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Forcipomyia bicolor and Related Species of the Subgenus Lepidohelea in Brazil (Diptera: Ceratopogonidae) Author(s): Willis W. Wirth Source: The Florida Entomologist, Vol. 74, No. 4 (Dec., 1991), pp. 506-517 Published by: Florida Entomological Society Stable URL: http://www.jstor.org/stable/3495404 Accessed: 15/06/2009 16:22 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showpublisher?publishercode=fes. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org. Florida Entomological Society is collaborating with JSTOR to digitize, preserve and extend access to The Florida Entomologist. http://www.jstor.org

506 Florida Entomologist 74(4) December, 1991 FORCIPOMYIA BICOLOR AND RELATED SPECIES OF THE SUBGENUS LEPIDOHELEA IN BRAZIL (DIPTERA: CERATOPOGONIDAE) WILLIS W. WIRTH Research Associate, Florida State Collection of Arthropods 1304 NW 94th St., Gainesville, Florida 32606, U.S.A. ABSTRACT Forcipomyia (Lepidohelea) bicolor Lutz from Brazil and two other closely related species, F. dubia Macfie and F. lacrimotorii Macfie, are redescribed from types. Forcipomyia discoloripes Macfie is a junior synonym of F. bicolor (NEW SYNONYMY). The Forcipomyia bicolor Group of species is diagnosed and a revised key is presented to the nine described Neotropical species. RESUMEN En base a prototipos se describe de nuevo la especie Foripomya (Lepidohelea) bicolor Lutz del Brazil y otras 2 especies estrechamente relacionadas, F. dubia Macfie y F. lacrimotorii Macfie. Forcipomyia discoloripes Macfie es un sinonimo de F. bicolor (Nuevo Sinonimo). Se hace una diagnosis del grupo de especies de Forcipomyia bicolor y se presenta una clave revisada describiendo las nueve especies neotropicales. After nearly 50 years of investigation, the most important pollinators of Theobroma cacao L., source of commercial chocolate and cocoa, have been found to belong to the ceratopogonid genus Forcipomyia Meigen (Billes 1941, Macfie 1944, Posnette 1950, Saunders 1956, 1959, Winder 1978, Young 1983). Earlier research indicated that the most frequent and important pollinators were species of the subgenus Euprojoannisia Brethes (Saunders 1959, Soria & Wirth 1974, Bystrak & Wirth 1976). More recently attention has also turned to members of other subgenera of Forcipomyia as more abundant in cacao plantings and also important as cacao pollinators. Thus Winder (1977, 1978) found that about a third of the Forcipomyia midges in cacao groves in Bahia, Brazil, belonged to the subgenus Forcipomyia s. str.; Young (1983) found the proportion much higher in Costa Rica. Kaufmann (1975), hovever, found F. (Microhelea) inornatipennis (Austen) and F. (Lepidohelea) squamipennis Ingram & Macfie to be more important in Ghana. Winder (1978) pointed out that there are so many variables in weather conditions, available terrestrial midge habitats, and flowering patterns of the cacao trees themselves, that relative importance of midges and other insects as pollinators is difficult to evaluate. Moreover, cage experiments with insects on cacao flowers also usually lead to unrealistic results. Nevertheless, recent collections that I have examined made by John Winder in the cacao plantation Fazenda Almirante in Bahia, Brazil, show that several species of the subgenus Lepidohelea Kieffer are abundant and at times dominant in cacao plantations. The systematics of the subgenus Lepidohelea remained extremely confused until Debenham (1987), in reviewing the Australasian species of the genus Forcipomyia, succeeded in finding reliable adult characters to separate species of Lepidohelea from the closely related Forcipomyia s. str. Species of Lepidohelea can be recognized by their spindle-shaped third palpal segment, legs with characterstic banding of alternating

Wirth: Forcipomyia bicolor Group 507 pale and dark bands, at least on hind tibia, male dististyle without long setae on the outer margin, and the larva with b hairs of body and p and q hairs swollen near base and becoming filamentous distally. Wirth (1991) followed with a key to divide the species of Lepidohelea into three species groups as follows: 1. Palpus with four segments; one spermatheca; male dististyle more or less expanded distally (Western Hemisphere species)... annulatipes Group 1'. Palpus with five segments; two spermathecae; male dististyle various... 2 2(1). Male dististyle straight or slightly curved, tapering to slender tip (Western Hemisphere species)... bicolor Group 2'. Male dististyle straight to sinuate, tip more or less expanded (Eastern Hemisphere species)... chrysolopha Group The species of the chrysolopha Group were admirably discussed and the Australasian species revised by Debenham (1987). The annulatipes Group has not been formally diagnosed, but is now known to comprise three Brazilian species, annulatipes Macfie, brasiliensis Macfie, and kuanosceles Macfie (Macfie 1939). I now have in preparation a revision of this group which is also represented in North America by at least two undescribed species. Wirth (1991) did not diagnose the bicolor Group, but presented a key to nine Neotropical species, of which one, F. winderi, was described as new. The bicolor Group is represented in North America by four described species, beckae Wirth, christiansoni Wirth & Hubert, seminole Wirth, and varipennis Wirth & Williams, and at least six undescribed species. The immature stages of members of the bicolor Group have not been described. Because of their abundance in collections of cacao pollinators, it is urgent that accurate names be ascribed to the important pollinators and that keys be presented for their identification. As a preliminary to planned revisions, first of the Nearctic species, and then of the Neotropical species, of the bicolor Group, it was necessary to study the available types of previously described species, whose identity remained questionable because of inadequate descriptions and figures. Notes are presented here on four Brazilian species described by Lutz (1914) and Macfie (1939): bicolor Lutz, discoloripes Macfie, dubia Macfie, and lacrimotorii Macfie. The remaining six previously described Neotropical species, abercrombyi Macfie, flavifemoris Macfie, seminole Wirth, squamithorax Clastrier, varipennis Wirth & Williams, and winderi Wirth, will be treated elsewhere when adequate taxonomic material is assembled. As a result of the present study a diagnosis is presented for the bicolor Group, a new synonymy is discovered, three species are redescribed and figured, and a corrected key is devised for the identification of the described Neotropical species. Explanation of the taxonomic characters used can be found in the general papers on Ceratopogonidae by Wirth et al. (1977) and Downes & Wirth (1981), and the revision of the North American Euprojoannisia by Bystrak & Wirth (1978). Forcipomyia bicolor Group Diagnosis.-Medium size to large species of Forcipomyia with legs more or less pale banded (Fig. 1). Wing length 0.7-1.5 mm. Female wing often with mottled pattern of pale areas; male wing usually extensively pale with restricted dark areas. Body with flattened scales, from long and 1-striated to broad and flattened and multi-striated. Head, body and legs with abundant semi-erect setae, these often long and bristly. Female antenna short to moderately long, segments vase-shaped to tapering; no sharp division in length between segments 10 and 11. Male antenna with plume usually yellowish

... 508 Florida Entomologist 74(4) December, 1991 BICOLOR.'-v.: ": : " v'..-:.'::.'.'.s......'..:-:..'-- : - ":-"*e '."'::" '"..!i':'?."".:'. ~! 'e ~.z' :~:.'"~ '.:'-..--..,..... @s.s*p.s j^00x@s@.ter9 ;0#@-@0 Xs B#;4g#43a@4>t0c~~~'o ":, DUBIA '!'-''': "r~:"'. *.' c'?p'*''.:-' 2X* '".-.;- N :..':;) LACRIMOTORI I,,,.,.!.,~,.,,..., ;,:,,,.,...s,.-,...e,<:,~ ~":""'"0:'"::: ABERCROMBYI g~~.,.,:::.:!:'.'-!'. F:.::..;':, ::. WINDERI E *.,,w t..-.' i FLAVIFEMORIS **^'s' WT~~~~,.; :*, :*:..;.**.*** ;^^~~~~~~,.^ P,, SEMINOLE ^.l^^^,6e,/'s, #;*'S,@5,0' t^ >, " :'.< 1 VAR I PENl I S Fig. 1. Diagram of pale and dark markings of femora (left) and tibiae) right) of (top to bottom) fore, mid, and hind legs of species of the Forcipomyia bicolor Group. apically. Palpus 5-segmrnented; third segment variably swollen in midportion, sensory pit small, shallow to deep. Female mandible without teeth. Hind tibia always with pale bands; femora and other tibiae with variable markings to unbanded. Hind tarsal ratio about 1.0. Tarsal claws curved and slender; outer claw of male fore tarsus often with

Wirth: Forcipomyia bicolor Group 509 blunt ventral tooth at midlength (Fig. 9). Female with two spermathecae. Female genital sclerotization (Fig. 7) usually an arcuate transverse ribbon bearing small spines on posterior margin, especially at lateral ends. Male genitalia (Fig. 11) often bicolored; dististyle not swollen or modified at tip. Aedeagus (Fig. 10) usually with low basal arch, more or less triangular in outline, and bearing 1-3 inconspicuous longitudinal ridges on ventral surface. Parameres not joined at base, consisting of simple, long, nearly straight rods tapering to filiform tip. Revised Key to Neotropical Species of the Forcipomyia bicolor Group (Fig. 1) 1. Femora dark from base to tip (narrow knee spots may be pale, narrow bases may be pale)...... 2 1' Femora extensively pale proximally, may be entirely pale... 5 2(1). All tibiae entirely dark; (antennae long, segments with narrow base swollen, abruptly narrowed to long slender distal portion; mesonotum with four pairs of lines of white scales)... squmithorax Clastrier 2'. At least hind tibia with broad sub-basal pale band... 3 3(2). All tibiae with pale band on proximal half... 4 3'. Fore and mid tibiae entirely dark; hind tibia with sub-basal and apical pale bands... dubia M acfie 4(3). Mesonotum with sublateral pair of compact tufts of long black setae; male aedeagus twice as long as basal breadth, basal arch to 1/5 of total length...... /acrim otorii M acfie 4'. Mesonotum without discal tufts of long black setae; male aedeagus short with broad base, total length subequal to basal breadth, basal arch to nearly half of total length... abercrombyi Macfie 5(1). Femora pale except median brown band on hind femur... seminole Wirth 5'. Femora with distal brown bands, at least on hind leg... 6 6(5). Femora dark on distal half or more; third palpal segment of female greatly swollen to past midportion, with deep pit; wing markings, tibial spur, and m ale fore claw various... 7 6'. Femora pale or with narrow basal and/or apical brown bands on mid and/or hind legs; third palpal segment of female moderately swollen with shallow pit; female wing with pale markings; spur of hind tibia short and nearly straight; male fore tarsus without ventral tooth on one claw... 8 7(6). Female wing uniformly dark brown; spur of hind tibia long, curved; one claw of male fore tarsus with blunt ventral tooth at midlength... bicolor Lutz 7'. Female wing mottled with pale markings; spur of hind tibia short and near straight; male fore tarsus without ventral tooth on one claw...... varipennis W irth & W illiam s 8(6). Fore and mid femora entirely pale; hind femur pale with broad apical brown band... wu nder W irth 8'. Fore and mid femora with narrow basal brown band; hind femur with narrow apical brown band... flavifemoris Macfie Forcipomyia (Lepidohelea) bicolor Lutz (Figs. 2-11) Forcipomyia bicolor Lutz, 1914: 89 (male; Brazil). Forcipomyia (Forcipomyia) bicolor Lutz; Wirth, 1974: 5 (catalog).

510 Florida Entomologist 74(4) December, 1991 C=Dt D_~ 72 2 723 '/_. 3 6 36 ^ ^\ "~ 6 /! 1 25 125.,~~~~~~~~~~~~ Figs. 2-11. Forcipomyia bicolor; 2, 5-7, female from Santa Catarina, Brazil (Plaumann); 3-4, 8-11, lectotype male, Santa Catarina, Brazil (Plaumann): 2, antennal segments 7-15; 3, antennal segments 12-15; 4, 6, palpus; 5, spermathecae; 7, genital sclerotization; 8, apex of tibia and basitarsus of hind leg; 9, fifth tarsomere and claws of fore leg; 10, aedeagus; 11, genitalia, aedeagus and one dististyle omitted (scale in microns). Forcipomyia discoloripes Macfie, 1939: 159 (male, female; Brazil; fig. female antenna, palpus). NEW SYNONYMY Forcipomyia (Forcipomyia) discoloripes Macfie; Wirth, 1974: 5 (catalog). A large species (wing length 1.30 mm in female) with uniformly dark brown wings in female, wing grayish with two distinct dark spots in male; femora dark brown on distal two-thirds; female palpus greatly swollen with deep sensory pit; hind tibial spur long and curved; outer claw on male fore tarsus with blunt ventral tooth at midlength. Female Plesiotype.-Wing length 1.39 mm; breadth 0.52 mm; costal ratio 0.47. Head: Dark brown including antenna and palpus; long blackish bristles arching over eyes. Antenna (Fig. 2) with lengths of flagellar segments 72-58-58-61-61-61-61-61-82-79-79-79-93 microns; antennal ratio 0.85; segments 4-10 short and somewhat flask-shaped, 11-15 longer and tapering distally; whorls of sensilla chaetica on 3-10 strong and bristle-like.

Wirth: Forcipomyia bicolor Group 511 Palpus (Fig. 6) with lengths of segments 36-36-93-3643 microns; third segment greatly swollen on proximal 0.7, with a deep sensory pit opening by asmaller pore; palpal ratio 2.00. Thorax: Dark brown; mesonotum with sparse, erect, dark brown bristles and sparse, pale brown, short appressed hairs. Legs with dark brown and yellowish brown bands as in Fig. 1; with rather long bristles, especially on tibiae, and numerous broad, appressed, striated scales. Spur of hind tibia long, curved and pubescent at base, as long as distal breadth of tibia, pale on basal half, dark distally. Tarsi dark brown with narrow pale rings at the joints, fifth tarsomeres pale; claws slender, curved, simple; hind tarsal ratio 1.00. Wing uniformly dark brown, with dense decumbent dark brown macrotrichia. Halter brownish. Abdomen: Dark brown, with dense vestiture of decumbent, dark brown, singlestriated scalelike hairs, mixed on proximal segments with broad appressed scales; last segment with long golden hairs. Genital sclerotization (Fig. 7) a narrow bowed ribbon with ends slightly broadened and bearing 2-3 slender spines, the concave posterior margin with three pairs of widely spaced, shorter, pale spines. Spermathecae (Fig. 5) ovoid with slender necks; unequal, overall measuring 86 by 53 microns and 61 by 48 microns. Male Lectotype.-Wing length 1.44 mm; breadth 0.36 mm; costal ratio 0.43. As in female with usual sexual differences. Wing with pale gray macrotrichia except two areas of darker, brownish macrotrichia, one over radial cells and the other over vein Cul. Halter pale. Legs with more extensive pale markings and fainter dark markings on fore and mid legs, their femora darker on about distal third and the dark bands on tibiae narrower. Antennal segments 12-15 (Fig. 3) with lengths 180-155-118-94 microns; plume brownish proximally, yellowish distally. Palpus as in Fig. 4; palpal ratio 2.30. Fore tarsus with outer claw (Fig. 9) broadened in midportion in a blunt tooth. Hind tibial spur (Fig. 10) greatly enlarged (length 115 microns) and curved, sickle-shaped, yellowish proximally and gradually darkened toward tip. Hind basitarsus somewhat bowed and bearing strong ventral spines; hind tarsal ratio 0.94. Genitalia (Fig. 11): Brown, base of basistyle and midportion of ninth sternum yellowish. Dististyle nearly straight. Aedeagus (Fig. 10) 1.4 times as long as basal breadth, appearing almost triangular with basal arms short and basal arch not developed, anterior margin well sclerotized, sides slightly convex; median ridge well-developed, lateral ridges displaced to lateral margins. Parameres of usual form in bicolor Group; moderately separated, the hyaline distal portions slender, slightly sinuate, and rather elongate, their tips greatly surpassing tip of aedeagus. Distribution.-Brazil, Costa Rica, Ecuador. Types.-Forcipomyia bicolor Lutz (1914):"A descricao e feita de um macho montado em preparado microscopico com gelatina glicerinada. Foi apanhado em Manguinhos num aparelho de luz." Through the kindness of Maria Luiza Felippe-Bauer, Lutz' types were borrowed from the Instituto Oswaldo Cruz in Rio de Janeiro, Brazil. These were mounted on four slides, apparently the work of Dr. A. da Costa Lima who studied the Lutz collection of Ceratopogonidae in 1937, as each slide bears the initials "C.L." 1. (left label): "Instituto Oswaldo Cruz / N. 3689 / Divisao 29 / Caixa 123." (right label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / Na Luz, Manguinhos / Prs. coll. Dr. A. Lutz / C. L." Under a large square cover slip is mounted one male cleared but with original coloration, intact but with genitalia dissected off and not present. 2. (left label): "Instituto Oswaldo Cruz / N. 3721 / Divisao 1 / Caixa 125." (right label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz o / terminalia do exemp.

512 Florida Entomologist 74(4) December, 1991 / da lam. 3689 / na Luz - Manguinhos / Dn. coll. Dr. A. Lutz / C.L." Genitalia mounted under large square coverslip. 3. (left label): "Instituto Oswaldo Cruz / N. 3691 / Divisao 1 / Caixa 124." (right label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / Na luz, Manguinhos / Da coll. A. Lutz / C.L. prep. x, 1937." Specimen apparently cleared in caustic and clorless; intact except genitalia not present. 4. (left label): "Instituto Oswaldo Cruz / N. 3690 / Divisao 30 / Caixa 123." (right label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / N. luz, Manguinhos / Da coll. A. Lutz / Do expmpl. lam. 3691 / C.L. prep x,1937." Male genitalia preparation cleared and much flattened. Slides nos. 1 and 2 as above are hereby selected as the LECTOTYPE of Forcipomyia bicolor Lutz. Types of F. discoloripes: Macfie's (1939) description is a combined description of the male and female sexes. No types were designated, but 16 male and 17 female specimens examined were listed from Nova Teutonia, Brazil, with dates from 29.vii.1936 to 17.v. 1937. These specimens may be considered syntypes. Through the courtesy of Bruce C. Townsend the British Museum (Nat. Hist.) loaned me two of Macfie's slides which were marked "Holotype male" and "Allotype female." The holotype male is hereby selected as LECTOTYPE. It bears the locality data: "Brasilien/ Nova Teutonia/ 27? 11' B 52? 23' L / Fritz Plaumann / 29,vii, 1936." It has been dissected and mounted under four coverslips and is in poor condition, but what remains agrees in all respects with the male lectotype specimen of F. bicolor Lutz, under which name F. discoloripes Macfie becomes a junior synonym (NEW SYNONYMY). Other Specimens Examined.-BRAZIL: Santa Catarina, Nova Teutonia, 7 males, 12 females collected by F. Plaumann, with dates as follows: ix. 1962, x.1962, viii. 1963, ix.1965, vii. 1966, viii-ix.1970, ix. 1970 (USNM). COSTA RICA: Puntarenas Prov., Sabalito, viii.1952, F. S. Blanton, light trap, 1 male. ECUADOR: Quevedo, Pichilingue, INAP, vi. 1978, J. Mendoza, reared from rotting vegetation, 5 males, 8 females (USNM). Forcipomyia (Lepidohelea) dubia Macfie (Figs. 12-19) Forcipomyia dubia Macfie, 1939: 162 (male, female; Brazil). Forcipomyia (Forcipomyia) dubia Macfie; Wirth, 1974: 5 (catalog). A large species (wing length 1.26 mm in female) with uniformly dark wings in female, wing whitish with two faint dark spots in male; legs dark brown, only hind tibia with pale bands; legs without broad scales; female palpus moderately swollen with deep sensory pit; hind tibial spur not modified; outer claw of male fore tarsus with blunt ventral tooth at midlength. Similar to Forcipomyia bicolor, but differing as follows: Female Plesiotype.-Wing length 1.26 mm; breadth 0.49 mm; costal ratio 0.43. Head: Antennae (Fig. 12) with lengths of flagellar segments 72-65-61-61-61-61-61-61- 65-65-72-72-90 microns; antennal ratio 0.78. Palpus (Fig. 15) with lengths of segments 25-42-97-36-43 microns; third segment moderately swollen, with deep sensory pit; palpal ratio 2.24. Thorax: Legs (Fig. 1) dark brown, narrow knee spots yellowish; all femora and fore and mid tibiae entirely dark brown, hind tibia with two prominent pale yellowish bands as in bicolor. Hind tibial spur slender, length 75 microns. Legs with abundant long, narrow, appressed one-striated scales, broad flat scales absent. Hind tarsal ratio 1.10. Wing uniformly dark brown as in bicolor, with abundant long dark-brown macrotrichia. Halter brownish.

Wirth: Forcipomyia bicolor Group 513 > 6< 6i DC 9 12 13</ 3 '.(Wt\ 0 ( )? w"''$l15 72 14 Figs. 12-19. Forcipomyia dubia: 12-13, 15-16, female from Santa Catarina, Brazil (Plaumann); 14, 17-19, lectotype male, Santa Catarina, Brazil (Plaumann): 12, antennal segments 7-15; 13, spermathecae; 14, apex of hind tibia; 15, palpus; 16, genital sclerotization; 17, fifth tarsomere and claws of fore leg; 18, aedeagus; 19, genitalia, aedeagus and one dististyle omitted (scale in microns). Abdomen: Spermatheca (Fig. 13) larger, apparently subequal (one tipped in orientation with neck hidden), slightly ovoid, 94 by 72 microns. Genital sclerotization (Fig. 16) as in bicolor but lateral spines short and relatively stout. Male Lectotype.-Wing length 1.61 mm; breadth 0.47 mm; costal ratio 0.42. Head damaged (measurements from plesiotype: antennal segments 12-15 with lengths 234-165- 119-119 microns; palpal segments with lengths 36-43-119-47-40 microns, palpal ratio 4.10). Hind tarsal ratio 0.93; hind tibial spur (Fig. 14) 79 microns long. Fore tarsus with blunt ventral tooth on outer claw as in bicolor (Fig. 17). Wing with pale macrotrichia except two dark spots of black macrotrichia, one over radial cells and other on vein Ml. Halter pale. Genitalia (Fig. 19): Brown, basal half of basistyle, median portion of ninth sternum, and all except tip of dististyle yellowish. Dististyle slender and curved. Aedeagus (Fig. 18) nearly triangular in ventral view, 1.5 times as long as basal breadth, apex acutely pointed; basal arch low but distinct, anterior margin well sclerotized; a median ridge and two lateral ridges well marked. Parameres as in bicolor Group; bases widely separated, distal portions hyaline, slender, reaching tip of aedeagus. Distribution.-Brazil. Types.-Syntypes, 2 males, Nova Teutonia, Brazil, 16 and 17.ix.1936 (BMNH). I have studied the two syntypes loaned through the courtesy of Bruce Townsend and the British Museum (Nat. Hist.). The specimen with date 16.ix.1936 had been mounted on

514 Florida Entomologist 74(4) December, 1991 a slide under two coverslips, and bears a blue circular label "Syntype" and a label with red border printed "Type." This male is hereby designated LECTOTYPE of Forcipomyia dubia Macfie. The second male, with date 17.ix.1936, is pinned and badly damaged, consisting essentially of the thorax and one wing, with the genitalia mounted on a plastic strip preparation on the pin below the specimen. Other Specimens Examined.-BRAZIL: Santa Catarina, Nova Teutonia, ix. 1962, F. Plaumann (BMNH slide 1966-590), 1 female (Plesiotype). One male (Plesiotype), same data but date viii-ix.1970, 1 male (USNM). Forcipomyia (Lepidohelea) lacrimotorii Macfie (Figs. 20-20) Forcipomyia lacrimotorii Macfie, 1939:161 (male, female; Brazil; fig. antenna, palpus). Forcipomyia (Forcipomyia) lacrimotorii Macfie; Wirth, 1974; 5 (catalog). A moderately large species (wing length 0.99 mm in female) with uniformly dark brownish wings in both sexes; mesonotum in both sexes with a prominent pair of tufts of long black setae; legs dark brown, tibiae with sub-basal pale bands and hind tibia with apex broadly pale; legs with abundant broad striated scales; female palpus slightly swollen with small shallow sensory pit; hind tibial spur not modified; outer claw of male fore tarsus with blunt ventral tooth at midlength. Very similar to Forcipomuyia dubia Macfie, differing as follows: Female Plesiotype.-Wing length 0.99 mm; breadth 0.39 mm; costal ratio 0.42. Head: Antenna (Fig. 20) with lengths of segments 57-61-65-65-65-65-65-65-61-65-65-72-86 microns; antennal ratio 0.69; all segments with bulbous bases, then narrowing and slightly tapering distally, less so on distal five segments. Palpus (Fig. 21) slender; segments with lengths 29-36-79-24-29; third segment slightly swollen on proximal half, palpal ratio 3.10; sensory pit small and shallow. Thorax: Dark brown; mesonotum with sparse long erect black bristles and abundant small appressed flattened whitish hairs; a pair of dense tufts of moderately long black hairs sublaterally at midlength, about 10 hairs in each tuft. Legs (Fig. 1) dark brown, including all femora; knees with narrow pale ring on tibiae; tibiae with moderately narrow sub-basal pale ring and hind tibia with moderately broad apex pale; femora and tibiae with abundant broad, striated, appressed scales. Hind tibial spur (Fig. 23) 65 microns long, straight and slender. Wing dark brown, slightly shaggy appearance due to abundant long scalelike decumbent macrotrichia, especially prominent and dense over radial veins. Halter brownish. Abdomen: Spermathecae (Fig. 24) small, deeply pigmented; ovoid with scarcely apparent necks; slightly unequal, measuring 61 by 44 microns and 52 by 40 microns. Genital sclerotization a slender, bowed, transverse ribbon as in bicolor, but ends not expanded and with no well-developed spines on posterior margin. Male Plesiotype.-Wing length 1.36 mm; breadth 0.40 mm; costal ratio 0.42. Head brown; antennal plume brown at base, yellowish distally. Antennal segments 12-15 with lengths 161-169-130-122 microns. Palpal segments with lengths 29-36-100-29- 30 microns; palpal ratio 1.60. Mesonotum with pair of prominent dark brown hair tufts as in female. Hind tibial spur 90 microns long, straight and slender. Hind tarsal ratio 1.08. Fore tarsus with blunt ventral tooth on outer claw as in bicolor and dubia (Fig. 22). Wing brownish, paler than in female but darker along veins. Halter pale. Abdomen dark brown; with abundant, semi-erect, long, single-striated scalelike hairs. Genitalia (Fig. 26): Brown; proximal half of basistyle yellowish. Dististyle straight, relatively stout, tapered to pointed tip. Aedeagus (Fig. 25) shorter and broader than

Wirth: Forcipomyia bicolor Group 515 72 20 72 21 72 23 29 22 72 4 0 0 Figs. 20-26. Forcipomyia lacrimotorii; 20-21, 24, female; 22-26, male; from Santa Catarina, Brazil (Plaumann): 20, antennal segments 8-15; 21, palpus; 22, fifth tarsomere and claws of fore leg; 23, apex of hind tibia and basitarsus of hind leg; 24, spermathecae; 25, aedeagus; 26, genitalia, aedeagus and one dististyle omitted (scale in microns). in bicolor and dubia, 1.4 times as long as basal breadth, basal arms and basal arch well developed, anterior margin well sclerotized; sides slightly convex, tapering to blunt apex; three well-marked ventral ridges at base. Parameres typical of bicolor Group; bases well separated, distal portions straight, slender, tips slightly surpassing tip of aedeagus. Distribution.-Brazil. Types.-Syntypes, 10 males, 13 females, Nova Teutonia, Santa Catarina, Brazil, F. Plaumann, dates from 13.v.1937 to 7.ix.1938 (BMNH). Through the courtesy of Bruce Townsend I have studied one male and two female syntypes from the British Museum (Nat. Hist.). They are mounted on one slide with the following labels: "Brasilien / Nova Teutonia / 27? 11' B 52? 23' L / Fritz Plaumann / 13.5.37." and "Forcipomyia / lacrimotorii / o o; SYNTYPE" in blue-bordered circular label and "PARATYPES" in red-bordered blue label. The two females are not dissected but mounted whole on their sides under one large coverslip. One of these is in excellent condition; the other is damaged and has the

516 Florida Entomologist 74(4) December, 1991 antennae mounted separately under a small square coverslip. The male is mounted under a separate small square coverslip and the genitalia and legs from one side have been dissected off. The head is missing. This male is hereby designated LECTOTYPE. I have also examined three male and three female pinned syntypes with data as above. These are in moderately good condition, some with mesonotum somewhat rubbed and vestiture missing. The characteristic mesonotal hair tufts are present on the better specimens. Other Specimens Examined.-Seven males, 10 females, same data as types but dates viii. 1945, v. 1963, viii. 1963, viii-ix.1970, ix.1970, i. 1971, ii.1971 (USNM). Specimens from this series have been selected as PLESIOTYPES and described and illustrated. ACKNOWLEDGMENTS I wish to express special appreciation to Maria Luiza Felippe-Bauer of the Instituto Oswaldo Cruz in Rio de Janeiro, Brazil, and to Bruce C. Townsend of the Natural History Museum, London, U.K. (BMNH) for the loan of types without which this study would not have been possible. Most of the Brazilian specimens for this study were purchased by the author from Fritz Plaumann of Nova Teutonia, Santa Catarina, Brazil, many years ago and deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Financial support for this publication by Mars Incorporated, Effem Services Inc., Information Services International Division, Mt. Olive, NJ, is gratefully acknowledged. REFERENCES CITED BILLES, D. J. 1941. Pollination of Theobroma cacao L. in Trinidad, B.W.I. Trop. Agric. (Trinidad) 18: 151-156. BYSTRAK, P. G. AND W. W. WIRTH. 1978. The North American species of Forcipomyia, subgenus Euprojoannisia (Diptera: Ceratopogonidae). U.S. Dept. Agric. Tech. Bull. 159: 1-51. DEBENHAM, M. L. 1987. The biting midge genus Forcipomyia (Diptera: Ceratopogonidae) in the Australasian Region (exclusive of New Zealand) III. The subgenera Forcipomyia s. s., and Lepidohelea. Invertebr. Taxon. 1: 167-199. DOWNES, J. A., AND W. W. WIRTH. 1981. Ceratopogonidae, pp. 393-421, in McAlpine, J. F., et al. [eds.], Manual of Nearctic Diptera, Vol. 1. Agric. Canada Monogr. no. 27: 674 pp. KAUFMANN, T. 1975a. Ecology and behavior of cocoa pollinating Ceratopogonidae in Ghana, W., Africa. Environ. Entomol. 4: 347-351. KAUFMANN, T. 1975b. Studies on the ecology and biology of a cocoa pollinator, Forcipomyia squamipennis I. & M. (Diptera, Ceratopogonidae), in Ghana. Bull. Entomol. Res. 65: 263-268. LUTZ, A. 1914. Contribuicao para o conhecimento das Ceratopogoninas do Brazil. Terceira memoria. Aditamento terceiro e descricao de especies que nao sugam sangue. Mem. Inst. Oswaldo Cruz 6: 81-99, 2 plates. MACFIE, J.W.S. 1939. A report on a collection of Brazilian Ceratopogonidae (Dipt.). Revta. Entomol. 10: 137-219. MACFIE, J.W.S. 1944. Ceratopogonidae collected in Trinidad from cacao flowers. Bull. Entomol. Res. 35: 297-300. POSNETTE, A. F. 1950. The pollination of cacao in the Gold Coast. Hort. Sci. J. 45: 155-163. SAUNDERS, L. G. 1956. Revision of the genus Forcipomyia based on characters of all stages (Diptera, Ceratopogonidae). Canadian J. Zool. 34: 657-705. SAUNDERS, L. G. 1959. Methods for studying Forcipomyia midges, with special reference to cacao-pollinating species (Diptera, Ceratopogonidae). Canadian J. Zool. 37: 33-51.

Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 517 SORIA, S. DE J., AND W. W. WIRTH. 1974. Identidade e caracterizacao taxonomica preliminar das mosquinhas Forcipomyia (Diptera, Ceratopogonidae) associadas comrn a polinazacao do cacaueiro na Bahia. Revta. Theobroma 5(54): 3-22. WINDER, J. A. 1977. Field observations on Ceratopogonidae and other Diptera: Nematocera associated with cocoa flowers in Brazil. Bull. Entomol. Res. 67: 57-63. WINDER, J. A. 1978. Cocoa flower Diptera: their identity, pollinating activity and breeding sites. PANS 24: 5-18. WIRTH, W. W. 1974. A catalogue of the Diptera of the Americas south of the United States. 14. Family Ceratopogonidae. Mus. Zool. Univ. Sao Paulo 14: 1-89. WIRTH W. W. 1991. New and little-known species of Forcipomyia (Diptera: Ceratopogonidae) associated with cocoa pollination in Brazil. Proc. Entomol. Soc. Washington. 93: 163-175. WIRTH, W. W., N. C. RATANAWORABHAN, AND D. H. MESSERSMITH. 1977. Natural History of Plummers Island, Maryland. XXII. Biting midges (Diptera: Ceratopogonidae). 1. Introduction and key to genera. Proc. Biol. Soc. Washington 90: 615-647. YOUNG, A. M. 1983. Seasonal differences in abundance and distribution of cocoa-pol- linating midges in relation to flowering and fruit set between shaded and sunny habitats of the La Lola cocoa farm in Costa Rica. J. Appl. Ecol. 20: 801-831. FIVE NEW SPECIES OF APHODIUS (COLEOPTERA: SCARABAEIDAE) FROM FLORIDA POCKET GOPHER BURROWS PAUL E. SKELLEY Department of Entomology & Nematology Building 970, Hull Road University of Florida Gainesville, FL 32611-0740 ROBERT E. WOODRUFF Florida State Collection of Arthropods P. 0. Box 147100 Gainesville, FL 32614-7100 ABSTRACT Five new species of Aphodius (dyspistus, tanytarsus, hubbelli, platypleurus, and pholetus), collected in the burrows of pocket gophers (Geomys pinetus Rafinesque), are described. Similarities between Florida and Great Plains pocket gopher burrow faunas are discussed, and reasons for the diversity of Aphodius in this habitat are postulated. RESUMEN Se describen cinco nuevas especies de Aphodius (dyspistus, tanytarsus, hubbelli, platypleurus, y pholetus), colectadas en las galerias de Geomys pinetus Rafinesque. Se discuten las similitudes entre la fauna de las galerias del genero Geomys de Florida y de las Grandes Planicies. Se postulan las razones de la diversidad de Aphodius en estos habitats.