the chloropidae (diptera) of the gal pagos islands,...

copy Insect Systematics amp Evolution (Group 6)

The Chloropidae (Diptera) of the Galaacutepagos IslandsEcuadorTERRY A WHEELER and JESSICA FORREST

Wheeler T A amp Forrest J The Chloropidae (Diptera) of the Galaacutepagos Islands EcuadorInsect Syst Evol 34 265-280 Copenhagen September 2003 ISSN 1399-560X

Thirteen species of Chloropidae are recorded from the Galaacutepagos Islands Ecuador Seven newspecies are described Diplotoxa loma sp n (subfamily Chloropinae) Conioscinella emphe-ria sp n Gaurax gethosyne sp n Hippelates alyscus sp n Liohippelates baptipalpis sp nOlcella anaclasta sp n Olcella lupina sp n (subfamily Oscinellinae) Another species in thegenus Apallates is apparently undescribed but there is insufficient information to justify a for-mal description Monochaetoscinella anonyma is recorded for the first time from the archi-pelago Four species previously recorded from the archipelago were also identified Cadremapallida Conioscinella galapagensis Elachiptera cultrata Liohippelates galapagensisPreviously published Galaacutepagos records of Liohippelates pusio apparently refer to L galapa-gensis A key to the Galaacutepagos species of Chloropidae is given Geographic affinities of theGalaacutepagos chloropid fauna are similar to those of other Diptera from the archipelago with fewpantropical species some species also found in the northern Neotropical and southernNearctic regions and endemic species apparently with Neotropical sister groups

T A Wheeler amp J Forrest Department of Natural Resource Sciences McGill UniversityMacdonald Campus Ste-Anne-de-Bellevue Quebec H9X 3V9 Canada (wheelernrsmcgillca)

Insect SystEvol

Introduction

Like many families of Diptera the Chloropidae ofthe Galaacutepagos Islands are incompletely knownOnly four species of Chloropidae have been previ-ously recorded from the archipelago the endemicspecies Conioscinella galapagensis (Curran1932) Liohippelates galapagensis (Curran 1932)and Elachiptera cultrata Wheeler amp Forrest 2002and the widespread species Cadrema pallida(Loew 1866) (Curran 1932 1934 Wheeler ampForrest 2002) Coquillettrsquos (1901) and Johnsonrsquos(1924) records of Liohippelates pusio (Loew1872) from the islands actually refer to L galapa-gensis Additional surveys of Galaacutepagos insects inthe 1960s especially by the California Academyof Sciences and an ongoing inventory of Galaacutep-agos arthropods since the 1980s by S B Peck andcolleagues (Peck 1996 Peck et al 1998) haveresulted in the collection of more than 1200 spec-imens of Chloropidae representing 13 species Inthis paper the chloropid fauna of the Galaacutepagos is

revised including the description of seven newspecies

Material and methods

Specimens examined are housed in the followinginstitutions (acronyms used in the text are inparentheses) Bernice P Bishop Museum Hono-lulu HI USA (BPBM) California Academy ofSciences San Francisco CA USA (CAS) Cana-dian National Collection of Insects Ottawa ONCanada (CNC) Lyman Entomological MuseumMcGill University Ste-Anne-de-Bellevue QCCanada (LEM) National Museum of NaturalHistory Washington DC USA (USNM) Zool-ogical Museum Oslo University Oslo Norway(ZMO)

Most specimens collected were preserved in70 ethanol and subsequently critical point driedusing liquid CO2 or hexamethyldisilazane Prepar-ations of genitalia were made by removing the

abdomens of mounted specimens (older air-driedspecimens were first relaxed in high humidity) andclearing them in 85 lactic acid heated in amicrowave oven for 1-2 30-second intervals sepa-rated by a cooling period Cleared abdomens werethen placed in glycerin for further dissection andexamination Dissected abdomens were stored inglycerin in plastic microvials pinned beneath thesource specimen

Key to species of Chloropidae of theGalaacutepagos Islands

1 Wing with costa extending to R4+5 crossveindm-cu absent Diplotoxa loma

- Wing with costa extending to M1+2 crossveindm-cu present 2

2 Arista broad sword shaped scutellum trape-zoidal body mostly yellowElachiptera cultrata

- Arista narrow rarely enlarged at base scutel-lum rounded posteriorly body colour variable 3

3 Hind tibia with apical or pre-apical ventral spurat least as long as tibial diameter 4

- Hind tibial spur absent or much shorter thantibial diameter 8

4 Thorax yellow in ground colour 5- Thorax black in ground colour 65 Hind tibial spur as long as basitarsus frontal

triangle shining scutum dorsally with palereddish stripes Cadrema pallida

- Hind tibial spur less than half as long as basi-tarsus frontal triangle dull scutum dorsally with black stripes Hippelates alyscus

6 Scutum dull eyes hairy hind tibial spur equal

in length to tibial diameter Apallates sp- Scutum shining eyes bare hind tibial spur

longer than tibial diameter 77 Legs yellow frontal triangle with sides straight

or nearly so extending slightly past middle offrons palp black basally yellow distally Liohippelates baptipalpis

- Legs partly brown hind femur and tibia dark-est frontal triangle with sides concave extend-ing almost to anterior margin of frons palpentirely yellow Liohippelates galapagensis

8 Vibrissal angle strongly produced proboscisgreatly elongate sclerotized geniculate (as inFig 17) 9

- Vibrissal angle rounded or only slightly pro-jecting proboscis not greatly elongate 10

9 Scutum with deeply incised longitudinal linesof punctures thoracic pleurites entirely pru-inose except for area between mid and hindcoxa Postgena very broad posteroventralangle less then 90deg (Fig 17) palp as long ashead many setae on frons and gena arisingfrom distinct punctures Olcella anaclasta

- Scutum without deeply incised lines of punc-tures thoracic pleurites with broad shiningstripe Postgena not as broad posteroventralangle greater than 90deg (Fig 20) palp at most07 times as long as head setae on head notarising from distinct punctures Olcella lupina

10 Frontal triangle shining notopleuron with 1anterior and 1 strong posterior bristle 11

- Frontal triangle dull notopleuron with 1 ante-rior and 2 strong posterior bristles 12

11 Frons with 1 outstanding fronto-orbital bristlescutellum black Monochaetoscinella anonyma

- Frons with several long fronto-orbital setaescutellum yellow Gaurax gethosyne

266 Wheeler T A amp Forrest J INSECT SYST EVOL 343 (2003)

1 2

epd

proj

sur

phap

hyp

cer

Figures 1-2 Diplotoxa loma (1) Male genitalia lateral (2) Male genitalia posterior Abbreviations cer - remnant ofcerci epd - epandrium hyp - hypandrium phap - phallapodeme proj - posteroventral projection sur - surstylusScale bar = 01mm

12 Male cercus short blunt not projecting (Fig3-4) surstylus parallel-sided in lateral view(Fig 3) Conioscinella empheria

- Male cercus projecting as triangular lobe (Fig5-6) surstylus broadest basally in lateral view(Fig 5) Conioscinella galapagensis

Subfamily Chloropinae

Diplotoxa loma Wheeler amp Forrest sp n

(Figs 1-2)

Description ndash Total length 14-20 mm Frons paleyellow roughly as long as broad slightly nar-rowed anteriorly frontal triangle pale yellowexcept for black ocellar tubercle subshining notclearly differentiated from rest of frons antennamostly yellow first flagellomere orbicular withdorsobasal black spot arista black pubescentthickened basally tapering apically occiputbrown medially face pale yellow eye with shortsparse ommatrichia long axis diagonal gena paleyellow 025 times height of eye higher posterior-ly vibrissal angle not projecting palp and pro-boscis short pale yellow Cephalic chaetotaxy 4reclinate to slightly lateroclinate fronto-orbitalsetae the posterior two longer and stronger sever-al moderately long medioclinate to proclinateinterfrontal setulae (some of these forming themargin of the frontal triangle) ocellar bristlesstrong proclinate divergent postocellar bristlesshort convergent inner and outer vertical bristleswell-developed the outer longer fine dark setulaepresent along occipital margin pale setulae pres-ent on lower margin of gena vibrissa long paleScutum pale yellow sparsely pruinose with fivebrown stripes median stripe on anteriortwo-thirds lateral stripes extending from post-pronotum to level of dorsocentral bristlessupraalar stripes extending from notopleuron tolevel of postalar bristle scutum approximately aslong as broad postpronotum yellow with brownspot scutellum pale yellow rounded broader thanlong thoracic pleurites pale yellow except forbrown spot around anterior spiracle brown lowerhalf of katepisternum black spot on meron anddark stripes on anterior margins of anepisternumand anepimeron Thoracic chaetotaxy 1 post-pronotal 1 weak anterior and 2 strong posteriornotopleural 1 dorsocentral 1 postalar 1 pair ofwidely separated apical scutellar and 1 shorter pairof lateral scutellar bristles postpronotum scutumand katepisternum setulose scutellum without

setulae Legs dark yellow Wing length 13-15mm second costal sector 10-11 times as long asthird costal sector cross-vein dm-cu absent Halterwhite Abdominal tergites pale brown setuloseand shining

Male postabdomen (Fig 1-2) Terminalia most-ly pale yellow subshining surstylus fused withepandrium surstylus short triangular in lateralview with several setae and setulae on posteriorhalf posteroventral margin of epandrium withshort dark projection (see Remarks) fused cerci(mesolobus) desclerotized present only as 3-4short stout setae

Type material ndash Holotype = ECUADORGALAacutePAGOS ISLANDS SANTA CRUZ 4km NBellavista MediaLuna 620m 14v-13vii1985 S amp JPeck Miconia zone FIT (CNC) Paratypes 28= 73Rsame data as holotype (CNC LEM)

Etymology ndash The species name is from the Spanish loma(hill) referring to the high elevation of the type locality

Remarks ndash This species would key to ElliponeuraLoew 1869 in existing keys to chloropid generabased on the lack of crossvein dm-cu The pres-ence or absence of this crossvein however isweak justification for recognition of a separategenus and the distinction between ElliponeuraPseudopachychaeta Strobl 1902 and DiplotoxaLoew 1863 is so tenuous that Spencer (1986)combined all three under Diplotoxa A compre-hensive systematic revision of this group ofldquogenerardquo is badly needed but for the present wefollow Spencer and recognize only the genusDiplotoxa Sabrosky amp Paganellirsquos (1984) recordof an undescribed species of Elliponeura from theGalaacutepagos Islands probably refers to this species

The male cerci of Chloropinae are usually fusedinto a single median structure (mesolobus)Spencer (1977) divided the New Zealand speciesof Diplotoxa sl into three groups based on varia-tion in the male cerci Group I with a typicalchloropine mesolobus Group II with the cerci sep-arate (as in most Oscinellinae) and conspicuouslylong and dark and Group III with the cerci absentDiplotoxa loma would belong to Spencerrsquos GroupII based on the separate dark projections (Fig 2)However we suggest that the dark projections ofSpencerrsquos Group II species are not the true cercibut instead are novel structures derived from theposteroventral epandrial margin andor thesubepandrial sclerite In D loma three or fourshort setae arising from sockets are visible at highmagnification in the same location as the

INSECT SYST EVOL 343 (2003) Chloropidae of the Galaacutepagos Is 267

mesolobus in most Chloropinae These setaeapparently mark the location of a secondarilydesclerotized mesolobus in D loma and possiblythe New Zealand species of Groups II and III(although we have not examined those species)

Given what is known of the biology of otherspecies of Diplotoxa sl in the New World Dloma is probably phytophagous in the flowerheads of sedges (Cyperaceae)

Subfamily Oscinellinae

Apallates sp

Material examined ndash ECUADOR GALAacutePAGOSISLANDS SANTA CRUZ 1= 1km E Charles DarwinResearch Station lagoon littoral 30iii-7iv1989malaise Peck amp Sinclair (CNC) 2= 1R Academy BayDarwin Research Station sweeping coastal plants25i1964 D Q Cavagnaro amp R O Schuster (CAS)

Remarks ndash This is the first record of the genusApallates Sabrosky 1980 from the GalaacutepagosAlthough it is apparently undescribed this speciesis similar to a number of described but poorly dif-ferentiated Neotropical species and we cannot besure of its status without consideration of severaladditional species Accordingly we prefer to leavethis species undescribed pending a complete revi-sion of the Neotropical Apallates

Cadrema pallida (Loew 1866)

Hippelates pallida Loew 1866 184Prohippelates pallidus Curran 1934 159

Material examined ndash ECUADOR GALAacutePAGOSISLANDS BALTRA [as S Seymour I] 1= 6R11vi1932 M Willows Jr Templeton Crocker Exped1932 (CAS) 1= 2R same data except 11vii1932(CAS) ISABELA 1R Pto Villamil 7iii1989 inter-tidal rocks B J Sinclair (CNC) SANTA CRUZ 1=3R Tortuga Bay littoral 29i1989 B J Sinclair MVlights (CNC) 1= 4R Darwin Research Station 0m lit-toral zone 28iv1991 J Heraty (CNC) 1R CharlesDarwin Research Station arid zone 16i1989 MVlights B J Sinclair (CNC) 1R Academy Bay DarwinResearch Station sweeping coastal plants 25i1964 DQ Cavagnaro amp R O Schuster (CAS) 1R AcademyBay Darwin Research Station 26i1964 R O Schuster(CAS) 1= Academy Bay at light 17ii1964 P DAshlock (BPBM)

Remarks ndash Although Curran (1934) recorded Cpallida from the Galaacutepagos Sabrosky amp Paganelli(1984) did not list the species from the islands nordid they cite Curran (1934) This species ispantropical with records from North Central and

South America Australia Africa and islands in thePacific Atlantic and Indian oceans The 11 July1932 record from Baltra is most likely an error onthe specimen label the correct date is probably 11June 1932 (as for other specimens from the samelocality) Nartshuk (2002) illustrated the male gen-italia of C pallida

Conioscinella empheriaWheeler amp Forrest sp n

(Figs 3-4)

Description (male only) ndash Total length 16-21mm Frons mainly yellow sometimes darker(grey-brown) posteriorly frontal triangle grey04-05 times length of frons frons includingfrontal triangle dull antenna mainly yellow firstflagellomere reniform darkened on dorsal half ormore arista short black occiput grey-brown faceyellow to pale brown with carina eye with shortdense ommatrichia gena yellow on anteriortwo-thirds or more grey-brown posteriorly 03times height of eye vibrissal angle projectingslightly in most specimens palp yellow proboscisbrown sclerotized geniculate Cephalic chaeto-taxy 7-8 short reclinate fronto-orbital setae sev-eral proclinate to medioclinate interfrontal setulaeocellar and postocellar bristles short cruciateinner and outer vertical bristles slightly longerthan surrounding setulae setulae along occipitalmargin and on ventral one-third to one-half ofgena vibrissa slightly longer than subvibrissalsetulae Scutum and scutellum heavily pruinosedark grey-brown scutum longer than broad post-pronotum and notopleuron dull usually paler thanscutum scutellum rounded apically broader thanlong thoracic pleurites variable in colour rangingfrom yellow to dark brown posterodorsal half ofanepisternum anterodorsal half of anepimeronand ventral corner of katepisternum at least sparse-ly pruinose Thoracic chaetotaxy 1 (occasionally2) postpronotal 1 anterior and 2 posterior noto-pleural 1 dorsocentral 1 postalar and 1 intrapos-talar bristles scutellum with 1 pair of convergentapical and 1 or 2 pairs of subapical bristles post-pronotum scutum and scutellum densely and uni-formly setulose katepisternum with a few finesetulae Legs yellow or variably darkened withbrown (see Remarks) Wing length 13-18 mmsecond costal sector 15-175 times length ofthird costal sector dm-cu oblique not parallel


with r-m Halter white Abdominal tergites brownsetulose subshining sternites paler brown setu-lose

Male postabdomen (Fig 3-4) Terminalia yel-low to pale brown with short dense pubescenceventral margin of epandrium almost straightbetween surstylus and cercus in lateral viewsurstylus parallel-sided evenly curved in lateral

view cercus short blunt with a single long setasituated far from median cleftType material ndash Holotype = ECUADORGALAacutePAGOS ISLANDS ESPANtildeOLA BahiacuteaManzanilla 5-10vi85 Samp J Peck littoral cryptocar-pus amp Prosopis FIT malaise (CNC) ParatypesBARTOLOMEacute 1= lava bed in contact with mangroveswamp N end of island 5rsquo alt 3ii1967 I L Wiggins(CAS) ESPANtildeOLA 14= Bahiacutea Manzanilla 5-


basph

distph

3 4

5 6

Figures 3-4 Conioscinella empheria (3) Male genitalia lateral (4) Male genitalia posterior Figures 5-6Conioscinella galapagensis (5) Male genitalia lateral (6) Male genitalia posterior Abbreviations basph - basiphal-lus distph - distiphallus Scale bars = 01mm

10vi1985 S amp J Peck littoral cryptocarpus ampProsopis FIT malaise (CNC LEM) 8= same dataexcept Prosopis grove behind beach carrion traps(CNC) 1= Punta Juarez 10-12ii1967 I L Wigginstrap among Lycium minimum Prosopis doleis andCryptocarpus pyriformus (Gubarsbia) (CAS) FLORE-ANA 2= littoral zone 23iii1989 sweeping white-sand beach B J Sinclair (CNC) 1= Black Beach 21-28iii1989 10m littoral-arid FIT Peck amp Sinclair(CNC) GENOVESA 3= S side of island 200 ydsfrom beach 4-6ii1967 I L Wiggins in flight trapamong Bursera graveolens (CAS) ISABELA 3=Villamil 1km W 2-15iii1989 FIT littoral scrub onsand 1m Peck amp Sinclair (CNC) SAN CRISTOacuteBAL43= Puerto Baquerizo south beach 21ii1989 marineiguana nesting sites littoral B J Sinclair (CNC LEM)SANTA CRUZ 1= 1km E Charles Darwin ResearchStation (lagoon) littoral 30iii-7iv1989 malaise Peckamp Sinclair (CNC) SANTA FE 5= 4iv1989 B JSinclair sweeping sea lion beach (CNC) 1= littoralzone yellow pan trap 4-6ii1989 B J Sinclair (CNC)2= 3-6vi1989 B J Sinclair sea lion dung trap in lit-toral zone (CNC) 1= littoral beach malaise 4-6iv1989 Peck amp Sinclair (CNC)

Etymology ndash The species name is from the Greekempheres (resembling like) referring to the great simi-larity between this species and the sympatric C galapa-gensis

Remarks ndash More than 280 female specimens ofConioscinella Duda 1929 from 14 islands (BaltraBartolomeacute Espantildeola Floreana Genovesa Isa-bela Marchena Pinta Raacutebida San CristoacutebalSanta Cruz Santa Fe Seymour Wolf) were notincluded in the list of paratypes or specimens exa-mined because the two species of Conioscinellarecorded from the Galaacutepagos can only be distin-guished on the basis of male genitalic charactersThere is considerable variation in leg colour with-in C empheria in which the legs vary from yellowto mostly dark brown The darkest specimens of Cempheria were taken from a mixed series of bothspecies on San Cristoacutebal where they were clearlydistinguishable from the yellow-legged C galapa-gensis In series from other islands comprisingonly C empheria most specimens had bright yel-low legs similar to those of C galapagensis Thepossible occurrence of character displacement insympatric populations of these two species wouldbe worthy of further study

Conioscinella galapagensis (Curran 1932) (Figs 5-6)

Oscinella galapagensis Curran 1932 357Conioscinella galapagensis Sabrosky amp Paganelli

1984 10

Redescription (male only) ndash As for C empheria

except as follows total length 14-18 mm anepis-ternum anepimeron katepisternum and meronalways brown legs usually entirely yellow (seeRemarks under C empheria)

Male postabdomen (Figs 5-6) ventral margin ofepandrium tuberculate between surstylus and cer-cus in lateral view surstylus broadest in basal halfnot curving in lateral view cercus projecting astriangular lobe with long seta at distal end

Type material examined ndash Holotype R ECUADORGALAacutePAGOS ISLANDS FLOREANA Post OfficeBay Seaside 12-14xi1925 A Wollebaek (ZMO)Paratypes 4R same data as holotype (ZMO)

Other material examined ndash ECUADOR GALAacutePAGOSISLANDS DARWIN 1= 29i1964 D Q Cavagnaro(CAS) ESPANtildeOLA 2= Punta Juarez 10-12ii1967 IL Wiggins trap among Lycium minimum Prosopisdoleis and Cryptocarpus pyriformus (Gubarsbia) (CAS)FLOREANA 11= several hundred R Post Office Bay8xii1925 A Wollebaek (ZMO) ISABELA 19=Puerto Villamil coastal pools 7iii1989 brackish B JSinclair (CNC LEM) 4= Puerto Villamil arid zone5iii1989 5m MV-lights B J Sinclair (LEM) PINTA6= littoral zone 2m carrion traps 14-20iii1992 SPeck (LEM) SAN CRISTOacuteBAL 10= PuertoBaquerizo south beach 21ii1989 marine iguana nest-ing sites littoral B J Sinclair (CNC) 1= PuertoBaquerizo arid zone 20ii1989 sweeping B JSinclair (CNC) SANTA FE 1= 4iv1989 B JSinclair sweeping sea lion beach (CNC) WOLF 4=1ii1964 D Q Cavagnaro (CAS) 1= 31i-01ii1964P D Ashlock (BPBM)

Remarks ndash Although the holotype and paratypesare all females there is a long series of specimensin the ZMO collected at the same time and locali-ty All male specimens in that series are conspecif-ic and our definition of this species is based onthose specimens See Remarks under C empheriafor notes on the difficulty in identifying femalespecimens

Elachiptera cultrata Wheeler amp Forrest 2002Elachiptera cultrata Wheeler amp Forrest 2002 2

Remarks ndash This species can easily be distin-guished from all other recorded Galaacutepagoschloropids on the basis of the broad flattenedarista the mainly yellow body and the trapezoidalscutellum Wheeler amp Forrest (2002) providedcharacters for distinguishing E cultrata fromother species of Elachiptera Macquart 1835

This species apparently belongs to a New Worldgroup of yellow or reddish species of ElachipteraMany published records of species in this groupare from low-lying habitats often coastal or other-


wise arid The habitats from which E cultrata wascollected were more diverse although primarilyhumid ranging from the Arid Zone near the coastthrough the Transition Scalesia Miconia andPampa Zones up to elevations of 1000m Spe-cimens were collected in native habitats as well asdisturbed agricultural areas and guava forestComplete label data were provided by Wheeler ampForrest (2002) and we have since examined twoadditional specimens (SANTA CRUZ 1= BellaVista 10km N Academy Bay 21ii1964 P DAshlock (BPBM) 1R 8km N Academy Bay aban-doned garden 24i1964 P D Ashlock (BPBM))

Despite its apparently broad habitat preferenceE cultrata was primarily collected (117 of 121specimens) on two large islands (Santa CruzIsabela) with long-term permanent human habita-tion and extensive habitat disturbance Only fourspecimens were collected from other islands(Floreana Seymour) in the course of a large scalemulti-year survey of Galaacutepagos insects Thisapparent association with human-altered islandsand absence from undisturbed islands may indi-cate that the species has been introduced fromelsewhere probably western South America orCentral America by human activity Finding Ecultrata in Central or South America would lendsupport to that possibility

Gaurax gethosyne Wheeler amp Forrest sp n

(Figs 7-8)

Description ndash Total length 17-23 mm Fronsshining yellowish brown anteriorly darker brownposteriorly frontal triangle including ocellartubercle polished brown not clearly delimitedanteriorly first flagellomere reniform entirelyyellow in male black at apex in female arista withbasal segments yellow terminal segment darkocciput dark brown pruinose face yellow withoutcarina eye apparently bare with only scatteredshort setulae iridescent long axis diagonal genamostly shining brown with black margin 008times height of eye vibrissal angle and parafacialshining yellow vibrissal angle not projecting palpblack proboscis short with brown sclerotizationCephalic chaetotaxy 5 reclinate fronto-orbitalsetae increasing in length posteriorly single rowof setulae on border of frontal triangle 1 pair oflonger cruciate interfrontal setulae near anteriormargin of frons ocellar and postocellar bristlesquite long erect cruciate inner and outer verticalbristles long the outer stronger short setulaealong occipital margin longer setulae along genalmargin the posterior 2 hairs notably longer vib-rissa slightly longer than subvibrissal setulaeThorax entirely polished brown except for pale


7 8

Figures 7-8 Gaurax gethosyne (7) Male postabdominal segments 6-8 and genitalia lateral (8) Male genitalia pos-terior Scale bar = 01mm

yellow scutellum and slight pruinosity aroundwing base scutum approximately as long asbroad with faint pattern of lighter brown postsu-tural dorsocentral and narrower supraalar stripesmerged posteriorly scutellum rounded apicallybroader than long thoracic pleurites lighter brownthan postpronotum and scutum Thoracic chaeto-taxy 1 postpronotal 1 anterior and 1 posteriornotopleural 1 dorsocentral 1 postalar 1 pair ofapical and 1 pair of shorter subapical scutellarbristles 1 weak katepisternal bristle postprono-tum scutum and katepisternum with long setulaescutellum microsetulose along margin and lateral-ly at base Fore leg (including coxa) yellow tarsusdarker mid and hind leg with coxa and basal thirdto half of femur yellow rest of leg brown to blackWing length 15-20 mm second costal sector 12-14 times length of third costal sector cell r1

broad but less so than in most Nearctic species ofGaurax Halter white Abdominal tergites palebrown syntergite 1+2 very pale medially tergitesevenly microsetulose and with scattered longersetulae sternites very pale with long setulae

Male postabdomen (Fig 7-8) Sternite 6 appar-ently present forming a complete ventral bandfused laterally with synsternite 7+8 ventral medi-al region of sternite 6 projecting ventrally as a pale

ventral processes anterior to epandriumEpandrium not greatly enlarged relative toabdomen epandrium dark brown with dense longhairs Surstylus roughly rectangular in posteriorview pointed at ventral medial corner with addi-tional tooth-like projection on posterior surfacePhallus long cylindrical membranous Cercuslong black well-sclerotized

Type material ndash Holotype = ECUADORGALAacutePAGOS ISLANDS ISABELA Sto Tomas 4-15iii1989 300m humid forest Malaise Peck ampSinclair (CNC) Paratypes SANTA CRUZ 3= 1RAcademy Bay Darwin Research Station sweepingcoastal plants 25i1964 D Q Cavagnaro amp R OSchuster (CAS)

Etymology ndash The species name is a noun in appositionderived from the Greek gethosyne (delight joy) refer-ring to the attractive appearance of this fly and the excel-lent condition of the holotype

Remarks ndash Although most Nearctic species ofGaurax Loew 1863 are distinguished by the pos-session of an enlarged epandrium the male geni-talia of this species are not greatly enlarged Thereis a range of development in the size of the epan-drium among other undescribed Neotropicalspecies of Gaurax Other than the size of the epan-drium the genitalia are typical of New World


9 10

Figures 9-10 Hippelates alyscus (9) Male genitalia lateral (10) Male genitalia posterior Scale bar = 01mm

species of Gaurax Cell r1 is also narrower than inmost Nearctic species of Gaurax but similar to theNeotropical species

Hippelates alyscus Wheeler amp Forrest sp n

(Figs 9-10)

Description ndash Total length 16 mm Frons yellowdull slightly longer than broad frontal triangleincluding ocellar tubercle black sparsely pruin-ose about 05 times length of frons antenna yel-low at least ventrally variably darkened dorsallyfirst flagellomere suborbicular arista dark slen-der occiput yellow variably infuscated face yel-low without carina eye densely short-haired genayellow 02 times height of eye vibrissal angle notprojecting palp and proboscis yellow proboscisshort mostly unsclerotized Cephalic chaetotaxy5-7 short reclinate fronto-orbital setae numerousshort interfrontal setulae on anterior of frons andforming single row along border of frontal trian-gle ocellar bristles short erect convergent posto-cellar bristles longer convergent to cruciate innerand outer vertical bristles equal in length to posto-cellars setulae along occipital margin and ventralhalf of gena vibrissa not differentiated from sub-vibrissal setulae Scutum yellow pruinose withfive black stripes broad median stripe on anteriortwo-thirds to three-fourths lateral stripes extend-ing from postpronotum to level just beyond end ofmedian stripe narrow paler supraalar stripes ex-tending from notopleuron to level of postalar bris-tle scutum as long as broad postpronotum andscutellum yellow pruinose scutellum rounded api-cally broader than long thoracic pleurites pruin-ose on dorsal half shining below yellow exceptfor large ventral black spot on katepisternum darkanterior margins of anepisternum and anepimeronand black posterior margin of meron Thoracicchaetotaxy 1-2 postpronotal 1 anterior and 2posterior notopleural 1 dorsocentral 1 postalar 1intrapostalar 1 pair of long and strong apical and2 pairs of shorter weaker subapical scutellar bris-tles postpronotum scutum scutellum andkatepist-ernum setulose Legs mostly yellow tarsi and dis-tal portions of tibiae variably darkened hind tibiawith curved pre-apical ventral spur approximatelyequal in length to diameter of tibia Wing length13-15 mm second costal sector 15-18 timeslength of third costal sector Halter white Abdo-minal tergites brown moderately long-haired ster-nites paler setulose

Male postabdomen (Fig 9-10) Epandriumbrown with long hairs on posterior surface sur-stylus broad black with anterior fringe of long se-tae cercus bilobed with 1 long seta and severalshort setae

Type material ndash Holotype = ECUADOR GALAacute-PAGOS ISLANDS FERNANDINA Cabo Hammond0m 3v1991 J Heraty beach grasses (CNC) Para-types ISABELA 1= 1R arid zone 5iii1989 B JSinclair sweeping (CNC)

Etymology ndash The species name is from the Greek alysko(wander uneasily) referring to the combination of char-acters that left this species suspended between at leastthree poorly defined genera for much of this study

Remarks ndash This species belongs to a group ofspecies that have been assigned by some authors tothe genus Olcanabates Enderlein 1911 AlthoughPaganelli amp Sabrosky (1993) treated the two gen-era as synonyms this was not based on phyloge-netic analysis and further study of the generic lim-its is required For the present we prefer to recog-nize a broader concept of Hippelates Loew 1863

Liohippelates baptipalpisWheeler amp Forrest sp n

(Figs 11-12)

Description ndash Total length 18-23 mm Fronsdark yellow anteriorly grading to dark brown orblack posteriorly subshining usually with somepruinosity bordering frontal triangle frontal trian-gle including ocellar tubercle polished darkbrown to black with sides straight or slightly con-cave 06-07 times length of frons antenna yel-low first flagellomere suborbicular usually slight-ly infuscated dorsally arista pale basally dark-ened distally occiput black pruinose face darkyellow with weak carina eye bare gena 02 timesheight of eye yellow to black shining microscop-ically striate with darker polished margin vib-rissal angle not projecting palp black basally yel-low distally proboscis sclerotized brown to blackgeniculate but not elongate Cephalic chaetotaxy1-2 short weak anterior and 4-5 longer strongerposterior reclinate fronto-orbital setae severalproclinate to medioclinate interfrontal setulae onanterior of frons and forming single row of setulaealong border of frontal triangle ocellar and posto-cellar bristles erect cruciate inner vertical bristlesapproximately equal in length to postocellarsouter vertical bristles slightly longer and strongersetulae along occipital and genal margins vibrissa


approximately twice as long as subvibrissal setu-lae Thorax entirely shining black except for pru-inosity around wing base and base of scutellumscutum slightly longer than broad scutellumrugose slightly broader than long subconicalThoracic chaetotaxy 1-3 weak postpronotal 1weak erect intrahumeral 1 anterior and 2 posteri-or notopleural 1 dorsocentral 1 postalar and 1

weak intrapostalar bristles (the latter may be indis-tinguishable from surrounding setulae) scutellumwith 1 pair of cruciate apical and 2 or more pairsof much shorter subapical bristles all borne onshort tubercles postpronotum scutum scutellumand katepisternum setulose setulae in median anddorsocentral rows of scutum slightly divergent3-4 rows of setulae between median and dorsocen-


11 12

13 14

Figures 11-12 Liohippelates baptipalpis (11) Male genitalia lateral (12) Male genitalia posterior Figures 13-14Liohippelates galapagensis (13) Male genitalia lateral (14) Male genitalia posterior Scale bars = 01mm

tral rows Legs pale yellow hind tibia with strong-ly curved preapical ventral spur longer than diam-eter of tibia Wing length 16-21 mm secondcostal sector 17-19 times length of third costalsector Halter white Abdominal tergites sparselypruinose setulose syntergite 1+2 mostly pale yel-low remaining tergites and sometimes posterolat-eral corners of syntergite 1+2 brown sternitesvery pale setulose

Male postabdomen (Figs 11-12) Epandriumlong-haired surstylus curved posteromediallytapering distally with several long setae on anteri-or half cercus elongate projecting posteroventral-ly cerci convergent in posterior view each withone long seta and several short setae phallus shortmembranous

Type material ndash Holotype = ECUADOR GALAacute-PAGOS ISLANDS SANTA CRUZ 1km E CharlesDarwin Research Station lagoon littoral 30iii-7iv1989 malaise Peck amp Sinclair (CNC) ParatypesBARTOLOMEacute 1R lava bed in contact with mangroveswamp N end of island 5rsquo alt 3ii1967 I L Wiggins(CAS) ISABELA 1= 10R Puerto Villamil 7iii1989intertidal rocks B J Sinclair (CNC USNM) SANTACRUZ 2= 9R 1km E Charles Darwin Research Stationlagoon littoral 30iii-7iv1989 malaise Peck amp Sin-clair (CNC LEM USNM) 1= 1R Academy BayECCD 30m arid zone thornscrub malaise-FIT 10v-14vii1985 S amp J Peck (CNC) 2R Charles DarwinResearch Station littoral zone 4-9ii1989 dung trap BJ Sinclair (CNC) 1R Academy Bay Darwin ResearchStation sweeping coastal plants 25i1964 D Q Ca-vagnaro amp R O Schuster (CAS) 1R Academy BayDarwin Research Station 7ii1964 R O Schuster(CAS)

Etymology ndash The species name is derived from theGreek bapto (dip dye) and palpis referring to the yel-low distal portion of the palps

Remarks ndashThis species may be distinguished fromthe externally similar Liohippelates collusor(Townsend 1895) by the bicoloured palps (all yel-low in L collusor) presence of six small but dis-tinct tubercles on the scutellum (only very smallapical tubercles in L collusor) and usually moreextensive yellow on the frons (only anterior mar-gin yellow in L collusor) Because of the similar-ity between the two species recent records of Lcollusor from other Pacific islands such as Hawaii(Beardsley amp al 1999) should be treated with cau-tion

Liohippelates galapagensis (Curran 1932)

(Figs 13-14)

Hippelates galapagensis Curran 1932 356

Liohipellates galapagensis Sabrosky amp Paganelli1984 21

Liohippelates pusio of Coquillett 1901

Redescription ndash Total length 12-24 mm Fronssubshining mainly dark yellow grading to brownat vertex frontal triangle including ocellar tuber-cle polished dark brown to black with sides dis-tinctly concave extending in narrow point almostto anterior margin of frons antenna mostly yel-low first flagellomere suborbicular infuscated ondorsal half arista brown palest basally occiputblack pruinose face yellow with weak carina eyebare gena 014-019 times height of eye yellowshining microscopically striate with polishedbrown margin vibrissal angle not projecting palpyellow proboscis sclerotized brown geniculatebut not elongate Cephalic chaetotaxy 1 to severalshort weak anterior and 4-5 longer stronger pos-terior reclinate fronto-orbital setae several procli-nate to medioclinate interfrontal setulae on anteri-or of frons and forming single row of setulae alongborder of frontal triangle ocellar bristles shortreclinate convergent postocellar bristles shorterect convergent to cruciate inner vertical bristlesonly slightly stronger than surrounding setulaeouter vertical bristles approximately equal inlength to postocellars setulae along occipital andgenal margins vibrissa approximately 3 times aslong as subvibrissal setulae Thorax entirely shin-ing black except around wing base and posteriormargin of scutum scutum slightly longer thanbroad scutellum somewhat rugose rounded api-cally slightly broader than long Thoracic chaeto-taxy 1 weak postpronotal 1 anterior and 2 poste-rior notopleural 1 dorsocentral 1 postalar and 1weak intrapostalar bristles (the latter may be indis-tinguishable from surrounding setulae) scutellumwith 1 pair of cruciate apical bristles borne onshort tubercles and 2 pairs of short subapical bris-tles postpronotum scutum scutellum and kate-pisternum setulose setulae in median and dorso-central rows of scutum slightly divergent 3-4 rowsof setulae between median and dorsocentral rowsFore leg and all tarsi mostly yellow fore femurand distal tarsomeres sometimes with somebrown remainder of legs various shades of yellowthrough brown hind leg darkest hind tibia withstrongly curved preapical ventral spur longer thandiameter of tibia Wing length 12-21 mm secondcostal sector 18-20 times length of third costalsector Halter white Abdominal tergites shiningsetulose syntergite 1+2 mostly pale yellow with


brown lateral margins and a faint brown medianspot tergites 3-5 mostly brown medially with yel-low anterior and posterior margins sternites verypale setulose

Male postabdomen (Figs 13-14) Epandriumlong-haired surstylus curved posteromediallyparallel-sided distally with several short setaecercus elongate projecting posteroventrally cerciparallel in posterior view each with one long setaand several short setae phallus membranous

Type material examined ndash Holotype R ECUADORGALAacutePAGOS ISLANDS FLOREANA Post OfficeBay 20-30ix1925 A Wollebaek (ZMO) Paratypes2R FLOREANA Post Office Bay Seaside Ranvik 10-20xi1925 (ZMO)

Other material examined ndash 114= 315R from the fol-lowing islands and localities Bartolomeacute Espantildeola(Bahiacutea Manzanilla Punta Juarez) Floreana (BlackBeach island record only) Genovesa (south side ofisland) Isabela (Puerto Villamil near Punta Tortuga)Marchena (SW Playa) Pinta San Cristoacutebal (PuertoBaquerizo) Santa Cruz (Academy Bay Charles DarwinResearch Station Bella Vista Trail) Santa Fe (allmonths from January to June) (CAS CNC LEM)

Remarks ndash C W Sabrosky (in litt) considered Lgalapagensis a possible synonym of Liohippelatesincompletus (Becker 1912) from Peru Althoughthe two species are externally similar there areminor differences unfortunately the lectotype ofL incompletus is a female so male genitalic char-acters cannot be compared We prefer to treat L

galapagensis as currently recognized pendingexamination of male specimens of L incompletusfrom the type locality Coquillettrsquos (1901) recordof L pusio from the Galaacutepagos refers to L gala-pagensis (C W Sabrosky in litt) specimensidentified by Johnson (1924) as L pusio may alsobe this species but they were not examined

Specimens examined in this study were collect-ed primarily in the Littoral Zone including largeseries from marine iguana nesting sites and sealion beaches Specimens were also collected in theArid Zone and Transition Zone and agriculturalareas on several islands

Monochaetoscinella anonyma(Williston 1896)

(Figs 15-16)

Oscinis anonyma Williston 1896 423Monochaetoscinella anonyma Duda 1930 107

Material examined ndash ECUADOR GALAacutePAGOSISLANDS ISABELA 1= 1R Guava-pampa zone600m 4iii1989 B J Sinclair (CNC) SANCRISTOacuteBAL 1= 1km E Sierra San Janquin agricul-ture-pasture 13ii1989 B J Sinclair (CNC)

Remarks ndash This species is widespread in thesouthern Nearctic (Arizona Texas FloridaMexico) Central America the West Indies andSouth America as far south as northern ArgentinaThis is the first record from the Galaacutepagos and


15 16

Figures 15-16 Monochaetoscinella anonyma (15) Male genitalia lateral (16) Male genitalia posterior Scale bar =01mm

almost certainly represents a recent introduction toagricultural areas of the archipelago from theNeotropical region

Olcella anaclasta Wheeler amp Forrest sp n(Figs 17-19)

Description ndash Total length 15-19 mm Fronspunctate pruinose grey-brown posteriorly yel-lower anteriorly longer than broad projectinganteriorly frontal triangle including ocellar tuber-cle grey-brown slightly darker than or indistin-guishable from rest of frons pruinose about 05times length of frons antenna yellowish ventro-medially grey-brown dorsally and laterally firstflagellomere reniform arista short grey-brownappearing bare except at very high magnificationocciput dark grey-brown face short with pro-nounced carina lunule forming triangular platebetween antennae eye with very short omma-trichia long axis diagonal gena pale yellow ante-riorly grey-brown posteriorly and often alonggenal margin 02 times height of eye higher pos-teriorly vibrissal angle less than 90 degrees pro-jecting to same extent as frons palp and proboscisremarkably slender and elongate palp pale yellowas long as head proboscis brown well-sclerotizedgeniculate labellum 15-16 times length of headCephalic chaetotaxy 5 short reclinatefronto-orbital setae several proclinate interfrontalsetulae on anterior of frons single row of setulaeforming border of frontal triangle ocellar and pos-tocellar bristles pale erect or slightly reclinateconvergent inner vertical bristles not easily distin-guished from surrounding setulae outer verticalslonger pale setulae present on occipital and genalmargins vibrissa short pale Scutum grey dense-ly pruinose and punctate with three deeply incisedbrown lines of punctures the median one narrow-est and not always apparent scutum approximate-ly as long as broad postpronotum and scutellumdensely pruinose concolorous with scutumscutellum flattened dorsally and blunt apicallythoracic pleurites grey to brown mostly pruinoseexcept for area between mid and hind coxaThoracic chaetotaxy 1-3 postpronotal bristlesusually 1 longer than others 1 anterior and 2 pos-terior notopleural 1 dorsocentral 1 postalar 1intrapostalar 1 pair of short apical and 2 pairs ofshorter subapical scutellar bristles postpronotumscutum and scutellum setulose katepisternumwith a few short setulae Legs yellow to brown

Wing length 12-16 mm second costal sector 18times length of third costal sector dm-cu obliquenot parallel with r-m Halter white Abdominal ter-gites 1-5 brown (at least on anterior half) setulosesubshining sternites pale yellow setulose

Male postabdomen (Figs 18-19) Epandriumsubshining mainly yellow pale brown anterodor-sally short-haired cercus and surstylus mainlyyellow subshining surstylus long curved nar-rowed distally with a few long setae medially cer-cus triangular in posterior view pointed apicallyphallus long membranous

Type material ndash Holotype = ECUADORGALAacutePAGOS ISLANDS PINTA 220m sticky traps08ii1982 Y Lubin (CNC) Paratypes GENOVESA2R S side of island 200 yards from beach 4-6ii1967I L Wiggins in flight trap among Bursera graveolens(CAS) PINTA 29= 220m sticky traps 08ii1982 YLubin (CNC LEM)

Etymology ndash The species name is from the Greekanaklastos (bent back) referring to the greatly elongategeniculate mouthparts

Remarks ndash Although the elongation of the headand mouthparts in O anaclasta is more extremethan in other described species of OlcellaEnderlein 1911 there are undescribed speciesfrom Central America with intermediate degreesof development of the vibrissal angle proboscisand palps

Olcella lupina Wheeler amp Forrest sp n

(Figs 20-22)

Description ndash Total length 18-21 mm Frons palebrown to dark grey-brown entirely pruinose13-14 times as long as broad frontal trianglegrey not clearly differentiated from rest of fronsantenna mostly grey-brown first flagellomerereniform usually with some yellow ventromedial-ly arista short dark appearing bare except at veryhigh magnification occiput dark grey pruinoseface black on ventral margin paler dorsally withstrong carina visible in lateral view eye roughlyround with extremely short ommatrichia genadark grey posteriorly pale brown anteriorly02-03 times height of eye higher posteriorlyvibrissal angle strongly projecting palp yellow07 times length of genal margin proboscis darkshining geniculate each segment 12 times lengthof genal margin Cephalic chaetotaxy 5-6 recli-nate fronto-orbital setae several proclinate inter-frontal setulae and a row of setulae forming the



17

18

19

20

21

22

Figures 17-19 Olcella anaclasta (17) Head lateral (18) Male genitalia lateral (19) Male genitalia posteriorFigures 20-22 Olcella lupina (20) Head lateral (21) Male genitalia lateral (22) Male genitalia posterior Scalebars = 01mm

margin of the frontal triangle ocellar and posto-cellar bristles convergent to cruciate the postocel-lars slightly longer outer vertical bristle approxi-mately equal in length and strength to postocellarsinner vertical slightly shorter genal margin withshort pale hairs vibrissa pale weak but distinct-ly longer than subvibrissal setulae palp with anoutstanding seta at tip Thorax entirely dark pru-inose except for shining metanotum and broadshining stripe on pleura (consisting of anteroven-tral margin of anepisternum dorsal half of katepis-ternum ventral spot on anepimeron and anteriorspot on meron) scutum slightly longer than broadwithout incised lines of punctures scutellumbroader than long rounded not flattened dorsallyThoracic chaetotaxy 1 postpronotal 1 anteriorand 2 posterior notopleural 1 long dorsocentral 1long postalar 1 intrapostalar 1 pair of long apicaland 2 pairs of shorter subapical scutellar bristles(anterior pair shortest) postpronotum scutum andscutellum setulose katepisternum with a few shortsetulae Legs mostly dark coxae and femorabrown or grey-brown knees paler tibiae and tarsiyellow to pale brown fore tibia and tarsus palestWing with 2 fairly long strong setae basally on Cwing length 18-20 mm second costal sector 16-19 times length of third costal sector dm-cuoblique not parallel with r-m Halter whiteAbdominal tergites and sternites dark grey-brownpruinose

Male postabdomen (Figs 21-22) Epandriumbrown with short hairs surstylus parallel-sidedevenly curved posteriorly in lateral view withshort setulae cercus short only slightly convexwith a single long seta phallus long membranous

Type material ndash Holotype = ECUADORGALAacutePAGOS ISLANDS WOLF 31i-1ii1964 P DAshlock (BPBM) Paratypes DARWIN 9R 29i1964D QCavagnaro (CAS) WOLF 3= 1R 1 31i-1ii1964 P D Ashlock (BPBM) 5R 1ii1964 D QCavagnaro (CAS LEM)

Etymology ndash The species name is from the Latin lupinus(of wolves) referring to the type locality and to the largeand menacing mouthparts of these flies

Remarks ndash The great similarity between the malegenitalia of O lupina and those of C empheria(Figs 3-4) is indicative of a widespread problem inthe generic classification of the ChloropidaeAlthough many species of Olcella andConioscinella may be distinguished on the basis ofexternal characters (eg long geniculate mouth-parts projecting vibrissal angle incised lines on

scutum) there are species that are intermediate inall these characters Furthermore there are some-times greater genitalic similarities between speciescurrently assigned to different genera (as seenhere) than between species within the same genusThe current limits of chloropid genera based oncombinations of external characters will continueto break down as new species are described andwill almost certainly not stand up to rigorouscomprehensive phylogenetic analyses at thespecies level

Discussion

This study has more than tripled the number ofknown species of Chloropidae in the GalaacutepagosIslands The number of widespread chloropidspecies was lower than expected however giventhe increasing human traffic between theGalaacutepagos and the mainland of South America itis probable that additional synanthropic species ofChloropidae will be introduced to the islandsespecially to disturbed agricultural areas

The geographic affinities of the Galaacutepagoschloropid fauna are similar to those of most otherDiptera families in the region Cadrema pallida isa pantropical species found in low-lying andcoastal habitats on a number of oceanic islandsand continental land masses Monochaetoscinellaanonyma is widespread in the New World tropicsand subtropics Both of these species were proba-bly introduced by human activity All otherchloropid species are so far known only from thearchipelago but all show affinities to New Worldlineages especially to those in the northernNeotropical region Elachiptera cultrata is relatedto a group of species found from the southernUnited States to temperate South America(Wheeler amp Forrest 2002) Apallates sp Lioh-ippelates baptipalpis L galapagensis and Hippel-ates alyscus all belong to New World taxa that areespecially diverse in the southern Nearctic andnorthern Neotropics Olcella is primarily a NewWorld genus with species similar to O anaclastaand O lupina in Central America AlthoughConioscinella as currently recognized is cosmo-politan C empheria and C galapagensis appar-ently belong to the northern Neotropical C solutagroup The relationships of Diplotoxa loma andGaurax gethosyne are more obscure both generaare widespread diverse and poorly defined Theyare almost certainly not monophyletic as currently


defined and there is no information on phyloge-netic relationships within either genus

With additional collecting effort at least someof the currently ldquoendemicrdquo chloropid species mayalso be found on the west coast of Central Americaor northern South America This has been the casewith the chyromyid fly Aphaniosoma rabidaWheeler 1994 which was originally describedfrom Isla Raacutebida (Wheeler amp Sinclair 1994) buthas also been collected at Playa TamarindoGuanacaste Province Costa Rica (specimens inUSNM)

Species of Chloropidae were not as restricted tothe Littoral and Arid Zones as many other acalyp-trate families recorded from the islands (egAsteiidae (Forrest amp Wheeler 2002) Canacidae(Wirth 1969) Carnidae (Wheeler 2000)Chyromyidae (Wheeler amp Sinclair 1994)) TheGalaacutepagos chloropids like those in other regionshave a wide range of habitat preferences and werecollected in almost all habitat types in the archi-pelago often in high numbers

AcknowledgementsOur late colleague Curtis W Sabrosky provided valuableadvice on the identity of these species in the early stagesof the study this paper is dedicated to his memory Wethank Dr B J Sinclair (Bonn Germany) for his invita-tion to participate in the study of the Galaacutepagos acalyp-trate Diptera J M Cumming (CNC) A L Norrbom(USNM) K Ribardo (CAS) and G Soli (ZMO)arranged loans of specimens Fieldwork in theGalaacutepagos by S B Peck and B J Sinclair was facilitat-ed by the Galaacutepagos National Park Service Departmentof Forestry Ministry of Agriculture Ecuador and theCharles Darwin Research Station Isla Santa CruzFunding was provided by the Natural Sciences andEngineering Research Council of Canada

ReferencesBeardsley J W Arakaki K T Uchida G K

Kumashiro B R amp Perreira W D (1999) Newrecords for Diptera in Hawairsquoi Bishop MuseumOccasional Papers 58 51-57

Coquillett D W (1901) Papers from the HopkinsStanford Galapagos Expedition 1898-1899 IIEntomological results (2) Diptera Proceedings of theWashington Academy of Sciences 3 371-379

Curran C H (1932) The Norwegian zoological expedi-tion to the Galapagos Islands 1925 conducted by AlfWollebaek IV Diptera (Excl of Tipulidae andCulicidae) Meddelelser fra det Zoologiske MuseumOslo 30 347-366

Curran C H (1934) The Templeton Crocker expeditionof the California Academy of Sciences 1932 No 13Diptera Proceedings of the California Academy ofSciences (Series 4) 21 147-172

Duda O (1930) Die neotropischen Chloropiden (Dipt)Folia Zoologica et Hydrobiologica 2 46-128

Forrest J amp Wheeler T A (2002) Asteiidae (Diptera)of the Galaacutepagos Islands Ecuador StudiaDipterologica 9 307-317

Johnson C W (1924) Diptera of the WilliamsGalapagos Expedition Zoologica (New YorkZoological Society) 5 85-92

Loew H (1866) Diptera Americae septentrionalis indi-gena Centuria sexta Berliner EntomologischeZeitschrift 9 127-186

Nartshuk E P (2002) A revision of species of the genusCadrema Walker (Diptera Chloropidae) from islandsin the Indian Ocean Entomologicheskoi Obozrenie81 235-256 [In Russian]

Paganelli C H amp Sabrosky C W (1993) Hippelatesflies (Diptera Chloropidae) possibly associated withBrazilian Purpuric Fever Proceedings of theEntomological Society of Washington 95 165-174

Peck S B (1996) Origin and development of an insectfauna on a remote archipelago the Galaacutepagos IslandsEcuador Pp 91-122 in Keast amp Miller The originand evolution of Pacific island biotas New Guinea toeastern Polynesia patterns and processes 531 ppAmsterdam

Peck S B Heraty J Landry B amp Sinclair B J(1998) Introduced insect fauna of an oceanic archipel-ago the Galaacutepagos Islands Ecuador AmericanEntomologist 44 219-237

Sabrosky C W amp Paganelli C H (1984) FamilyChloropidae A Catalogue of the Diptera of theAmericas south of the United States Fascicle 81 1-63 Satildeo Paulo

Spencer K A (1977) A revision of the New ZealandChloropidae Journal of the Royal Society of NewZealand 7 433-472

Spencer K A (1986) The Australian Chloropinae(Diptera Chloropidae) Journal of Natural History20 503-615

Wheeler T A (2000) Carnidae of the GalapagosIslands Ecuador description and phylogenetic rela-tionships of a new species of Neotropical MeoneuraRondani 1856 (Diptera Carnidae) StudiaDipterologica 7 115-120

Wheeler T A amp Forrest J (2002) A new species ofElachiptera Macquart from the Galaacutepagos IslandsEcuador and the taxonomic status of CeratobarysCoquillett (Diptera Chloropidae) Zootaxa 98 1-9

Wheeler T A amp Sinclair B J (1994) Chyromyidae(Diptera) from the Galaacutepagos Islands Ecuador threenew species of Aphaniosoma Becker Proceedings ofthe Entomological Society of Washington 96 440-453

Williston S W (1896) On the Diptera of St Vincent(West Indies) Transactions of the EntomologicalSociety of London 1896 253-446

Wirth W W (1969) New species and records ofGalaacutepagos Diptera Proceedings of the CaliforniaAcademy of Sciences (Series 4) 36 571-594

Revised manuscript accepted March 2003


abdomens of mounted specimens (older air-driedspecimens were first relaxed in high humidity) andclearing them in 85 lactic acid heated in amicrowave oven for 1-2 30-second intervals sepa-rated by a cooling period Cleared abdomens werethen placed in glycerin for further dissection andexamination Dissected abdomens were stored inglycerin in plastic microvials pinned beneath thesource specimen

Key to species of Chloropidae of theGalaacutepagos Islands

1 Wing with costa extending to R4+5 crossveindm-cu absent Diplotoxa loma

- Wing with costa extending to M1+2 crossveindm-cu present 2

2 Arista broad sword shaped scutellum trape-zoidal body mostly yellowElachiptera cultrata

- Arista narrow rarely enlarged at base scutel-lum rounded posteriorly body colour variable 3

3 Hind tibia with apical or pre-apical ventral spurat least as long as tibial diameter 4

- Hind tibial spur absent or much shorter thantibial diameter 8

4 Thorax yellow in ground colour 5- Thorax black in ground colour 65 Hind tibial spur as long as basitarsus frontal

triangle shining scutum dorsally with palereddish stripes Cadrema pallida

- Hind tibial spur less than half as long as basi-tarsus frontal triangle dull scutum dorsally with black stripes Hippelates alyscus

6 Scutum dull eyes hairy hind tibial spur equal

in length to tibial diameter Apallates sp- Scutum shining eyes bare hind tibial spur

longer than tibial diameter 77 Legs yellow frontal triangle with sides straight

or nearly so extending slightly past middle offrons palp black basally yellow distally Liohippelates baptipalpis

- Legs partly brown hind femur and tibia dark-est frontal triangle with sides concave extend-ing almost to anterior margin of frons palpentirely yellow Liohippelates galapagensis

8 Vibrissal angle strongly produced proboscisgreatly elongate sclerotized geniculate (as inFig 17) 9

- Vibrissal angle rounded or only slightly pro-jecting proboscis not greatly elongate 10

9 Scutum with deeply incised longitudinal linesof punctures thoracic pleurites entirely pru-inose except for area between mid and hindcoxa Postgena very broad posteroventralangle less then 90deg (Fig 17) palp as long ashead many setae on frons and gena arisingfrom distinct punctures Olcella anaclasta

- Scutum without deeply incised lines of punc-tures thoracic pleurites with broad shiningstripe Postgena not as broad posteroventralangle greater than 90deg (Fig 20) palp at most07 times as long as head setae on head notarising from distinct punctures Olcella lupina

10 Frontal triangle shining notopleuron with 1anterior and 1 strong posterior bristle 11

- Frontal triangle dull notopleuron with 1 ante-rior and 2 strong posterior bristles 12

11 Frons with 1 outstanding fronto-orbital bristlescutellum black Monochaetoscinella anonyma

- Frons with several long fronto-orbital setaescutellum yellow Gaurax gethosyne


1 2

epd

proj

sur

phap

hyp

cer

Figures 1-2 Diplotoxa loma (1) Male genitalia lateral (2) Male genitalia posterior Abbreviations cer - remnant ofcerci epd - epandrium hyp - hypandrium phap - phallapodeme proj - posteroventral projection sur - surstylusScale bar = 01mm





(Figs 1-2)












Apallates sp









(Figs 3-4)







basph

distph

3 4

5 6







1984 10














(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion



































(Figs 1-2)












Apallates sp









(Figs 3-4)







basph

distph

3 4

5 6







1984 10














(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion


































Apallates sp









(Figs 3-4)







basph

distph

3 4

5 6







1984 10














(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion



































basph

distph

3 4

5 6







1984 10














(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion




































1984 10














(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion


































(Figs 7-8)



7 8










9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion







































9 10




(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion

































(Figs 9-10)







(Figs 11-12)






11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion


































11 12

13 14








(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion





































(Figs 13-14)














(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion







































(Figs 15-16)





15 16











(Figs 20-22)




17

18

19

20

21

22








Discussion








































(Figs 20-22)




17

18

19

20

21

22








Discussion
































17

18

19

20

21

22








Discussion





































Discussion































the chloropidae (diptera) of the gal pagos islands,...

Documents