 56) L anodonta is excluded. The latter shares only 1
21 Surstylus ap—ical portion: 0 not expanded synapomorphy with the remainder of the species
(Fig. 56); 1 s—lightly to moderately expand- (character 20, state 1), which occurs convergently
ed (Fig. 62); 2 greatly expanded (Fig. 15) in I. acmophallus -l- /. regina.
22 Surstylus apical portion,—discal bristles: 0
none or 1 (Fig. 15); 1 several (Fig. 68) Biogeography
(polarity unknown)
23 Surstylus apical portion, peglike bristle(s): The /. anemosyris group is the only species group
0—absent (Fig. 15); 1—1 (Fig. 57); 2—2 of Inbiomyia restricted to South America. All
(Fig. 85) other groups are represented in both Central and
24 Surstylus—apical portion, marginal row of South America. Because of the still fragmentary
hairs: 0 —absent (Fig. 75); 1—irregular knowledge on Inbiomyia (many species still to be
(Fig. 47); 2 regular (Fig. 25) discovered, poorlyknownspecies distributions), it
8 Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae
is difficult to develop meaningful hypotheses on (Fig. 25). Female tergite 7 entire; spermatheca
the phylogeographyofthegroup. Interestingly, all asymmetrically egg-shaped (Fig. 23) .... 4
Central American species have their sister species 4 Ocellar plate glossy, without any trace of
in SouthAmerica (/. exul, occurringin both South microtomentum. Male foretibia slightly cla-
and Central America, and its Central American vate, with distinctly enlarged, nearly erect
sister species, Inbiomyia sp. 1, represent a special ventral bristles (Fig. 7). Slender basal portion
case). On the basis of current phylogeny, it of surstylus not extending beyond apex of
appears most likely that the stem species of posteroventral epandrial lobe (Fig. 46). Ven-
Inbiomyia lived in South America, and its tral epandrial process with distinctly bilobate
descendants colonized Central America on at apex; median lobe with several stout apical
least 3 different occasions. bristles (Fig. 49). Female sternite 7 partially
desclerotized and pale, especially sublaterally
KEY TO THE SPECIES OF INBIOMYIA BUCK (French Guiana) . . . /. regina Buck sp. nov.
(Note: Three unnamed species are known from - Ocellar plate shining but with thin microto-
females only and are not included in the key.) mentum. Male foretibia simple, with only
slightly enlarged, inclined ventral bristles
(Fig. 8). Slender basal portion of surstylus
1 Entire katepisternum (and sometimes also extending far beyond apex of posteroventral
ventral margin ofanepisternum) white, yellow epandrial lobe. Ventral epandrial process not
or brownish yellow, contrasting with medium bilobate, with minute, bare median projection
todark brownanepisternum (e.g.. Fig. 1) . . . (Fig. 25). Female sternite 7 evenly sclerotized
2 and brown (Colombia, Peru)
- At least dorsal portion of katepisternum /. anemosyris Buck sp. nov.
medium to dark brown and hardly paler than 5 Cs2/Cs3 < 2.5. Surstylus with greatly expand-
anepisternum 5 ed apical portion bearing discal as well as
2 Upper pleuron (anepisternum, anepimeron, marginal bristles (Figs. 68, 69); marginal
postnotum) uniformly dark brown, sharply bristles few in number and spaced widely.
contrastingwithwhitetowhitishyellowlower Female unknown. [Face not sclerotized below
pClse2u/rCosn3th<rou2g.h0o,utCsit3s/wChso4le>le2n.g5th ((FFiigg.. 91)).. larotwiecrulafatceidaltoriedpgaen.drVieunmt.r]al(Ceopalnodmrbiiaal)pr.o.ce.s.s
S(Fuirgs.ty5l6u)s. aHnyguplaantde,riunomt weixtphanldoendg,apfiocrakleldy - Cs2/Cs3 > 3.0. Sur/s.tyalnuosdwointthamBoudcekrastpe.lnyovo.r
ventral process (Fig. 53). Female tergite 7 greatly expanded apical portion, if greatly
entire (Fig. 52). Spermatheca tire-shaped expanded then with marginal bristles only
(Fig. 52) (Costa Rica, Ecuador)
I. exul Buck sp. nov. m(eo.gr.e. cFliogs.e-1s5e)t; marginal bristles, if present6,
- Upper pleuron not uniformly colored; at least
6 Inner vertical bristle almost as long as orbital
anepimeron paler than anepisternum, the
bristle. Anteriorpairofscutellar bristleslonger
former not sharply contrasting with yellow
lower pleuron. Cs2/Cs3 > 2.8, Cs3/Cs4 < 2.3 than acrostichals, ca. one-third as long as
posterior scutellars. Ventral epandrial process
(e.g.. Fig. 10). Surstylus either greatly expand-
ed apically or not angulate. Hypandrium biangulate in lateral view; surstylus lacking
without forked ventral process. Either female fringe of long bristles, with 2 stout, peglike
tergite 7 divided or spermatheca asymmetri- bristles apically (Fig. 75). Female tergite 8
cally egg-shaped 3 with long anterior apodeme (Fig. 81: ap);
3 Inner vertical bristle ca. 0.75X as long as tergite 7 broadly divided medially (Fig. 81)
0or.b4iXtalasblroisntglea.sApnotsetreriioorrsocnuet,eldliasrtibnrcitsltylelo0n.ge3r- - I(nCnoesrtaveRritciaca)l.br.istIl.es<col0i.o7stXylausslBouncgkasspo.rbniotva.l
and more robust than acrostichal hairs. 1-3 bristle (Fig. 3). Anterior pair of scutellar
dorsocentral setulae between postsutural and bristles short, hairlike and subequal to acro-
prescutellar dorsocentral bristles. Apex of stichals, or absent. Ventral epandrial process
surstylus slightly expanded, without fringe of straightinlateralview. Surstyluswithfringe of
long bristles (Fig. 84). Female tergite 7 nar- 6-16 long bristles, lacking stout peglike
rowly divided medially; spermatheca tire- bristles. Female tergite 8 without or with very
shaped (Fig. 82) (Colombia, French Guia- short anterior apodeme, tergite 7 entire or
na) /. zeugodonta Buck sp. nov. medially divided 7
- Inner vertical bristle at most half as long as 7 Male perianal area elliptical, sharplydelimited
orbital bristle. Anterior scutellar bristle small ventrally by dark brown, sclerotized subanal
and hairlike, subequal to acrostichal hairs. No plate (e.g.. Fig. 63). Surstylus only moderately
setulae between postsutural and prescutellar expanded apically, andwith fringe offew long
dorsocentral bristles. Apex ofsurstylus greatly bristles (e.g.. Fig. 62). Femaletergite 8 without
expanded, with fringe of long bristles anterior apodeme. Ca. 2 dorsocentral setulae
Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae 9
between postsutural and prescutellar dorso- colorous with remainder of leg. Expanded
central bristles 8 distal portion of surstylus broadly rounded
Male perianal area not elliptical, either poorly anteriorly (Fig. 40). Epiphallus greatly ex-
delimited ventrally from weakly sclerotized panded apically, anchor-shaped (Fig. 44).
subanal plate (e.g., Fig. 32) or its margins Ventral epandrial process not tapered, with
parallel to slightly divergent ventrally several long hairs on inner surface of apical
(Fig. 16). Surstylus broadly expanded distally, fourth (Fig. 43). Female sternite 7 darker
with fringe of numerous long bristles (e.g., medially (Bolivia)
Fig. 15). Either female tergite 8 with short I. pterygion Buck sp. nov.
anterior apodeme (Fig. 22) or scutum without
setulae between prescutellar and postsutural SPECIES ACCOUNTS
dorsocentral bristles 9
8 Subanal plate and hind margin of epandrium
almost forming a right angle in lateral view; Inbiomyia anemosyris species group
vrieonrtramlaregpiann;dripaolstpgrooncietses wdiitrhectsetrdaiaghntteproisotrel-y SPECIES INCLUDED. I. acmophallus sp. nov.,
m(eBduicakl,ly20v0e6ry: nfaigr.ro6wB).butFenmoatleintteerrrguiptteed7, pI.tearnyegmioosnyrspi.snsopv..,noavn.d,L/.rmeagitnaamaspt.anosvp.. nov., /.
DIAGNOSIS. Inner vertical bristle at most half
laterally fused to sternite (Buck, 2006: fig.
7A) (Costa Rica) ..../. mcalpineorum Buck as long as orbital bristle. Facial sclerotization
- Angle formed by subanal plate and hind usuallyextending belowlowerfacial ridge (Fig. 4;
margin ofepandrium obtuse (Fig. 62). Ventral except in /. acmophallus). Usually without dorso-
central setulae between postsutural and prescu-
epandrial process with a posterior tubercle at
dbiarseecteodf vaepnitcraalllyth(iFridg. 6(3Fi)g..F6e5m).alePotsetrggointeite7 tluesl)l.arRdo2r+3socneonttrsahlorbtreisntelde,s (Cesx2c/eCpts3in>I. a3c.0m.opMhaalle-
syntergite 5-1-6 with a pair of ventral lobes below
medially interrupted, laterally not fused to
spiracles 6 (Fig. 17: tl). Ventral epandrial process
sternite (Fig. 80) (Ecuador) . fused to epandrium, not curved medially, at or
I. empheres Buck sp. nov. near apex with a small, usually medially directed
9 Face not sclerotized below lower facial ridge
toothlike process. Surstylus with greatly enlarged
(Fig. 5). Male perianal field angulate ventro- apical portion, bearing distinct row of long
laterally (Fig. 16). Fringe of bristles of distal marginal bristles and no discal bristles. Flypan-
portion of surstylus extending far onto dorsal drium broadly connected to epandrium. Disti-
margin (Fig. 15). Ventral epandrial process phallus spinulose, without microtrichia.
with 2 long apical hairs and several stout,
blunt-tipped bristles on medial surface Inbiomyia acmophallus Buck sp. nov.
(Fig. 18). Female sternite 7 pigmented darker Figs. 5, 11, 14-22; Buck, 2006: fig. 5B
laterally and medially. Spermatheca tire-
shaped (Fig. 22) (Colombia) ADULT. Wing length 1.41-1.49 mm {$),
1. acmophallus Buck sp. nov. 1.44-1.56 mm ($). Sclerotized portions of head
- Facial sclerotization extending below level of dark brown excluding prementum, which is
lower facial ridge (Fig. 4). Perianal field yellow to brownish yellow. Antenna dark brown,
elliptical. Fringe of bristles of distal portion almost concolorous with frons; palpus slightly
of surstylus ending at or near posterior corner paler. Frons, besides usual long inclinate orbital
(e.g.. Fig. 40). Ventral epandrial process with- bristle, with 2 orbital hairs and a pair of ocellar
out apical hairs, medially with slender fine- setulae. Ocellar plate sparsely microtomentose.
tipped bristles only (e.g.. Fig. 43). Female Face not sclerotized below lower facial ridge
sternite 7 often evenly pigmented, if not then (Fig. 5). Thorax including pleuron dark brown,
darkermedially butnotlaterally. Spermatheca katepisternum slightly and gradually becoming
oval (Figs. 37, 38) 10 paler ventrally and anteriorly. 1-3 dorsocentral
10 Ocellar plate glossy and lacking microtomen- setulae between postsutural and prescutellar dor-
tum on each side of ocellar area. Forecoxa socentral bristles; acrostichal rows of hairs ex-
brown, at least basal half distinctly darker tending posteriorly to at least level of postsutural
thanremainderofleg. Expanded distalportion dorsocentral bristle. Scutellum with posterior pair
of surstylus with angulate anterior corner of bristles only, anterior pair absent. Legs in-
(Fig. 31). Epiphallus nearly parallel-sided cluding forecoxa brownish yellow. Both antero-
(Fig. 35). Ventral epandrial process tapered, and posteroventral rows of bristles of forefemur
without long hairs in apical fourth (Fig. 34). strongeranddarkerinmalethaninfemale;bristles
Female sternite 7 evenly pigmented (Vene- of posteroventral row slightly stronger than
zuela, Colombia) anteroventral row in both sexes. Distal half of
1. matamata Buck sp. nov. male foretibia with well-developed row of short,
- Ocellar plate shining but sparsely microto- semierect, blackish, ventral bristles. Wing slightly
mentose throughout. Forecoxa yellow, con- infuscate. Cs2/Cs3 = 3.0-3.8 Cs3/Cs4 = 1.6-2.1.
,
——
10 Contributions in Science, Number 512 Buck and Marshall: Revision ofInbiomyiidae
Figures 14-21 Inbiomyia acmophallus Buck sp. nov., male terminalia. 14. Left postgonite, anterior. 15.
Hypopygium, left lateral. 16. Ditto, posterior (bristles of postgonites and left surstylus omitted). 17. Segments 5-8,
ventral. 18.Leftventralepandrialprocess,anterior. 19.Phallapodemeandbasiphallus,dorsal.20.Phallus,ventral.21.
mm
Ejaculatory ap—odeme, posterodor—sal. Scale 0.05—(15-17, 19-20), 0.025 — (14, 18, 21). —Abbreviations: bp
basiphallus, d—p distiphall—us,eph epiphallus—, ha hypandr—ialapodeme,—pa phalla—podeme,pf perianalfield,pg
po—stgonite, S sternite, sa subanal plate,—sp spiracle, ss surstylus, T tergite, tl ventral lobe of syntergite 5+6,
vl posteroventral lobe ofepandrium, vp anteroventral process ofepandrium. Drawings: A. Cormier
MALE TERMINALIA (Figs. 14-21). Synster- of anterior apodemes dorsally, ventrally continu-
nite 5+6+7 without posterior process on ventral ous with subanal plate, which is less sclerotized
or lateral portion (Fig. 17). Ring-shaped sclerite and paler than epandrium. Perianal field ventrally
present but faint. Epandrium brown, withoutpair with parallel to slightly divergent lateral margins
