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June 1997
A revision of the Sclerocoelus galapagensis group (Diptera: Sphaeroceridae: Limosininae)
S. A. Marshall Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1, Canada
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Marshall, S. A., "A revision of the Sclerocoelus galapagensis group (Diptera: Sphaeroceridae: Limosininae)" (1997). Insecta Mundi. 260. https://digitalcommons.unl.edu/insectamundi/260
This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 11, No. 2, June, 1997 97
A revision of the Sclerocoelus galapagensis group (Diptera: Sphaeroceridae: Limosininae)
S. A. Marshall
Department of Environmental Biology, University of Guelph, Guelph, Ontario, N1G 2W1, Canada.
Abstract: The Sclerocoelus gcilapage~lsisgroup is defined and revised, including the description of S. galapage~~sisnew species from the Galapagos Islands; S. cariberlsisnew species from the Caribbeanand adjacent areas; S. brasilertsisnew species from Brazil, Ecuador, Colombia, ancl Panama; S. ltetttorrl~oidalisnew species from Ecuador and Venezuela; and S. andensis new species from Argentina, Bolivia, and Venezuela. The south Atlantic species Sclerocoelus sl~bbreuipert~iis(Frey), new combination, is rctlescribed as a member of the S. galapage~tsisgroup, and is considered the sister species to the rest of the species group. A lcey to species, character matrix, ant1 claclogram are provided.
Introduction Diagnosis of the genus Sclerocoelr~s:Mem- The genus Sclerocoelus Marshall is abundant bers of the genus Sclerocoelzrs can be recognized by in the New \Vorltl, with over 40 species clistributed thc combination of a very broad alula, an enlarged from Canada t,o Argent,ina, ant1 with one species, intra-alar bristle just anterior to the pre~cut~ellar Sclerocoelz~ss~rDbrevil)e~t,r~is (l?rey) from the south dorsocentral bristle, a very broad lunule and char- Atlantic. The latter species is a member of the acteristic tibia1 chaetotaxy including a micl ventral distinctive Sclerocoelu,sgalc~~)ager~sisgroup, and is bristle only in the female. Characteristics of the here redescribcd from Tristan (la Cunha. The S. male genitalia, especially the complex subcerci ancl gal~pager~sisgroup also includes five other new genital pouch, are definingfor the genus (Marshall, species from the Galapagos Islands, Mexico, the 1995). Caribbean, Central America, and South America Diagnosis of the Sclerocoelus galapagen- (M~I)). sis group: All mcmbcrs of the species group have Most Sclerocoell~sare a~sociat~cdwith wet, cle- strongly developed male perianal pacls, character- caying vegetation in cloucl forests, where they are istically shal)ccl surstyli, and one (or rarely 2) stout often collected in large numbers by sweeping a net apical bristle on each female cercus. All Neotropi- ovcr moist dead leaves. The species group revised cal specics have setae on the membrane flanking here, the S.gnlapcrg.ert.sis gronl), is the most clistinc- the prosternum. tive and easily diagnosecl group of Sclerocoelus. All Description of the Sclerocoelusgalapagen- species in the group can easily be distinguished sis group: Dark, heavily pollinose limosinines, from congeners by the conspicuous, sclerotized 2.0-3.0mm, usually ca. 2.5mm. Interfrontal area male perianal patls or by the spinose apical bristles tapcred, width at top equal to height; 3 equal of the female cerci. They are also generally larger interfrontal bristles. Lunule prominent, broadly than congeners ancl, as outlinecl in the diagnosis rounded. Fore cosa with 3-4 short, stout medial below, form a clearly monophyletic group charac- greapical bristles. Acrostichal bristles in 8 rows terized by other striking apomor1)hic characters. between anterior dorsocentrals; ~rescutellaracros- Other than the G species considered here, the tichal bristle and posterior intra-alar bristle tivice only described species of Sc1erocoelu.s are the wide- as long as anterior dorsocentral bristle; anterior spread, Nearctk, type spccies, S. sordipes (Adams), dorsocentral bristle slightly longer than acrostichal and the Ncot;ropical species S. plzi.~rsisela(Duda) setulae. Scutellum 1.3-1.5X as wide as long, four and S. regu,laris (Malloch). apical bristles 1.8-2.OX as long as scu tellum. Dorsal surface of mid tibia with 2 large anterodorsal bris- Sclerocoelus Marshall tles and 1 small posteroclorsal bristle proximally, Sclerococlrss Marshall, 1995: 283. Type species Lir~tosi~~aposterodorsal bristle midway between anterodor- sordipes Aclai~ls1904 sals; 2 anteroclorsal bristles elistally, the lower one twice as long as the upper; 1 very large posteroclor- 9 8 Volunle 11, No. 2, June, 1997, INSECTA MUNDI
sal distal bristle; 1 very small distal dorsal bristle at acters (18,19). The short, brown wings of gal- level of upper anterodorsal. Mid tibia of female upczger~sisare extremely similar to those of subbre- with a midventral bristle; ventral surface of male viper~r~is;however, shortening and darkening of mid tibia with a patch of short, stout bristles distal- wings is common in insular sphaerocericls and ly; mid tibia of both sexes with a strong apicoventral these characters are therefore considered especial- and weaker posteroventral and anteroventral api- ly subject to homoplasy. The third cladogram cal bristles. Miclfemur with short, stout anteroven- treatsgalapagerzsis as the sister group to the rest of tral and posteroventral bristles basally; mid basi- the genus, and links the Atlantic subbreuiper~r~isto tarsus with an enlargecl ventral bristle in basal mainland Neotropical and Caribbean species on the half. Wing with seconcl and third costal sectors basis of two characters of equivocal polarity (6,21). equal; R,,, sinuate, strongly curved up to costa. Character 21, the presence of a single stout apical Abdomen: Syntergite 1+2 with broad, short, bristle at the apex of the female cercus, is probably anterior pale area with a narrow posterior exten- a synapomorphy for the whole species group and is sion. Sinistral flange of synsternite 6+7 short, reversedin galapager~sisby the enlargement of the weakly developed; synst,ernite completely separat- inner preapical cercal bristle found in all members ed from sternite 8. Inner ventral corners of epan- of the species group. driurn widely separated, forming narrow, truncate The relationships between the remaining four lobes. Sternite 10 (subepandrial sclerite) com~lete species, all found in mainland South America and medially. Subcercus complex, with an outer (up- the Caribbean basin, are well supported. Sclerocoe- per) part usually projecting posteriorly into the lus brasilerlsis is the most plesiomorphic of the perianal pad and anteriorly to the posterior surst- mainlancl spccies, wit,h superficial similarities to ylar-cpanclrial junction, ancl an inner (lower) part the island endemics. Sclerocoelus hrasiler~sisis often with a long posterior arm; junction of inner treated as the sister group to the otther mainland and outer parts of subcercus with posterior bristles; and Caribbean species on the basis of six weak left ancl right subcerci meeting or almost meeting synapomorphies (7,9,13,14,20,22), tvhich are not near post,eromeclialpart of sternite 10 (subepandri- conflicted elsewhere on the cladogram. The mono- a1 sclerite). Distiphallus short, broad, dorsoven- phyly of a group containing the remaining three trally flattened, with broad, laterally-directed bas- species in the group (cariber~sis,ar~der~sis, Iternor- al sclerites, a basal ring, ancl a dist,al part compris- rhoidalis) is much more strongly supported. Rela- ing two curved lateral sclerites and a medial scler- tive to ot,her Sclerocoelus, members of this group ite which is usually distally bifid. Three spermath- seem to be characterized by progressive rccluction ecae, single one largest; s.ivollen apically with a of the complicate0 set of genital pouch sclerites shallow invagination; ducts short, inserted near (Character 1). Most members of the genus have an invagination; surface striate. apparently homologous series of sclerit,esincluding 4 or 5 separat,e sclerit,es bet,ween tjhe left side of Phylogeny of the Sclerocoell~sgalapagensis synsternit,e 6+7 and the spiracular sclerite on the species group right side of the abdomen. In the basal lineages of the galapcger~sisspecies group, 3 of these sclerites Analysis of a matrix of 22 characters, using are clearly visible, and 2 (sclerites C and D) are Hennig 86, yielclecl three equally parsimonious separate as in other Sclerocoelus. These sclerites cladograms (consistency index 82, reten tion index disappear in the highly distinctive Neotropical clacle 87, length 35) which differed only in the placement including cariher~sis,ar~der~sis, and hernorrhoida- of the two island endemics, the Atlantic subbreui- lis, and are replaced with a deep, evenly sclcrotized per~r~isancl the Pacific galapager~sis. I prefer the pouch. This group is also characterised by a num- cla(1ogram treating si~bbreviper~r~isas the sister ber of other unique synapomorphies of the male group to the rest of the genus, and placing gal- and female genitalia, and the monophyly of this apa,ger~sisas t,he sister group to the rest of the Netv clacle of three species seems unequivocal. World species (Fig. 1, table 1). The latter sister group relationship is supported by two unusual, Zoogeography of the Sclerocoelus independent characters (16,17) which probably rep- galapagensis species group resent good synapomorphies. An alternative cla- dogram treats the phenetically very similar island As is the case for the genus as a whole, members endemics as sister species based on two wing char- of the Sclerocoelus galapagensis group are almost INSECTA MUNDI, Vol. 11, No. 2, June, 1997
always associated with wet leaf litter in cloucl Membrane beside prosternum with short bris- forests or similar habitat. This might seem to sug- tles. Setaeofposteromedialpart of male skernite gest that these species are less likely to colonize 5 not flattened. Caribbean, Mexico, South and islands than are littoral species. It is possible, Central America, and Pacific...... 2 however, that association with wetlitterpreclispos- 2. Synsternite 6+7 of male broken into small, dark scler- es Sclerocoelus species to inclusion in flood debris itesposterior to sternite 5 (Fig. 34). Perianal pad washed out to sea after heavy rains. Peck (1994) with a narrowed ventral part articulating with found abundant fluvial debris in pleuston net sam- posterior arm ofouter part of subcercus (Fig. 14). ples taken between the Galapagos Islands during Posteroventral epandrial bristle shorter than the strong 1992 El Niiio event. The same pleuston epandrium (Fig. 16). Sternite 8 offemale strong- net samples included almost 4,000 arthropods. ly differentiated into almost verticallateral part,s Although none were identified as Sphaeroceridae, and concave ventral part (Fig. 58) ...... 3 transport in riverine flood debris is a plausible - Pocket-like genital pouch well developed and sternit,e 5 sometimes with 2 dark posteromedial lobes mechanism for the dispersal of cloucl forest species (Fig. 5), but no small, separate, dark sclerites to islands. Sclerocoelus sr~bbrevipe~u~isis the only behind posteromedial part of male sternite 5. endemic sphaerocerid in the remote Tristan da Perianal pad broad ventzally (Fig. 3). Poster- Cunha group, and S. g.nlq~cfgert.sisis one of a very oventral epandrial bristle longer than epanclri- few endemic sphaerocerids on the Galapagos is- um. Sternite 8 of female relatively simple, with- lands, so the group appears to be preaclapted to out prominent lateral parts ...... 4 overwater c1isl)ersal. Sclerocoelrrs szcbbrevi~)e~t,l~t:s probably originated from an eastern South Ameri- 3. Wing nlenlbrane brown, especially above and below can population of the common ancestor of the whole R,+,. Cerci of female dorsally concave, shining, ternlinating in 2 strong bristles (Fig. 38). Gal- group, subsequent to which the ancestral species of apagos...... the new rvorld clacle acquired synapomorphies 1G ...... Sclerocoelusgalapaget~sis,new species. and 17. Western South American populations of -Wing membrane clear. Cerciof female dorsally convex this species servetl as the source for the ancestor of and setulose, terminating in asingle short,&out Sclerocoel~rsg~o~l~z~~ctge~~sis,which is now restricted bristle (Fig. 20). Sout,h and Central America ... to wet Scalesia forest on t'he Galapagos Islands...... Sclerococl~~s Drasiler~sis, new species. Division of the mainland ancest,ral species into the eastern South American brasile~~sis,and a western 4. Area behind posteromedial part of male sternite 5 Neot,ropical-Caribbean clacle must have been the simple. Posterior arm ofsurstylus with 3-6 prom- inent, short, stout bristJes (Fig. 29) ...... 5 next occurrence in the zoogeogral~hichistory of the -Two prominent, bare, dark lobes arising behincl post,er- group. The ~vestcrnNeotropical-Caribbean clade omedial part of male sternite 5 (Fig. 5). Posterior then split int,o ccrrit)e~t,sisin the Caribbean basin, arm of surstyli~swith relatively snlall bristles. and an Andcan species anc~st,~alto nr~de~~sis and ...... Sclerocoelus ar~detrsis,new species. hemorrl~oidlalis. Totlay, cr.~t,del~sisis common in relatively unclisturbecl southern Andean cloud for- 5. Sternite 5of male short, shorter than genital pouch and ests, and has been collected in cloud forests as far halfas long as sternite 4, withposteromedial and north as Venezuela. Sclerocoelus h.ernorrhoidnlisis posterolateral areas of long setae (Fig. 24). Gen- known frorn relat,ively few specimens, mostly from ital pouch prominent, heavily sclerotized ...... Sclerocoel~tscariber~sis, new species. disturbed sites such as pasture and a riverbank in - Sternite 5 of Inale longer than genital pouch and an urban area near hiIerida, Venezuela. subequal in length to sternite 4, with two setose posterolneclial lobes; setosity of posterolateral Key to species of tlie Sclerocoell~s areasparse (Fig. 45). Genital pouch well tievel- galapcrgensis group oped, but relatively small and liglltly sclerotizeti 1. Posteroclorsal bristle of Iiatepistern~~mwell develogecl, ...... Sclerocoelr~sI~e~,~orrlt.oidalis, new species. reaching half way to \\ring base. Menlbrane be- sicleprost,ernum bare. Post,ero~rleclialpartofmale Sclerocoelus andensis, new species sternit,e 5 with flat,, scale-like bristles (Fig. 53). (Figs. 2- 12) Trist,an da Cunl~a...... Sclcrocoelus SLL~~)I.C~I~P~IL~L~S(Frey), new combi- Length 2.2-3.0mm; dark brown nation. Description: -Post.erodorsa1 kat,epist,ernal bristle fine, hair-lilie, reach- to black; legs, pleuron, and gena redclish-brown, ing at most one-third of the way to wing base. tarsi paler; interfrontal strips black, lower frons 100 Volume 11, No. 2, June, 1997, INSECTA MUNDI otherwise redclish; vibrissal angle reddish brown ly slightly expanded at point of contact with dis- like anterior gena and lateral parts of face, middle tiphallus. Distiphallus with distally expanded and part of face darker. Minimum interantennal dis- serrate laterally directed basal sclerites; central tance 1.7X width ofocellar triangle. Eye height 3.2- sclerite broad, distally pale and bifid. 3.5X genal height; anterior genal bristle as long as Female abdomen: Tergite 7 dark; tergite 8 genal height. divided into 2 large, dark lateral sclerites and an Prothorax with 3 small, bristle-bearing scler- indistinct, pale dorsal sclerite. Epiproct minutely ites in membrane beside prosternum. Icatepister- setulose, tapered between cerci; cerci tapered, with num with a short, fine posterodorsal bristle ancl a long, stout apical bristle and short stout dorsal minute anterodorsal setula. Male mid tibia strong- preapical bristle. Sternite 7 strongly convex pos- ly curved; ventral surface with a row of slightly teromedially. Main sclerit,e of sternite 8 trough- enlarged bristles on distal half, apical fifth covered like; narrow anteriorly, broaderposteriorly; anteri- with stout, curved bristles. Male mid femur with a or and lateral margins dark, middle part bare, pale row of 4 stout anteroventral and a patch of ca. 10 and concave; 2 small setae on sclerotized bases stout posteroventral bristles at base. Wing length present in membrane on each side. Hypoproct 2.5~width; membrane clear. projecting ventrad and densely long-setose on pos- Male abdomen: Tergite 5 with middle posteri- terior third. or bristles short but with 2 or 3 pairs of other Holotype(d ,CNC) and 43 Paratypes posterior marginal bristles 1.5X tergite length. (22d ,21 Q ,GUE,INESALT): Argentina. Salta. St,ernite 5 similar in size to sternite 4; uniformly 1500m, 40kmN Salta, Camino La Cornisa, sweep in dark brown except for short,, pale, striate portion roadside forest remnant, 27.ii. 1992, S.A. Marshall. normally telescoped over sternite 4; posteromeclial- Other paratypes: ARGENTINA. Salta. ly with a setulose patch from which arise 2 very 15kmW Chicoana, Canyada Gotera, 18.ii. 1992, clark, posteriorly directed lobes. Synsternite 6+7 sweep in forest remnant, S.A. Marshall almost ~ymmet~rical,right side (spiracular sclerite) (I ld,1 Q ,GUE);30kmE Campo Quijano, 18-28.1992, similar in size and shape to ventral part of left side, Flight intercept trap and sweep in forest remnant, middle part forming a continuous narrow band S.A. Marshall (ld,49, GUE); La Caldera, 1500m, ventral to a tleep, sclerotized central pouch. Ring forest sweep, 27.ii. 1992, S.A. Marshall sclerit,e well cleveloped, posterior to but separate (Gd ,9 Q ,GUE); 22kmN Lacaldera, El Ucumar, from spiracular sclerite. Epandrium with two malaise, subtropical humid forest, 2-30.xii. 1987, posterior bristles as long as epandrium ancl a 110s- S.&J. Peck (ld,CNC). BOLIVIA. Sta. Barbara, N. teroventral bristle twice as long as epandrium. Coroico, Yungas, 4-5.i. 1976, 1100m, L.E. Pena Length of dorsal part of epandrium greater than (4d ,3 9,CNC). VENEZUELA. Merida. Los Chor- height of anal fissure. Perianalpads strongly clevel- ros, 2100m, Flight intercept trap and sweep among oped, longer than epandrium, expancledventrally; leaf debris, 1-5.v.1988, S.A. Marshall anterior and dorsal parts well sclerotizetl but pos- (176,2Q,GUE,P\fIZA); Merida, 1500m, Calle Sta. teroventral part membranous, bulbous and set,u- Rosa trail, Flight intercept trap near spring, 24- lose. Pseutlocercus small, bearing 3 short brist,les; 30.iv. 1988, S.A. Marshall (8d,3Q ,GUE). connect,etl t,o epandrium by a pale band of tissue. Comnieiits: Sclerocoelus ar~der~sisis most Halves of sternite 10 (subepandrial sclerite) uni- easily separated from other Sclerocoelus on the formly narrow except at midclle where t,hey are basis of the 2 pairs of long posterior bristles on male slightly expanded and fused, middle part with tergite 5. This species is found in Andean cloud weak ventrolateral lobes. Subcercus with large forests from Venezuela south to northern Argenti- inner lobe, vent,rally expanded into posterior and na. In the northern part of its range it occurs anteriorlobes; outer part ofsubcercus narrow, with sympat,rically with the closely relatecl S. hernor- anterior arm only. Sur~t~yluslarge, twice as long as rl~oidalis. deep in lateral vic~v;anterior part with a cligitate ant,erotlorsal process, anter~vent~rallywith 3 inner Sclerocoelus brasilensis, new species bristles, a t,~ot,h-liIieapical process, and laterally (Figs. 13-23) directed ventral ridge; posterior part of surstylus long, tapered. Paramerevery broad basally, apical- Description: Length 2.4-2.Gmm; dark brown ly with a narrow, anteriorly hooked process. Ba- to black; legs ancl gena dark reddish-brown, t,arsi siphallus short, simple, posteriorly carinate; distal- paler; lower frons usually luteous only along nar- INSECTA MUNDI, Vol. 11, No. 2, June, 1997 101
row anterior margin; vibrissal angle reddish, con- deeper than posterior part, with 3 or 4 enlarged trasting in colour with the dark gena and face. bristles and a small flat, carina on inner surface; Minimum interantennal clistance 1.3X width of inner anterior corner with a narrow, sharply ta- ocellar triangle. Eye height 2.8-3.3X genal height; pered lobe. Paramere broad basally; distal half anterior genal bristle as long as genal height. narrow, strongly bent posteriorly; apex swollen, Prothorax with 2-3 small, bristle-bearing sclerites with a narrow, anteroventrally recurved part. Ba- in membrane beside prosternum. Icatepisternum siphallus simple, cylindrical, basally enlarged. Dis- with a short, fine posterodorsal bristle and a minute tiphallus short, dorsoventrally flattened, with lat- anterodorsal setula. Male mid tibia strongly curved; erally directed basal sclerites broad but pale and ventral surface with a row of slightly enlarged with only 2 or 3 serrations distally; central sclerite bristles on distal half, apical fifth covered with narrow, distally bifid. stout, curved bristles. Male mid femur with a row Female abdomen: Tergite 7 dark, shortened of 6 stout anteroventral and a patch of ca. 10 stout medially; tergite 8 divided into 2 elongate, dark posteroventral bristles at base, an teroventral bris- lateral sclerites and an indistinct, pale dorsal scler- tles stouter. Wing length 2.45 width; membrane ite. Epiproct setulose, tapered between cerci; cerci clear. tapered, with &out apical bristle and short stout Male abdomen: Tergite 5 with middle poste- dorsal preapical bristle. Sternite 7 short, posterior rior brist,les short but with 3 pairs of posterior margin appearing slightly concave because of pale bristles en. I. 1X tergite length and 1 pair of poste- posteromedial area. Sternite 8 almost egg-shaped, rolateral bristles ca. 1.3X tergite length. All stern- posterior margin weakly pigmented and appearing ites with a medial longitudinal pale stripe. Sternite concave, lateral margins folded under; 2 small 5 equal in wiclt,h and length to stcrnite 4; postero- setae on sclerotized bases present in membrane on medially with short, wicle, pale area, each side each side. Hypoproctprojectingventradancl densely bulging ancl with a pigmented, setose patch. Three long-setulose on posterior half, anterior part with small dark sclerites behind posteromedial part of medial and lateral bare, pigmented areas. sternite 5, of these t.he most sinistral (sclerite D) Holotype (d ,CNC) and 26 paratypes small and apparently fusecl with sternite 5, the (15d, 11Q ,GUE,MNR) paratypes BRAZIL. Para- other 2 (sclerites C and E) heavily sclerotized, with na. Curitiba, under fallen epiphytes near Natural dextrally curvecl, tapered, post,erior parts. Spirac- History Museum, 5.ii. 1990, S.A. Marshall. ular sclerite large, bilobed sinistrally, similar in Other paratypes: Bolivia. Coroico Yungas de size to remaincler of genital pouch; ring sclerite La Paz, 6-7.i.1976, 1100-1600m, L.E. Pena large, anterior edge fused to spiracular sclerite, (16 ,CNC). Brazil. Rio de Janeiro, Jacara Pagua, posterior edge t,hin. Right side of sternite 8 cleft; wet litter by stream, 22.i.90, S.A. Marshall anteroventral part contiguous with spiracularscler- (Gd, l4Q,GUE); Mury, near Nova Friburgo, 1000m, ite. Epandrium sparsely setose dorsally, more dense- swept off goat dung in wet area, 1l.iii. 1990, S.A. ly long-setose vent,rally, longest posterior bristles Marshall (1Q ,GUE); Siio Paulo, Jaragua, 8.ii. 1990, shorter than epandrium. Dorsal part of epandrium S.A. Marshall (3d,1 Q ,GUE); Parana, Londrina, slightly shorter than anal fissure. Perianal pads Parque Arthur Thomas, sweep, l.ii. 1990, S.A. Mar- flat, dark; t,aperecl ant1 bare ventrally, otherwise shall (3d,lO ,GUE); Curitiba, Flight intercept trap microsetulose; ventral lobe articulating with sub- in woods behind Natural History Museum cercus. Pseudocercus small, brown, free from epan- (ld,1 Q ,GUE); sweep of creek debris east of Curiti- drium, with 3 bristles, upper bristle twice as long as ba, 8.ii. 1990, S.A. Marshall (l6,GUE). ECUADOR. others andhalf as long as longest epandrial bristle. Pichincha prov., Rio Palenque Station, 47kmS Sto. Each half of sternite 10 (subepanclrial sclerite) Domingo, 29.iv-5.v. 1987, L.Coote and B. Brown, long, narrow, weakly arched; flattened, expanded 180m, Malaise trap, lowland rain forest (1Q ,GUE). and fused medially into a quadrate middle part. PANAMA. Barro Colorado Island, Canal Zone, Subcercus with inner part ventrally extended into 1.5.1929, C.H. Curran (ld,lQ ,AMNH). a distally expanded lobe but without prominent Comments: Males of this species are easily posterior process; outer part wit.h prominent ante- separated from other mainland species by the form rior and posterior arms, po~t~eriorarm articulating of the perianal pads and the separate sclerites with sclerotized part of perianal pad. Surstylus behindsternite 5. The name bra.sile~~sisreflects the twice as long as deep in lateral view; setulose on abundance of this species in southern Brasil. ventral half; posterior part flat; anterior part t,hick Volume 11, No. 2, June, 1997, INSECTA MUNDI
Sclerocoell~scaribensis, new species anterodorsal inner lobe, and a broad, carinate lobe (Figs. 24-33) on medial surface. Paramere short, broacl basally, sharply tapered on apical half. Basiphallus short, Description: Length 2.2-2.5mm, dark brown simple, compressed, partly contiguous with ventral to black, legs dark reddish-brown, lower frons usu- part of distiphallus. Distiphallus with broad, dis- ally red only lateral to interfrontal strips; vibrissal tally serrate, laterally directed basal sclerites; cen- angle reddish, contrasting with darker gena and t,ral sclerite narrow, cfist,ally pale and bifid. face. Minimum interantennal distance 1.5X width Female abdomen: Tergite 7 dark; tergite 8 of ocellar triangle. Eye height 3.5X genal height; divided into 2 large, dark lateral sclerites ancl an anterior genal bristle as long as genal height. indistinct, pale dorsal sclerite. Epiproct setulose, Prothorax with 2-4 small, bristle-bearing scler- truncate; cerci tapered, with long, stout apical ites in membrane beside prosternum. ICatepister- bristle and short stout dorsal preapical bristle. num with a short, fine posteroclorsal bristle and a Sternite 7 strongly convex posteromedially. Stern- minute anterodorsal setula. Male mid tibia strong- ite 8 narrow anteriorly, broadened and concave ly curved; ventral surface with a row of slightly posteriorly; 2 smallsetae on sclerotized bases present enlarged bristles on distal half, apical fifth covered in membrane on each side. Hypoproct projecting with stout,, curved bristles and with a large apicov- ventrad and densely long-setose on posterior half. entral bristle. Male mid femur with 2-3 stout an- Holotype (8,CNC) and 24 paratypes teroventral bristles and 6-8smaller posteroventral (loo", l4Q,GUE): DOMINICAN REPUBLIC. Near bristles at base. Wing length 2.4X width; mem- Cabo Rojo, 1300m, Las Abejas, cloud forest, brane clear. 19.i.1989, S.A. Marshall. Male abdomen: Tergite 5 with 1 pair of Other paratypes: Costa Rica. Mon teverde median posterior marginal bristles 1.5X tergite Reserve, 1500m, 15-20.viii.1986, L. Masner length, other posterior bristles 0.5X tergite length. (18d,15 Q ,CNC); Estrella Valley, Panclora, Sternite 5 0.5X as long as sternite 4; dark except l9.ii. 1984, Malaise trap, I-I. &A. Howden (I Q ,GUE); ant,erolateralcorners ancl 1)osteromcdial area; long- Cartago, 500m., Turrialba, Catie, 4.ix.1986, L. setose posterolaterally and ~ost~eromeclially.Area Masner (lo",GUE); San Jose, San Antonio de Esca- behind post,eromedial part ofsternite 5 occupied by zu, 25 Mar. to 9 April, 1984, Malaise trap, S.A. a deep sclerotized pouch (sclerite D), with a dark Cameron (SNOW, 1o", 19); Puntarenas, Monte sinistral lobe (sclerit,e C) and a spinulose pat,ch Verde, en. 1500m, lower montane wet forest, 4- (sclerite E) posterior to sac. Right side of sclerotized 1l.ii.89, D. Grimaldi (lo",AMNH). Cuba. Pinar del sac wit,h a posterior lip cont,iguous with spiracular Rio, Prov. Soroa, 27-28.iv.1983, W.N. Mat,his sclerit,e. Spiracular sclerite very large, well sclero- (1 Q ,USNM). Dominica, W.I. Trafalgar Falls, tized. Ring sclerite small, fused to posterolateral G.iii.1965, W.W. Wirtb (ld,USNM); Clarke Hall corner of spiracular sclerit,e. Epanclrium wilh dense, Est., 31.v,l.vi,2.vi,3.vi, 4.vi. 1966, G. Steyskal short bristles, posterior bristles longer and bristle (30",2Q,USNM); Manets Gutter, 5.iii. 1965, W.W. at posteroventral corner as long as epanclrium. Wirth (ld,2Q,USNM). El Salvaclor. Los Chorros, Length of dorsal part of epandrium grcat,er t,han 20.vi.1963, D.Q. Cavagnaro & M.E. Irwin height of anal fissure. Perianalpads st,rongly clevcl- (13d,CAS). Guatemala. Isabal, 300m, Las Escobas oped but shorter than epandrium, expanded ven- at light, 14-1G.xi.1986, M. Sharkey (1 ld,13 9 ,CNC). trally; anterior and clorsal parts well sclerotized bu t Jamaica. Hardwar Gap, 10.iii. 1970, stream mar- posteroventral part membranous, bulbous and set- gin, Wirth & Farr (lOd,3Q,USNM); Harclwar Gap, ulose. Pseudocercus small, weakly pigmented, free 25.vii. 1966, 4000t, Howden & Becker (1 Q ,CNC); from epandrium, with 3 small bristles. Halves of Runaway Bay, ii. 1969, W.W. Wirth, stream bed sternite 10 (subepandrial sclerite) dark, narrow; (46,2Q,USNM). MEXICO. Tepic, Nayarit, 3000t, contiguous medially, medial part with weak ven- viii. 1964, light trap, Blanton & Broce tral lateral lobes. Subcercus with a broad, elongate (3o",1 Q ,USNM); SLP, 2OkmW. Xilitia, lGOOm, 12.vi- inner lobe, posteroventrally extended into a point- G.viii, 1983, cloudforest flight intercept trap, S.&J. ed, prominent process; outer part of subcercus with Peck (lo" ,GUE); SLP, 40kmW. Xilitla, 12.vi- a short posterior arm and a longer ant,erior arm. 6.viii. 1983, 1700m, Pine-Oak forest, Flight inter- Surst,ylus large, wit,h a scoo1)-shapedposterior lobe cept trap, S.&J. Peck (1 Q ,GUE); Chiallas, Union with 4-6 short, stout posterior bristles; ant,erior Juarez, 4kmN., 22.iv.1983, W.N. Mathis part complex, with enlarged ventral bristles, an (1 8,USNM). Puerto Rico. Aguas Buenas Census, INSECTA MUNDI, Vol. 11, No. 2, June, 1997 103
Aguas Buenas Cave, 14.v.1973, 250m., S. Peck at posteroventral corner half as long as epandrium; (1 Q,BERI<).St. IGtts, W.I. Mt. Misery, 4-20.xii.1985, anteroventral bristles stout. Length of dorsal part 730m, pond/rainforest, L.D. Coote (ld,4Q,GUE). of epandrium subequal to height of anal fissure. Venezuela. Aragua, Rancho Grande, Parque Na- Perianal pad strongly developed, almost entirely cional Henry Pittier, 4.iii. 1967, flight trap, 1loom, sclerotized, shorter than epandrium, setulose, ven- M.E. Irwin (3d,CAS); Rancho Grande, Malaise tral part with a narrow lobe articulating with intercept, 2-30.xii. 1987, M. Sanborne (ld,GUE); subcercus. Pseudocercus small, weakly pigmented, Caripe, Guachero caves, 20.iv. 1986, oilbird refuse fused with corner of epandrium, with 3 bristles, the piles, S.A. Marshall (ld,MIZA). lower 2 very small, upper bristle as long as longest Other specimens examined (approx. 400,GUE): epandrial bristle. Halves of sternite 10 (subepan- Dominican Republic. Las Abejas Valley, near Cabo drial sclerite) dark, narrow; contiguous medially, Rojo; trail to Pico Duarte, near La Cienaga; Baraho- medial part deeply U-shaped and weakly lobate at na; Puerto Plata. middle. Subcercus with a broad, elongate inner Comments: This species is easily separated part; outer part of subcercus with a very broad from congeners by the escept,ionally distinctive anterior lobe ancl a quadrate posterior lobe articu- male postabdomen with spinose surstyli and a lating with sclerotized part of perianal pad; inner short sternite 5. The name ccrribel~sisreflects the and outer parts of subcercus broadly joined. Surst- great abundance of this species throughout the ylus large, with a scoop-shaped posterior lobe wit,h Caribbean region. small, densely packed posterior bristles; anterior part with four slightly enlarged ventral bristles on Sclerocoelus galapagensis, new species outer surface and a pair of stout ventral bristles on (Figs. 34-4 1) inner surface; anterior lobe of surstylus quadrate, flattened. Paramere relatively broad, with a deep Description: Length 2.0-2.2mm, dark brown anterior notch near base, apex boot-shaped. Ba- to black, legs and gena reddish-brown, lower frons siphallus with a short, quadrate epiphallus, distal extensively reddish; vibrissal angle concolorous part expanded clorsally. Distiphallus short and with reddish gena and face. hilinimum interanten- broad, with basal, laterally direct,ed sclerit,es blade- nal distance 2.OX width of ocellar triangle. Eye like, distally smooth; central sclerite broad, short. height 2.5-3.OX genal height; anterior genal bristle Female abdomen: Tergite 7 dark; tergite 8 shorter than genal height. divided into 2 large, dark lateral sclerites and a pale Prothorax with 1 or 2 small bristles in mem- dorsal sclerite. Epiproct bare except for 4 small brane beside pros t,ernum. I Comments: Sclerocoelusgalci~~ager~sisis high- meclially with weak ventral lobes. Subcercus with ly distinctive in both male and female genitalia, and a broad, elongate inner lobe, posterovent,rally ex- forms tlhe sist,er group t,o t,he rest of the Neotropical tended into a pointed, prominent process; outer members of the species group. It is named for its part of subcercus small, with an anterolateral arm restriction to the Galapagos islands. but not produced posteriorly. Surstylus large, with a scoop-shaped posterior lobe with 4-6 short, stout Sclerocoelr~sh,emorrh.oidalis, new species posterior bristles on the tapered apex; anterior part (Figs. 42-51) complex, with enlarged ventral, apically curled bristles, an anterodorsal inner lobe, and a broad, Description: Length 2.1-2.3mm, dark brown carinate lobe on medial surface. Paramere short, to black, legs retldish-brown, lower frons usually broatl basally, sharply tapered on apical half. Ba- redtlish only at corners; vibrissal angle reddish, siphallus short, simple, compressed, partly contig- contrasting with clark gena and face. Minimum uous withventral part of distiphallus. Distiphallus interantennal distance 1.5X width of ocellar trian- with laterally directed, basal, dist,ally serrate scler- gle. Eye height 3.5X genal height,; anterior genal ites; central sclerite thin, apically bifurcate. bristle as long as genal height. Prothorax with 3-5 Female abdomen: Tergite 7 dark; tergite 8 small, bristle-bearing scleritm in membrane beside tlivided into 2 large, tlark lateral sclerites and an prosternum, I Character matrix list, Sclerocoelus External characters galapagensis group 16) Membrane beside prosternum: 0, without bristles; 1, with bristles. Male terminalia 17) Posterior katepisternal bristle: 0, reaching half Sclerites C and D behind sternite 5: 0, present and way towing base; 1, minute, fine, reachingless than separate; 1, present and fused; 2, absent. one-third of distance to wing base. Deep, sclerotized pouch above posteromedial part of 18) Wing membrane: 0, clear; 1, brown. sternite 5: 0, absent; 1, present. 19) Wing length: 0, ca. 2.5X width; 1, ca. 2.0X width Surstylus with stout apical bristles on posterior lobe: (conspicuously shortened). 0, absent; 1, present. 20) Face and subvibrissal angle colour: 0, yellow brown; Surstylus with a narrow, finger-like anterodorsal 1, face dark brown to black and vibrissacontrasting- lobe: 0, absent; 1, present. ly pale. Surstylus with anterol~ledialcrest: 0, weak or ab- sent; 1, prominent. Female abdomen Posteroventral epandrial bristle: 0, short, like other 21) Cerci: 0, with two or more apical bristles; 1, termi- epandrial bristles; 1, longer than other epandrial nating in a single apical spine. bristles but shorter than epandrium; 2, prominent, 22) Sternite 8: 0, lateral parts more extensive than ven- longer than epandrium. tral part; 1, lateral parts smaller than ventral part; Large setulose area behind perianal pad: 0, absent; 2, lateral parts absent or folded under large concave 1, prominent. ventral part. Perianal pacl with finger-lilie posteroventral exten- sion: 0, present; 1, absent. Middle part of sternite 10: 0, medially rounded; 1, quadrate with lobes at corners. Inner (ventral) part of subcercus: 0, rounded, not projecting posteriorly; 1, with long, blade-like poste- rior lobe. Outer (upper) part of subcercus: 0, with posterior lobe articulating with apex of perianal pad; 1, with short posterior lobe not articulating with perianal pati; 2, without posterior lobe. Basiphallus: 0, broad basally and narrow distally; 1, short, laterally compressed; 2, with shortepiphallus anti distal expansion (unorderetl). Basal lateral arms of distiphalltts: 0, without teeth; 1, with 3 teeth; 2, with over 5 teeth. Mitldle part of tlistiphallus: 0, short and undevel- oped; 1, long and clistally bifurcate. Pseudocercus with three bristles: 0, upper bristle much longer than others; 1, all three bristles small. 108 Volume 11, No. 2, June, 1997, INSECTA MUNDI ++*A.Rii SG BCAH Map. Distribution of species in the Sclerocoelus gnlu~~nger~sisgroup. S, S. subbreviper~riis;G, S. gulnpagerisis; B, S. brasileri- sis; C, S. cnribetlsis; A, S. at~derlsis;H, S. hentorrhoidulis. INSECTA MUNDI, Vol. 11, No. 2, June, 1997 109 ancestor hemorrhoidalis Fig. 1. Phylogeny of the Sclerocoelus gc~lupcrgerrsisgroup. States marked with an asterisk are hornoplasies; numbers on right side are character states. 110 Volume 11, No. 2, June, 1997, INSECTA MUNDI Figs. 2-12. Sclcrocoelus crrtdettsis. 2, Male terminalia, ventral; 3, Male terminalia, posterior;$, Subcerci, surstyli, ancl subepnndrial sclerite,posteroventrnl; 5, Male sternites 5-7,ventral; 6, Male terminalia, left lateral; 7, Aedeagus and associatedstructures; 8, Subcerci, surstyli, ailcl subepandrial sclerite, left lateral; 9, Female terminalia, dorsal; 10, Female terminalia, ventral; 11, Female terminalia, lateral; 12, Sperinathecae. Abbreviations: pp, perinnal pad; sc, subcercus; sur, surstylus; sps, spiracular sclerite; s5, sternite 5; se, subepanclrial sclerite; pc, pseudocercus; 01, outer lobe of subcercus; il, inner lobe of subcercus; s6+7, synsternite 6+7; rs, ring sclerite; disti, distiphallus;pin, parainere; s7, sterilite 7. INSECTA MUNDI, Vol. 11, No. 2, June, 1997 11 1 Figs. 13-23. Sclerocoelus Drc~sile~rsis.13, Male terminalia, ventral; 14, Male terminalia, posterior; 15, Subcerci, right surstylus, and subepanclrial sclerite, posteroventral; 16, Male terminalia, left lateral; 17, Subcerci, surstyli, and subepnndrinl sclerite, left lateral; 18, Aedeagus and associated structures, lateral; 19, Terminalia, left dorsolateral, epandrium removed; 20, Female terminalia, dorsal;21, Female terminalia, ventral; 22, Female terminalia, lateral; 23, Spermathecae. Abbreviations: disti, distiphallus;pm, parameres; sc, subcercus; sur, surstylus; 8s, sternite 8. 112 Volume 11, No. 2, June, 1997, INSECTA MUNDI Figs. 24-33. Sclarocoelus curiber~sis.24, Male terminalia, ventral; 25, Male terminalia, posterior; 26, Subcerci, right surstylus, and subepnndrial sclerite, posteroventral; 27, Aedeagus and associated structures; 28, Male terminalia, left lateral; 29, Subcerci, surstyli, and subepandrial sclerite, left lnteral; 30, Spermnthecae; 31, Female terminalia, dorsal; 32, Female terminalia, ventral; 33, Female terminalia, lateral. INSECTA MUNDI, Vol. 11, No. 2, June, 1997 113 Figs. 34-41.Sclerocoelr~sgcrlctpcrger~sis. 34, Male termninalia, ventral; 35, Male termninalia, posterior; 36, Aedeagus and associated structures; 37, Male terminalia, left lateral; 38, Female terminalia, dorsal; 39, Female terminalia,ventral; 40, Spermnthecae; 41, Female terminalia, lateral. 114 Volume 11, No. 2, June, 1997, INSECTA MUNDI Figs. 42-51.Sclerocoel1~sIretnorrltoidalis. 42, Male terminalia, posterior; 43, Male terminalin, left lateral; 44, Aedeagus and associated structures; 45, Distiphnllus, ventml; 46, Male terminalia, ventral; 47, Subcerci, right surstylus, and subepandrinl sclerite, posteroventrnl; 48, Felnnle terminalia, dorsal; 49, Female terminalia, ventral; 50, Female terminalia, lateral; 51, Spermathecae. INSECTA MUNDI, Vol. 11, No. 2, June, 1997 115 Figs. 52-61. Sclerocoelr~ssr~bbreuipertrlis. 52, Male terminalia, posterior; 53, Male termninalia, left lateral; 54, Male sterilites 5-7; 55, Aedeagus ant1 associated structures; 5G, Male terininalia, ventral; 57, Subcerci, right surstylus, and subepanclrial sclerite, postero- ventral; 58, Felnale terminnlia, dorsal; 59, Female termninalia, ventral; GO, Female terminalia, lateral;Gl, Sperlnathecae. Abbrevintions: SG+7, sy~lsteriliteG+7; SC, sclerite C; SD, sclerite D; RS, ring sclerite; SPS, spiracular sclerite; RS, ring sclerite; S5, sternite 5.