New and little-known species of Forcipomyla (Diptera : Ceratopogonidae) associated with cocoa pollination in Brazil

13 Pages·1991·6 MB·

by  W W Wirth

PROC. ENTOMOL. SOC. WASH.
93(1), 1991, pp. 163-175
NEW AND LITTLE-KNOWN SPECIES OF FORCIPOMYIA
(DIPTERA: CERATOPOGONIDAE) ASSOCIATED WITH
COCOA POLLINATION IN BRAZIL
Willis W. Wirth
Research Associate, Rorida State Collection ofArthropods, and Cooperating Scientist,
NW
Systematic Entomology Laboratory, U.S. Department ofAgriculture, 1304 94th St.,
Gainesville, Florida 32606.
Abstract.—T'wo new species ofForcipomyia Meigen that are common in cocoa plan-
tations in Brazil, and are involved in cocoa pollination, are described and figured: For-
A
cipomyia {Forcipomyia) soriai Wirth and Forcipomyia (Lepidohelea) winderi Wirth. key
is given to distinguish the subgenus Forcipomyia s. str. from the closely related subgenus
Lepidohelea Kieffer. Keys are also presented to distinguish the Neotropical species ofthe
squamitibia Group ofForcipomyia s. str. which includes F. soriai, and the Neotropical
species ofthe bicolor Group ofLepidohelea, which includes F. winderi. Information on
the immature stages and biology for these and related species is summarized and the
immature stages ofF. (F.)pinamarensisSpinelli, also associated withcocoa, are described
and figured.
Key Words: Ceratopogonidae, Forcipomyia, cocoa, pollination
Biting midges ofthe genus Forcipomyia variation in the abundance and importance
Meigen are important pollinators ofcocoa in pollination between these taxa, between
{Theobroma cacao L.) wherever it is grown individual species, from region to region,
throughout the tropics. A voluminous lit- and also with the season.
eraturehasaccumulatedinthepast 30years From 1967 to 1983 intensivestudieswere
dealingwith the taxonomy, biology, and re- carried out by John A. Winder and Saulo
lationtococoapollinationofnumerousspe- deJesus SoriaattheCacao ResearchCenter
cies of Forcipomyia, especially ofthe sub- (CEPLAC) at Itabuna, Bahia, Brazil. More
genus Euprojoannisia Brethes. That which recentlyWinderhasresumed hiscocoa pol-
appeared up to 1977 was reviewed by Win- lination studies at the Fazenda Almirante,
der (1978). Approximately three-fourths of near Itajuipe in Bahia, and I have had the
the 70 speciesofCeratopogonidaethathave opportunity to cooperate in the determi-
been collected in cocoa flowers belong to nation ofthe ceratopogonids from his col-
thegenusForcipomyia. Nearlyathirdofthe lections. In Bahia, and especially at the Fa-
Forcipomyia midgesbelongto the subgenus zenda Almirante, by far the most abundant
Forcipomyia s. str., while in decreasing or- ceratopogonids collected in the cocoa plan-
der of importance the subgenera Eupro- tations belong to the Forcipomyia bicolor
joannisia Brethes, Thyridomyia Saunders, GroupinthesubgenusLepidohelea, andthe
Lepidohelea Kieffer, Microhelea Kieffer, F. squamitibia Group in the subgenus For-
Warmkea Saunders, and Lasiohelea Kief- cipomyia. The purpose of this paper is to
fer, are also involved. There is considerable describe the two most important species in
164 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
Winder's collections to make the names cies of Thyridomyia usually are found in
available for his biological studies, and to association with moss or algae in rather
provide means fordistinguishingthem from damp habitats. Immature stages of
closely related species. Warmkea species are frequently found in
Explanation ofthe taxonomic characters leafaxilsofPandanus, aroids,andepiphytic
used can be found in the general papers on and terrestrial bromeliads, and less often in
Ceratopogonidae by Wirth et al. (1977) and rotting cocoa leaves on the ground, in ba-
Downesand Wirth (1981), and the revision nana stems, and inbractsofHeliconia. Spe-
ofthe North American Euprojoannisia by cies of the subgenera Forcipomyia, Lepi-
Bystrak and Wirth (1978). Holotypes and dohelea, and Schizoforcipomyia are less
allotypesofnewspeciesaredepositedinthe aquatic and are more commonly associated
National Museum of Natural History with rotting plant material, which in cocoa
(USNM), Smithsonian Institution, Wash- plantations and their environs often in-
ington, D.C.; paratypes as available in the volves heaps of cocoa pods, old banana
Florida State Collection of Arthropods, stems, cocoa leaves, coconut debris, bracts
Gainesville,Rorida;CaliforniaAcademyof ofHeliconiaandCalathea, androttingfruits
Sciences, San Francisco, California; Museo ofcoconut, calabash, palm nuts, etc.
de La Plata, La Plata, Argentina; Instituto The taxonomic key to the subgenera of
Oswaldo Cruz, Rio de Janeiro, Brazil; Brit- Forcipomyia published by Wirth and Ra-
ishMuseum (Natural History), London; and tanaworabhan isout-datedand hasbeen re-
Museum National d'Histoire Naturelle, placed by that ofDebenham (1987a). Until
Paris. the work ofDebenham (1987a, b, c,) it was
Taxonomic work on the cocoa-pollinat- not possible with confidence to distinguish
ingForcipomyiamidgeshasasitsmainpur- between the subgenera Forcipomyia s. str.,
pose the identification ofthe most impor- Lepidohelea, andSchizoforcipomyia, butshe
tantpollinatingspecies, thelocation oftheir pointed out characters in all stages to make
mostimportantbreedingplaces, andtheac- possible a key to these taxa for the Western
cumulation ofenough information on their Hemisphere.
biology so that a practical regimen ofcocoa
cultureandmidgehabitatmanipulationcan
be devised to maximize cocoa pollination Key TO Subgenera of Forcipomyia
and crop production (Young 1982, 1983). Related to Lepidohelea
Although the study ofForcipomyia breed- \. Hind tarsal ratioabout 1.0; wing usually with
ing places and identification of immature palecostalspotorpalemottling;secondradial
stages has been pursued intensively in cell short, costal ratio about0.5 2
America since the early work of Saunders - Hind tarsal ratio less than 0.5 orgreaterthan
(1956, 1959),progresshasbeendifficultand r1a.t5i;owainngdusseucaolnlydwriatdihaolutcepllalveamrairokuisngs;costal
slow. Nevertheless, important contribu- miscellaneous subgenera ofForcipomyia
tions have been made by Winder and Silva 2. Bodywings,and/orlegswithflattenedstriated
(1972), Winder (1977a), Soria et al. (1979), scales ranging from long and 1-striated setae
and Soria and Wirth (1979) in Brazil, and tobroad,short,multistriatedscales;palpuswith
Young (1982, 1983, 1986) in Costa Rica. third segment more or less spindle-shaped,
slightlymoreswollenonbasalhalfwithsensory
LarvalhabitatsoiForcipomyiamidgesare pit located at about 'A length; legs with char-
somewhat correlated with their taxonomic acteristicbandingofalternatingpaleanddark
group. The habitats o{Euprojoannisia spe- bands, at least on hind tibia; male genitalia
cies are usually semiaquatic situations such oftenwithwhitishdististyleandbicoloredninth
asalgae-covered rocks ormud, wet moss or sternum and basistyle; dististyle without long
setaeonoutermargin;aedeagusvarious;larva
leaves, matsofdecayingaquaticvegetation, with h hairs ofbody, andp and q head hairs
and leafaxils ofwater-holding plants. Spe- swollen near base and becoming filamentous
VOLUME NUMBER
93, 1 165
distally, usuallysomeaboutaslongasbodyof nentsetosetubercles;respiratoryhornlargeand
larva; anal pseudopod oflarva cleft with each globularwith spiracularopenings in astraight
branch bearing 6-8 booklets apically and 3-4 row squamitibia Group
booklets on innerlateral side 3 Femaletibiaewithouthastatespines;palewing
Body,wings,andlegsrarelywithflattenedstri- spotsmaller,notcoveringendofsecondradial
ated scales; palpuswith thirdsegmentswollen cell; hind tarsal ratio 0.50-0.76; male para-
onbasalVj-'A,slenderdistally,withsensorypit mereswithbasesjoinedinabroadplate;pupa
located at basal 'A; legs, if banded, not with with setose tubercles that are often elongate
alternatingpaleand darkbandson hind tibia; andprominent;respiratoryhornsmalltomod-
maledististylenotcontrastingwhitish,butwith erate in size, with longer neck and spiracular
well-developed setae on outer margin; aede- openings in a more or less irregularrow ....
agusshield-shaped,apex pointedwithmedian argenteola Group
ridge; larvalbhairsandpandqheadhairsnot
as above; anal pseudopod of larva not cleft,
bearingunbroken row ofstrong hooks Forcipomyia {Forcipomyia)
Subgenus Forcipomyia
pinamarensis Spinelli
Maleparamereswith basalapodemesforming
a U-shaped arch, parameres a pair ofslender Figs. 1-11
to stout, straight or bowed, well-sclerotized Forcipomyia {Forcipomyia) pinamarensis
bladesorrodsreachingtonomorethanmiddle
ofbasistyles;aedeagusvarious,withlowtohigh Spinelli, 1983: 121 (male, female; Argen-
basal arch, distal process various, often with tina; figures).
recurved points; dististyle straight or slightly —
Types. Holotype female, allotype male,
curved, tapering to slendertips; palpus 5-seg-
mented;twospermathecae;scalesofbodyand 12 female, 29 male paratypes: Argentina,
legsnevershort and broad Buenos Aires Prov., Pinamar, Partido de
SubgenusSchizoforcipomyia General Madariaga, 27.iii.1981, G. R. Spi-
Maleparameresseparateorindistinctlyjoined nelli (in Museo de La Plata).
in an anteromesal bridge; parameres various, —
Note. Shortly before his death in 1968,
usually straight slender rods nearly as long as
basistyle and tapering to filamentous tip; ae- Professor L. G. Saunders ofthe University
deagus usually much longerthan broad, basal of Saskatchewan very generously donated
arch low or absent, usually stout distally; dis- to the Smithsonian Institution in Washing-
tistyleoften sinuate withexpandedtip; palpus ton his extensive worldwide collection of
4- or 5-segmented; one or two spermathecae;
Forcipomyiamidges, mostlyreared from the
scales ofbody and legs often short and broad
Subgenus Lepidohelea immature stages, alongwith his manuscript
notes and drawings. Amonghis manuscript
notes were drawings and the description of
Subgenus Forcipomyia, s. str. aspeciesthathecollectedandrearedin Bra-
Wirth(1982)discussedthetaxonomyand zil in 1923 and designated as "B-70." Dur-
biology of the Neotropical Forcipomyia s. ingthe present study itwasdetermined that
str. associated with cocoa pollination, in- Saunders' "B-70" wasthe same speciesthat
cluding four species which he placed in the Spinelli described as Forcipomyia pina-
""argenteolaGroup." Healsodiagnosedand marensis from Argentina. This species is
keyed out F. (F.) squamitibia Lutz, which abundant and widespread in cocoa planta-
with similar species could be confused with tions; therefore this opportunity is taken to
species ofthe argenteola Group. Species of publish Saunders' description and figures of
these two groups can be separated by the the adults and immature stages, and to list
following key: new distribution records. The following de-
scriptions are adapted from Saunders'
1. Female tibiae with row of hastate spines on manuscript notes:
extensorsurface;palewingspotatendofcosta "Larvae ofthis species were taken in fair
larger,includingmostofsecondradialcell;hind
tarsal ratio about 1.0; male parameres with numbers under the bark ofa rotting log in
slender common base; pupa without promi- the forests ofCorcavado, Rio, 18.viii.l923.
166 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
150
Figs. 1-n. Forcipomyiapinamarensis: 1^, female; 5, 6, pupa; 7-10, larva; 11, male: 1, antenna; 2, radial
vemsofwing; 3, hastatespinesoftibia;4, hindleg; 5, respiratoryhorn;6, pupa; 7, lasttwoabdominalsegments;
8, head and body setae, letteringas in text; 9, 10, integumental armature ofbody; 11, genitalia, partial (drawn
by L. G. Saunders; scale in microns).
VOLUME 93, NUMBER 1 167
Pupae and adults were obtained from them light brown on all the usual sclerites; a very
in due time.: small scleritein frontofthebaseofthewing
"Larva. Color white, the macrochaetae darkbrown. Legsochreous, foreandmiddle
black, their papillae light brown. Head coxaebrown,hindcoxapaler;afuscousband
ochreous to black around mouth. Protho- justbefore extremity ofhind femur(Fig. 4);
racic pseudopod ofmoderate length, forked duskiness oftibiae due to presence ofnar-
in its distal half, with the crown of hooks row black squamous hairs; outer surface of
disposed equally over the end of each all tibiae with a few ochreous lanceolate to
branch. Chaetotaxy (Fig. 8): p and q hairs squamoushairs(Fig. 3), notstriatedlikethe
ofhead lanceolate; a hairs ofbody lanceo- more usual decumbent hairs oflegs and ab-
late; b and d separate, b short, curved, spi- domen. Proportional length of hind tarsal
nulate distally. Dorsal hairs oflast abdom- segments, 15:12:7:6:5. Wingdensely clothed
inal segment arising from a cap of chitin with dark hairs, narrowly squamous; area
(Fig. 7). Cuticulararmature in form ofshort of pale yellowish hairs at end of costa.
stout stem with a number of apical arms Branchesofradiusdividingto form distinct
(Fig. 10); simple cones or hooks on pleura cell (Fig. 2); end ofcosta at middle ofwing,
and last abdominal segment (Fig. 9). Cauda fork of Cu directly below. Halteres white.
pointed. Anal blood gills bilobed, the outer Abdominaltergitesuniformlybrownexcept
lobe much smaller than the inner. 8 + 9 which are pale ochreous with faint
"Pupa (Fig. 6). Color (exuviae) pale lateralbrown spots; stemites 3-7 palebrown
ochreous throughout. Head devoid of pa- with stronger lateral coloring, 8 yellowish.
pillae. Thorax with one dorsal pair ofsetig- Cerci pale. Tergites and pleural membrane
erous papillae and three other pairs ofvery clothed with narrow black-striated hairs;
small papillae. Abdomen with no cuticular pale simple hairs on venter. Spermathecae
spines,fewmicrochaetae.Terminalabdom- two; 0.090 by 0.056 mm, subspherical, no
inal processes ofmoderate length. Protho- chitinization on neck. Length 1.6 mm; wing
racic respiratory horn (Fig. 5) in form of 2.1 1 by 0.45 mm; antenna 0.6 mm.
subspherical knob on short, broad-based "Male. Specimen very imperfect. Palps
stem. About 16 spiracular papillae having much as in female, but terminal segment
smallexternalmembranescurveupthepos- longer; proportionallengths, 9:1 1:38:20:16.
teriorsideandoverthetop. Length2.3 mm. Hypopygium (Fig. 11): Aedeagus long,
"AdultFemale. Facepale, ocellartriangle rounded distally, hairy, without lateral ca-
faintly brown; dark ring around bases of rinae. Parameresjoinedinmiddle,aslender
antennae. Palpus brown with light bands at pairofbowed styletsextendsbackwardsbe-
distal articulations; segment three much yondend ofaedeagus. Sidepiecesofforceps
swollen in proximal one-third, distal neck long, tapered slightly, with distinct brush of
tapering gradually; proportional length of fine hairs on inner side near anterior end;
segments, 9:11:37:21:13. Antenna (Fig. 1): claspers lost.
Scape light brown, flagellum white tinged "This is a very typical Forcipomyia con-
faintly with brown distally; proximal seg- formingclosely to type. The larvae are very
ments flask-shaped with broad elongated close to F. bipunctata (Linnaeus), differing
neck;proportionoffirst8 tolast 5 segments, only in the presence ofa chitinous cap on
43:34; last 5, 6:6:6:6:10; terminal style a the last abdominal segment and in the ex-
largeknobwith indentedextremity. Thorax traordinary dorsal cuticular armature. The
uniformly brown on dorsum, with coarse verybare, spinelesspupaisdistinctive, pos-
golden pubescence; humeral angles pale; sessing as it does a row ofunusually small
scutellum brown to dark brown around respiratorypapillaeon the prothoracic horn.
margins; postscutellum dark brown. Pleura Theadult female belongsto the smallgroup
168 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON
exhibiting a row oflanceolate scales on the distal segments unusually short for the ge-
tibiae; apart from this character they may nus; proximal verticils appearing bushy.
readily be distinguished from most other Palpus (Fig. 14) with lengths ofsegments in
species by the minute dark sclerite in front proportion of 10-11-22-13-10; third seg-
ofthe wing base." ment slightly swollen at base (palpal L/W
Distribution.—Argentina, Brazil, Costa ratio 2.2), gradually taperingto tip, a small,
Rica, Panama. round, shallow, sensory pit near base; seg-
Specimens examined.—ARGENTINA: ments 4 and 5 with faint separation. Man-
Pinamar, Bs. As., 27.iii.1981, G. SpineUi, 1 dible without teeth.
female (paratype). BRAZIL: Bahia, Itabu- Thorax: Pale yellowish brown, darker on
na, 1970, J. Winder, emergence traps, 5 pleura, scutellum, postscutellum, and cox-
males, 6 females; Bahia, Ilheus, CEPEC, ae. Mesonotum with suberectfineyellowish
15.vi.l974, S. Soria, 1 female; Para, km 3, hairs. Legs uniformly yellowish except tib-
Rodovia Castanhal-Curuca, Faz. Agricul- iae distally and all tarsi proximally, slightly
tura Bandeirante, 31.v.1980, S. Soria, ex infuscated;withyellowishandbrownish,fine
cacao cherelles, 4 males, 2 females; Rio de hairs, no broad flat scales; tibiae, and tarsi
Janeiro, Corcavado, 18.viii.l923, L. G. to a lesserdegree, with long, slender, brown
Saunders, under bark, 4 larvae, 1 pupal ex- setae on extensor side; tarsomeres 1-3 with
uviae, 2 females. COSTA RICA: Heredia row of strong, blackish, ventral spines, a
Prov., vie. LaVirgen, FincaLaTigra, 30.vi- pair at apices oftarsomeres much stronger;
3.vii.l980, 25.vii.1981, 1 l,30,vii.l982, A. all tibiaewith a rowoferect, nearly hyaline,
M. Young, 2 males, 3 females. PANAMA: hastate setae (Fig. 12) with petiolate bases
Canal Zone, Pipeline Road, vii.1967, F. S. and bluntly pointed or rounded tips. Claws
Blanton, light trap, 1 female; Canal Zone, moderately long and slender, strongly
Balboa, fogging Cassia tree, 19.vii.l979, E. curved, tapering to sharp tips; empodium
Broadhead, 1 female; Canal Zone, Gatun, small. Hind leg with lengths from femur to
humid forest, fogging Spondias and Cordia tarsomere 5 as 82-80-24-26-16-12-8; tarsal
tree canopy, 9-10.vii.1979, E. Broadhead, ratio (T1/T2) 0.92; hind tibial spur yellow-
2 males, 3 females. ish, straight, slender, taperingto sharppoint.
Wing densely and uniformly covered with
Forcipomyia {Forcipomyia) soriai Wirth, brownish macrotrichia, except two anterior
New Species
patches of blackish, one-striated, scalelike
Figs. 12-21 macrotrichia over radial field andjust past
A
hairy yellowish species with dark hal- end ofcosta, the two patches separated by
teres; femalewithabundantlong, dark, one- a small pale spot ofyellowish scales at end
striatedscalelikesetaeonradialfieldofwing ofcosta. Halter brownish.
and on abdomen; wing with one small yel- Abdomen: Brownish, darker posteriorly
lowish costal spot; tibiae with row ofblunt- withnumerouslong, brownsetae,especially
pointed, pale, hastatesetae; hindtarsalratio on pleura and on posterior segments. Sper-
0.92. mathecae (Fig. 15) two, dark brown, rela-
Allotype female.—Winglength 1.14mm; tively large; unequal, measuring 0.118 by
breadth 0.52 mm; costal ratio 0.47. 0.087 mmand0.087 by0.075 mm;thelarg-
Head:Pale brownish; overall clothed with er oval, the smaller subspherical; both with
numerous long, brown, one-striated setae lightly sclerotized, slender, almost thread-
giving a bushy appearance. Antenna (Fig. like entrances to the ducts.
13)withlengthsofflagellarsegmentsin pro- Holotype male.—Wing length 1.21 mm;
portion of20-15-15-15-15-15-15-16-15-15- breadth 0.43 mm; costal ratio 0.44. Similar
16-16-22,antennalratio(11-15/3-10)0.67; to the female with the usual sexual differ-
VOLUME 93, NUMBER 1 169
Figs. 12-21. Forcipomyiasoriai: 12-15, female; 16, pupa; 17, 18, male; 19-21, larva: 12, hastate spines of
tibia; 13, antenna; 14, palpus; 15, spermathecae; 16, respiratory horn; 17, aedeagus; 18, genitalia, aedeagus
omitted; 19, head; 20, bodysegment, lateral view; 21, lasttwoabdominal segments, dorsal view(letteringasin
text; scale in microns).
ences. Antenna with well-developed plume in female, palpal ratio 3.4. Legs with exten-
ofyellowish-brown verticils; lengths offla- sorsetaelongerthaninfemale,hastatespines
gellar segments in proportion of25-15-15- absent on tibiae; tarsi with numerous long,
15-15-15-15-15-16-55-28-20-32, antennal slender, black spines ventrally; claws long,
ratio (12-15/3-1 1) 1.03. Palpuswith lengths slenderandcurved;empodiumslenderwith
of segments in proportion of 9-1 1-27-12- shortrays. Hindlegwithlengthsfromfemur
10; third segment more slenderat base than to tarsomere 5 as 86-80-24-26-17-13-10;
170 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
tarsal ratio 0.92; tibial spur pale yellow, tory horn (Fig. 16) irregularly oval in out-
straight, slender, Vy as long as basitarsus, line, with very short, stout peduncle; in
taperingto sharp point. Wingwith pale yel- cross-section appearing slightly flattened
lowish macrotrichia overall, a small clump with the spiracularopenings forminga slight
of strong blackish macrotrichia over first rim; 16-18 spiracularopenings in a straight
radialcellandanotherjustpastendofcosta, row up posterior side and around tip. Body
the two separated by a small area ofyellow- tubercles not developed, setae scarcely vis-
ish macrotrichia. Halter brownish. Abdo- ible under high magnification.
menyellow, withbrownishpigmentedareas Distribution.—Brazil, Puerto Rico, Trin-
laterally on each segment; with abundant, idad.
—
long, brownish, bristlelike setae, without Types. Holotype male, allotype female,
broad striated scales. Brazil, Itajuipe, Fazenda Almirante,
Genitalia(Fig. 18).—Ninthsternumabout 9.V.1988, J. A. Winder, in emergence trap
twiceasbroadaslong,caudalmarginslight- (deposited in USNM). Paratypes, 37 males,
ly convex with shallow mesal concavity; 44 females, 1 larva, 2 pupalexuviae: Brazil,
basistyleabout twiceaslongasbroad, with- same data as types, but dates iv.l987-
out stout setae on mesal face but with nu- vii.1989, 28 males, 35 females; Bahia, Ita-
merous long, bristlelike setae laterally and buna, 1970, J. A. Winder, emergence trap,
distally that are about twice as long as dis- 6 males, 4 females; Bahia, Ilheus, CEPEC,
tistyle, the latter straight, slender, and ta- 4-9.viii.1972, S. Soria, at flowers ofcacao,
pered to slender tip. Aedeagus (Fig. 17) 1 male, 3 females. Puerto Rico, Maricao,
shield-shaped, broad basally, length equal 21.i.l953, L. G. Saunders, reared from ba-
to basal breadth, with short basal arms, the nanaleaves, 1 larva, 1 pupalexuviae, 1 male.
basalarchonlyaslightconcavity; sidescon- Trinidad, no locality, 7.V.1953, L. G. Saun-
vex, taperingtoabluntcaudomedianpoint. ders, reared from wet leaves in stream, 1
Paramereswith slender, straight, basal apo- larva, 1 pupal exuviae, 1 male.
demes;joined mesally on basal fourth; dis- Discussion.—This species is dedicated to
tal portions relatively stout, nearly straight Saulo de Jesus Soria of Bento Goncalves,
and parallel, with the blunt tips slightly R.S., Brazil, in recognition ofhis long and
curved laterad. continued interest and his many important
Larva.—Length 4-5 mm when mature. contributions pertaining to the role ofcer-
Color yellowish white; head capsule pale atopogonids in cocoa pollination through-
brownish; dorsal body hairs borne on out Central and South America.
brownishconicalprominences(Figs. 20, 21). Lane's description and figure ofthe male
Hairs p and q of head (Fig. 19) long and genitaliaofF. argenteolaMacfiefrom Brazil
slender,withspear-shapedtip, thatofphair probably refers to F. squamitibia Lutz, as
much larger than q. Antenna long and ta- all species ofthisgroup have the hind tarsal
pering, borneonalargesubconicalpedestal. ratio ofabout 1.0 and the parameres with
Body hairs as in Fig. 20; a hairs with long narrow basal separation and quite slender
slender stem and spear-shaped tip, even on proximally. Forcipomyia argenteola, to the
last segment (Fig. 21). Cuticular armature contrary, belongs to a group of species re-
absent. Prothoracic pseudopod cleft, each viewed by Wirth (1982) which has the hind
lobe with 6-7 small brown hooks; posterior tarsal ratio about 0.5 and the male para-
pseudopod with a row of 14-16 brown mereswith basesfusedinabroadplatewith
hooks. Cauda short but pointed. posteriorconcavity and the caudal process-
Pupa.—Exuviae only, length not mea- es relatively stout and tapering to a sharp-
sured. Color pale yellowish, integument pointedtip. Wirth(1982)describedthe male
without spinules or shagreening. Respira- ofF. argenteolafrom maleswithassociated
.
VOLUME NUMBER
93, 1 171
females from Nova Teutonia, Brazil, the of the subgenus Lepidohelea, including
type locality. Species of the Forcipomyia characters that may be used to separate the
squamitibia Group can be distinguished by speciesgroupsoccurringintheAustralasian
the following key: Region. TheAmericanspeciesappeartofall
in two groups differing from the Old World
Key to Species of the species, most of which would fall within
Forcipomyia squamitibia Group whatDebenhamhasnamedthechrysolopha
1 Hastate spines offemale tibiae in two dense Group. The American species may be dis-
irregularrows; largedark brown species, wing tinguished from the chrysolopha Group by
length 1.7-2.0 mm; hind tarsal ratio 1.0 .... the following key:
chilensis (Philippi)
- Hastate spines offemale tibiae in one sparse 1. Palpus with four segments; one spermatheca;
row; large orsmall, pale ordark species 2 male dististyle more or less expanded distally
2. Mesonotumwithdarkvittae;smalldarkbrown (Western Hemisphere species)
species, wing length 0.93 mm; hastate tibial annulatipesGroup
spinesbrown, short,andbroadwithsharptip; - Palpuswith five segments; two spermathecae;
hind tarsal ratio 0.9 sexvittata Wirth male dististyle various 2
- Mesonotum not vittate; color, size and tibial 2. Male dististyle straight or slightly curved, ta-
spines various 3 peringtoslendertip(WesternHemispherespe-
3. Hastatetibialspineslongerthanwidthoftibia 4 cies) bicolorGroup
- Hastate tibial spines as long as width oftibia - Maledististyle straight to sinuate, tip moreor
or shorter 5 less expanded (Eastern Hemisphere species)
4. Yellowish brown species with unmarked yel- chrysolophaGroup
lowish legs; tibial spines slightly longer than
widthoftibia,swollenin midportion,tapering Forcipomyia {Lepidohelea) bicolor
tosharptip;antennapale;thirdpalpalsegment Species Group
swollen on proximal '/s; wing length 1.3 mm;
hind tarsal ratio 1.0; male parameres with fi- Forcipomyia {Lepidohelea) winderi Wirth,
liform tips straight and parallel squamitibia Lutz New Species
- Dark brown species with brown legs; tibial Figs. 22-30
spinesmuchlongerthanwidthoftibia,slender
A
and only slightly swollen in midportion; an- hairy, predominantly yellowish species
tenna dark brown; third palpal segment swol- with brownish markings; female wing dark
len on proximal %; wing length 1.5 mm; hind with one large pale anteriorspot, male wing
tarsal ratio 1.3; male unknown paleexceptfordarkcostalfield;palpusstub-
tenuisquamipesWirth
5. Legspaleyellow,hindfemurwithnarrowapex by, third segment not swollen, with small
brownish; hastate tibial spines petiolate with shallow pit; antenna moderately long; legs
broadapices; winglength 1.1 mm; hind tarsal banded with bicolor Group type markings,
ratio 0.9; spermathecae pale; male parameres femora pale except hind femur with apical
with filiform tips crossing(Fig. 11) dark brown band, all tibiae and tarsi band-
pinamarensisSpinelli
- Legsduskyyellowishwithtibiaebrownishdis- ed; male genitalia with brownish markings
tally,hindfemurwithoutdistaldarkband;has- laterally on ninth sternum, distal halves of
tate tibial spines slightly petiolate with tips basistyles, and all ofdististyles.
moderatelybroadandslightlytaperedtopoint; Femaleholotype.—Winglength 1.11 mm,
wing length 1.14 mm; hind tarsal ratio 0.92; breadth 0.44 mm; costal ratio 0.47.
spermathecae dark brown; male parameres
stout distally with tips abruptly bent a short Head: Palebrownish, overall clothed with
way, divergent (Fig. 18) numerous long, brown, one-striated setae
soriaiWirth, new species giving head a bushy appearance, especially
strong and numerous on clypeus. Antenna
Subgenus Lepidohelea Kieffer
(Fig. 22) with lengths of flagellar segments
Debenham (1987b) has presented an ad- in proportion of20-16-16-16-16-16-15-15-
mirablediscussionofthedifficulttaxonomy 20-20-20-20-23,antennal ratio0.79; anten-
172 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
50
©
23
.^^y^^'
Figs. 22-30. Forcipomyia windert:22-25, 29, 30, female;26-28,male: 22,antenna; 23,palpus; 24,26,wing;
25, striated scales ofbody and legs; 27, aedeagus; 28, genitalia, aedeagus omitted; 29, fore, mid, and hind legs
(top to bottom); 30, spermathecae (scale in microns).
nal verticils short and strong. Palpus (Fig. figured: fore and mid femora pale, tibiae
23) with lengths ofsegments in proportion with narrow basal and broad apical bands;
of8-12-17-9-8; third segment slightly swol- hindfemurwithbroadapical,andhindtibia
len in midportion, spindle-shaped, with un- with broad basal and narrow subapical
usually small, shallow, round, sensory pit. bands; bandsmoreintenseonhindleg; fem-
Mandible without teeth. ora and tibiae with some long, brownish,
Thorax: Reddish brown; mesonotum bristly setae, and abundantly clothed with
somewhat paler in midportion, clothed broad, sharp-pointed, 4- or 5-striated scales
abundantly with long, pale brown, bristly, (Fig. 25). Hindtibialspurshortandpointed,
erect setae and fine, yellowish, appressed nearly straight, about as long as breadth of
hairs. Scutellum paler in midportion. Legs tibia. Tarsomeres dark banded in midpor-
(Fig. 29) yellowish, with brownish bands as tion; hind tarsal ratio 1.07; claws slender