(BIG-HEADED ) 54

Jose Albertino Rafael and Jeffrey H. Skevington

Fig. 54.1. Male of luteicornis Cresson, (Nearctic, MND, fig. 53.1).

Diagnosis Abdomen subcylindrical, coloration and tomentum similar to thorax, occasionally with yell ow markings. Tergite 1 usu­ Adu lts (Fig. 1) small (body length 2.0- 11.5 mm), dark. ally with row or cluster of bristles laterally. Male terminalia Head semiglobose with narrow occiput in Chalarinae or globose twisted left (Fig. 17). Male syntergosternite 8 large, forming with broadened occiput in Nephrocerinae and . tip of abdomen (Fig. 17). Female terminali a modified into Compound eyes occupying most of head (Figs. 1--4). Female piercing ovipositor (Fig. 16). with enlarged anterior ommatidia (not distinct in ). Pipunculids can be differentiated from syrphids by the Antenna with first flagellomere appearing as single segment, large compound eyes that occupy most of their globose or + obtuse to fi li form in shape, with dorsal arista. Thorax with semiglobose head, lack of spurious vein, and open r4 5 cell. darkish ground color. Proepisternum sometimes bearing fan­ A few bombyliids such as Metacosmus Coquillett resemble like tuft of bristles (Fig. 3). Claws and pulvilli larger in pipunculids when encountered in the fie ld, but these round­ females. Wings long and narrow, usually hyaline, iridescent headed bombyliids can be easily differentiated by their + in direct light, pterostigma present or absent, cell r4 5 open. antennae that lack the dorsal arista found in pipunculids.

793 794 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME 2

Biology Region (Skevington & Marshall, 1997) and the Australian Region (Skevington, 2001). Most New World pipunculid Most pipunculids that have been studied are endoparasitoids have no biological data available. In the Neotropical of several Hemiptera () families, including Region and Mexico, the following papers treat pipunculid­ Cicadellidae, Cercopidae, Delphacidae, Membracidae, Issidae, host associations: Hardy (1963) for southern Colombia and Cixiidae, and Flatidae. The only known exception occurs in northern Ecuador, Vega et al. (1991) for Mexico, Virla & Ra­ the genus Nephrocerns. These unusual flies attack adult crane fael (1996) for Argentina, and recently Virla et al. (2009) for flies (Tipulidae) (Koenig & Young, 2007). The potential val­ all Neotropical species. ue of Pipunculidae for biological control has stimulated some Most pipunculid species attack more than one species of work on the bionomics of this family. For example, research host, but show a preference for a particular set of host spe­ into the control ofthe potato leafi1opper, Empoascafabae (Har­ cies. Only two species have been reared from more than one ris), a major pest of alfalfa in midwestem and eastern USA and subfamily of host (Skevington, 2001). Superparasitism and Canada, involved exploration in Europe for natural enemies to multiple parasitism are rare, and survival of more than one be introduced into the USA (Jervis, 1992) and included rear­ pipunculid in these cases is exceptional (Jervis, 1980; Mora­ ing of Chalarns for release (Skevington & Marshall, 1997). kote & Yano, 1988; Waloff & Jervis, 1987). Despite the economic importance of the group, few data on host relationships exist, and some genera have no known hosts Classification (for example Allomethus, Basileunculus, Elmohardyia, and Protonephrocerns ). Pipunculidae are considered to be a monophyletic lineage Adults are frequently seen hovering among vegetation, and and sister group of the flower flies (Syrphidae). Together, their flight is similar to that of Syrphidae. Adults utilize all ter­ pipunculids and syrphids are grouped in the monophyletic restrial habitats, but diversity and numbers are greatest in forest c1ade , sister group of (Cumming openings and along forest edges. In semi-arid regions, adults are et aI., 1995). Worldwide, 140 I valid species of pipunculids commonly found at seeps or along small creeks. Adult flies feed are recognized, and it is likely that over 2700 species exist on honeydew and can occasionally be found in large, mixed (De Meyer, 1996; De Meyer & Skevington, 2000; J. H. Skev­ feeding groups on Auchenorrhyncha secretions (Skevington & ington, unpublished data). Twenty-one extant genera are Marshall, 1998). Because of this behavior, they are often at­ currently recognized in three subfamilies: Chalarinae, Neph­ tracted to artificial honeydew sprayed on vegetation (a mixture rocerinae, and Pipunculinae (De Meyer, 1996; De Meyer & of honey, Coca-Cola®, and water). Many pipunculids employ Skevington, 2000; Rafael & De Meyer, 1992; Skevington & the landmark mating system known as hilltopping (Skevington, Yeates, 200 I). The latter are divided into five tribes: Pipun­ 2000,2001). Hilltop collecting is an excellent way to sample culini, Cephalopsini, Microcephalopsini, Eudorylini, and much of the pipunculid diversity in an area and often provides Tomosvaryellini. Molecular analyses made by Skevington & large numbers of specimens. Other effective methods frequent­ Yeates (2000) corroborated most of Rafael & DeMeyer's ly employed to capture pipunculids include sweeping, Malaise (1992) phylogenetic results, except for the position of the trapping, and less successfully, using yellow pan traps. Occa­ Nephrocerinae genera (Nephrocerus and Protonephrocerus). sionally, mating pairs are discovered hovering in copula or tem­ The combined molecular data suggested that Nephrocerlls porarily settled on vegetation. Pronounced sexual dimorphism is a basal lineage of subfamily Chalarinae, and Protoneph­ in pipunculids makes association of the sexes difficult, so these rocerus is a basal lineage of subfamily Pipunculinae. When occasional discoveries are useful. these data were combined with the morphological data set Females of most pipunculid species deposit their eggs in ear­ ofRafael & De Meyer (1992), Nephrocerus and Protoneph­ ly instar nymphs or adults of their hosts. After visually locating rocerns appeared between the basal Chalarinae and the ter­ a host, some species remove the host from the food plant and minal Pipunculinae, reflecting the hypothesis of Rafael & oviposit one egg inside each host while in flight (Huq, 1982; De Meyer with the difference that these two genera do not May, 1979; Williams, 1919), whereas others leave the host form a monophyletic group. These results suggest that the in situ (Benton, 1975). Larvae develop to maturity within the subfamilial ofthe pipunculids may need modifica­ nymph or adult hosts. When mature, the leaves the host tion and further study. Generic concepts are relatively well through a break in one of the dorsal intersegmental membranes resolved but more work is still needed to clarify Eudorylini of the abdomen in order to pupate in the soil, in leaf litter, at genera. Despite recent work on Eudorylini, problems remain the base of plants, or rarely, attached to leaves (Ferrar, 1987; within this large lineage (Skevington & Yeates, 2001). Perkins, 1905). More detailed information on immature stages together with information on rearing techniques can be found in Benton (1975), Coe (1966), Ferrar (1987), Freytag (1988), Huq Identification (1984), Jervis (1980), Morakote & Yano (1987), and Perkins (1906). Because of their small size, black coloration, and the pau­ Pipunculid- host associations are best known for the Euro­ city of external characters, pipunculid species are difficult pean fauna (Waloff & Jervis, 1987), followed by the Nearctic to identify. It is particularly important to have male and PIPUNCULlDAE (BIG-HEADED FLIES) 54 795 fema le specimens from the same sample in order to associate In order to secure the ground color and tomentum of pip un­ the sexes. For secure identification, it is necessary to dissect culids, specimens should ultimately be dried and pinned. Pin­ the male terminalia. To accomplish this, remove the abdo­ ning fresh material killed with cyanide or ethyl acetate is the men and heat in 85% lactic acid for at least 30 minutes. Re­ easiest method, and a number of techniques can be used. The move the macerated abdomen and place it directly into clean best method for obtaining views of the 's genitalia in­ glycerin. Dissection involves separating syntergostemite 8 volves double mounting. Pinning through the right anepis­ from the remainder of the abdomen. Store the abdomen and temum with a minuten pin, then pinning to pith is more dissected terminalia in glycerin in plastic microvials on the expensive and time consuming than pointing, but produces same pin as the source specimen. Surstyli shape, presence or excellent specimens that are less likely to require dissection absence of a membranous area on syntergostemite 8, phal­ than other pinning techniques. Material collected into alcohol lic guide shape, phallus shape and ramification, sperm pump is preferably dried with a critical point drier and then pointed shape and sclerotization, ejaculatory apodeme shape, hypan­ or glued directly to the side of a pin. Chemical drying using drium shape, and modifications of the stemites are often in­ HMDS (hexamethyldisilazane) or ethyl acetate is also accept­ formative. Tn female specimens, the ovipositor shape is often able. Although some researchers prefer to study pipunculids useful, but few other diagnostic characters are available. preserved in 70% alcohol, specimens preserved in this way Description of new species on the basis offemales often com­ discolor over time and become brittle and difficult to dissect. plicates the taxonomy of the family and should be avoided Terminology used to describe pipunculid morphology unless new character systems are discovered for females. We has varied considerably over time and between authors, but encourage DNA barcoding (sequencing cytochrome oxidase recent efforts have been made to standardize this language. f) of new species of pipunculids to faciliate association of Most of the current morphological terminology is illustrated sexes and larvae and to stabilize nomenclature across wider here (Figs. 1- 37), but see Skevington & Yeates (2001) for a geographical areas. more thorough treatment of this topic.

Key to the genera of Pipunculidae of Central America and Nearctic Mexico (modified from Albrecht, 1990; Rafael, 1996; Skevington & Yeates, 2001)

I. Hind margin of eye deeply excised medially (Fig. 4); first f1agellomere reniform (Fig. 4); NEPHROCERINAE ...... N ephrocerus Zetterstedt Hind margin of eye straight or nearly so (Figs. 2, 3, 8); first f1agellomere acute (Fig. 5), filiform (Fig. 6) to aristiform ...... 2

2. Ocellar bristles distinct; occiput narrow, scarcely projecting behind eyes (Fig. 8); head hemispherical (Fig. 8); margin of scutum and scutellum with strong bristles (Fig. 8); CHALARINAE ...... 3 Ocellar bristles reduced or absent (Figs. 2, 3, 7); occiput swollen and plainly visible in lateral view (Figs. 2, 3, 7); head spherical (Figs. 2, 3, 7); margin of scutum and scutellum without strong bristles (Fig. 7); PIPUNCULTNAE ...... 5

3. Wing venation incomplete, cross vein dm-cu absent, vein M reduced (Fig. 9) .. Walker Wing venation complete, M and dm-cu present (as in Fig. 12) ...... 4

4. M2 present (Fig. 11) ...... Verrallia Mik M2 absent (as in Fig. 13) ...... Jassidophaga Aczel

5. Proepistemum with fanlike tuft of bristles (Fig. 3) ...... 6 Proepistemum bare ...... 11

6. At least anterior portion of scutum evenly setose (Fig. I) ...... Pipunculus Latreille Scutum bare except for differentiated rows of dorsocentral bristles (Fig. 7) ...... 7 796 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME 2

2 Collinias d' 3 Cephalosphaera d'

7 Dasydorylas d'

6 Eudorylas d' 8 Jassidophaga d' c Se

10 Collinias d' plslg

\;", . ~~ c I'tt.l'\ ~ ..... \ '\ '''It,~ ...... ,' .,,' ... ,", ...... ,...... ",,,,,,,...... ,, .. ,,, ' ..... "..... ~ A1 +cuA2 dm-eu 2 II Verrallia d' 12 Pipunculus d'

13 d' 14 d'

c..--"'- - - ~«Z\\Z~t-==:~-~- ~ --:: ~ .... 15 Cephalops d'

Figs. 54.2-15. Heads, wings, and associated structures: lateral view of head of(2) Collinias sp., (Australasian, Skevington & Yeates, 2001, fig. 2B); dorsolateral view of head and portion of thorax of (3) Cephalosphaera par/henopipis Skevington, (Australasian, Skevington & Yeates, 200 I, fig. 21); lateral view of head of (4) Nephrocerus slossonae Johnson, (Nearctic, MNO, fig . 53.7); lateral view of antenna of (5) Elmohardyia doelloi (Shannon), (South America, Rafael, 1987b, fig. 9); (6) Eudorylas mu/ilia/us (Loew), (Australasian, Skevington & Yeates, 2001, fig. 2A); dorsal view of head, thorax, and base of abdomen of (7) Dasydorylas eucalyp/i (Perkins), (Australasian, Skevington & Yeates, 200 I, fig. 3A); same, with head rotated almost 90° to right, of (8) Jassidophaga sp., (Australasian, Skevington & Yeates, 200 I, fig. 3C); dorsal view of wing of (9) Chalarus sp., (Australasian, Skevington & Yeates, 200 I, fig. 40); (10) Collinias sp., (Australasian, Skevington & Yeates, 2001, fig. 4C); (11) Verrallia virginica Banks, (Nearctic, MNO, fig. 53.3); (12) Pipunculusfuscus Loew, (Nearctic, MNO, fig. 53.6); (13) Dorylomorpha exilis (Ma \loch), (Nearctic, MNO, fig. 53.4); (14) Tomosvaryella sylva/ica (Meigen), (Holarctic, MND, fig . 53.5); dorsal view of hind leg of (15) Cephalops cochlea/us De Meyer, (Australasian, Skevington & Yeates, 200 I, fig . 3F). Abbreviations: ocp, occiput; prepst s, proepisternal setae; ptstg, pterostigma. PIPUNCULlDAE (BIG-HEADED FLIES) 54 797

7. Vein M2 present (as in Fig. 11); Cephalosphaera Enderlein ...... 8 Vein M2 absent (as in Fig. 13) ...... 9

8. Hind tibia with comb oflonger setae apically; male abdominal syntergostemite 8 with membranous area reaching epandrium (Fig. 23) ...... Cephalosphaera (Neocephalosphaera) De Meyer Hind tibia without comb of longer setae apically; male abdominal syntergostemite 8 with mem­ branous area not reaching epandrium (Fig. 22) ... Cephalosphaera (Cephalosphaera) Enderlein

9. Face narrower than lower portion of frons; frons inflated in lateral view (as in Fig. 2); first f1agellomere small, slightly larger than pedicel (as in Fig. 2); cell dm not expanded medially (as in Fig. 10) ...... Microcephalops De Meyer Face equal or subequal in width to lower portion of frons; frons not inflated in lateral view; first f1agellomere distinctly larger than pedicel (as in Fig. 3); cell dm expanded medially (as in Fig. 13); Cephalops Fallen ...... 10

10. Tibiae lacking strong median and apical spinelike setae; abdomen broad and shortened; male abdominal syntergostemite 8 with membranous area reaching epandrium (as in Fig. 23) ...... Cephalops (Semicephalops) De Meyer Tibiae with strong median and apical spinelike setae (Fig. 15); abdomen long and narrow; male abdominal syntergostemite 8 with membranous area not reaching epandrium (as in Fig. 22) ...... Cephalops (Cephalops) Fallen

11. Pterostigma absent (Figs. 13 , 14) ...... 12 Pterostigma present (Figs. 9-12); Eudorylini (only males identifiable to genus) ...... 16

12. Crossvein r-m meeting cell dm near middle (Fig. 14); abdominal tergites usually with at least some grey or brown tomentum laterally; abdomen shorter and not distinctly clavate, usually widest at middle ...... Tomosvaryella Aczel Crossvein r-m meeting cell dm near basal third (Fig. 13); abdominal tergites (except tergite 1) shining, entirely without tomentum; abdomen elongate and clavate, widening distally; Dorylomorpha Aczel ...... 13

13 . Males ...... 14 Females ...... 15

14. Frons at narrowest point at least as wide as anterior ocellus; inner lobe of surstylus with ventral projection (Figs. 36b, c); apices of surstyli usually dilated (normally visible without dissection) (Fig. 36a) ...... Dorylomorpha (Dorylomyia) Albrecht Frons at narrowest point less than half as wide as anterior ocellus or eyes contiguous; inner lobe of surstyli without ventral projection (Figs. 37b, c); apices of surstyli not dilated (Fig. 37a); Neotropical (Colombia, Brazil) and Holarctic Regions (USA: south to Colorado and Utah in Nearctic Region) ...... [Dorylomorpha (Pipunculina) Albrecht]

15 . Piercer without lateral flanges (Fig. 18) or if distinct flanges present then abdomen short and robust in form ...... Dorylomorpha (Dorylomyia) Albrecht Piercer of ovipositor with distinct lateral flanges (Fig. 19); abdomen long and slender; Neotropical (Colombia, Brazil) and Holarctic Regions (USA: south to Colorado and Utah in Nearctic Region) ...... [Do rylomorpha (Pipunculina) Albrecht]

16. Tergite 6 visible dorsally in males (Fig. 20) ...... 17 Tergite 6 not visible dorsally in males (Fig. 21) ...... 18 798 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME 2

spr 6 tg 6 tg 7

spr 7 hypd tg 7

tg 8 pmp "----tg9 17 Dorylomorpha 0' cere

16 Pipunculus Q

18 Dorylomorpha (Dorylomyia) Q 19 Dorylomorpha (Pipunculina) Q 20 Elmohardyia 0'

syntgst 8

tg1 tg2 tg3 tg4 tg5 22 Cephalosphaera (Cephalosphaera) 0' 21 Allomethus 0' syntgst 8

mem

23 Cephalosphaera (Neocephalosphaera) 0'

Figs. 54.16-23. Abdomens and terminalia: lateral view of female abdomen of (16) Pipunculus torus Skevington, (Nearctic, MND, fig. 53.9); lateral of male abdomen of (17) DorylomOlpha canadensis Hardy, (Nearctic, MND, fig. 53.8); ventral view of ovipositor of (18) D. (Dorylomyia) beckeri Aczel, (Holarctic, Albrecht, 1990, fig. 44); (19) D. (Pipunculina) borealis Wah1gren, (Holarctic, Albrecht, 1990, fig . 62); dorsal view of male abdomen of (20) Elmohardyia atlantica (Hough), (Nearctic, Skevington & Yeates, 200 I, fig. 4J); (21) A llomethus rotundicornis (Hardy), (Neotropical, Skevington & Yeates, 2001, fig. 4M); ventral view of male terminalia of(22) Cephalosphaera (Cephalosphaera) prionotaina Skevington, (Australasian, Skevington, 1999, fig. 5D); (23) Ce. (Neocephalosphaera) pelila Skevington, (Australasian, Skevington, 1999, fig. 4e). Abbreviations: cere, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; mem, membranous area; ovp, ovipositor; ph, phallus; spr, spiracle; spm pmp, sperm pump; st, sternite; sur, surstylus; syntgst, syntergosternite; tg, tergite. PIPUNCULlDAE (BIG-HEADED FLIES) 54 799

ph

25 Eudorylas d'

26 Eudorylas d'

hypd 27 Basileuneulus d' hypd

28 Basileuneulus d'

29 Allomelhus d' 30 Allomelhus d' 31 Allomelhus d'

32 Elmohardyia d' 33 Elmohardyia d' 34 Elmohardyia d'

35 Elmohardyia d'

Figs. 54.24-35. Male terminalia: dorsal view of male terminalia, phallus removed, of(24) Eudorylas moffattensis Skevington, (Australasian, Skevington & Yeates, 200 I, fig. 61); lateral view of phallus and phallic guide of (25) Eu. sehreileri (Shannon), (South America, Rafael, 1987b, fig. 6); lateral (a) and dorsolateral (b) views of ejaculatory apodeme and sperm pump of (26) Eu. mulillalus (Loew), (Australasian, Skevington & Yeates, 200 I, figs. 8E-F); dorsal view of surstyli, phallic guide, and hypandrium of (27) Basileuneulus rex (Curran), (Skevington & Yeates, 2001, fig. 6E); dorsal view of phallus, sperm pump, and ejaculatory apodeme of (28) B. rex, (Skevington & Yeates, 2001, fig. 7J); lateral view of male sperm pump and ejaculatory apodeme of (29) A. brimleyi Hardy, (Nearctic, Rafael, 1990b, fig. 14); dorsal view of male surstyli , phallic guide, and hypandrium of (30) A. rotundieornis Hardy, (Neotropical, Skevington & Yeates, 200 I, fig. 6F); dorsal view of phallus of (31) A. brimleyi Hardy, (Nearctic, Skevington & Yeates, 200 I, fig. 8C); lateral (a) and dorsal (b) views of male ejaculatory apodeme and sperm pump (32) and dorsal view of male terminalia, phallus removed (33) of Elmohardyia allanliea (Hough), (Nearctic, Skevington & Yeates, 2001, figs. 8P-Q and fig. 7C); right lateral view of phallus of (34) El. doelloi (Shannon), (South America, Rafael, 1987b, fi g. 13); right lateral view of phallus of(35) El. alianliea (Hough), (Nearctic, Skevington & Yeates, 2001, fig . 71). Abbreviations: ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; ph, phallus; ph gd, phallic guide; sbepand scl, subepan­ drial sclerite; spm pmp, sperm pump; st, stemite; sur, surstylus; syntgst, syntergostemite. 800 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME 2

17. Apex of first flagellomere obtuse below (Fig. 5); ventral spinelike setae (ctenidium) present on midfemur only; at least tergite 5 with oblique gray tomentose spots posterolaterally; phallus simple, apically swollen and membranous (Figs. 34, 35); sperm pump usually elongate with square distal process (Fig. 32) ...... Elmohardyia Rafael Apex of first flagellomere acuminate (Fig. 6) to subfiliform or aristiform in some species; ventral spinelike setae present on forefemur and midfemur; tergites without oblique gray tomentose spots; phallus trifid (Fig. 25); sperm pump usually elongate with two distal processes (Fig. 26) ...... Eudorylas Aczel, in part

18. Abdomen with small, inconspicuous bristles (as in Fig. 20) ...... Eudory las Aczel, in part Abdomen with conspicuous bristles, notably laterally (Fig. 21) ...... 19

19. Distal section of vein M\ relatively straight (as in Fig. 12); phallus trifid, with hooks at apices and carpet of setae laterally (Fig. 31); ejaculatory apodeme linear (Fig. 29) ...... Allomethus Hardy Distal section of vein M\ curved (as in Fig. 13); phallus simple and heavily sclerotized, swollen distally (Fig. 28); ejaculatory apodeme shaped li ke ball on stem (Fig. 28) ...... Basileunculus Rafael

36 Dorylomorpha (Dorylomyia) 0' 37 Dorylomorpha (Pipunculina) 0'

Figs. 54.36-37. Male surstyli : (a) ventral view of both surstyli, (b) left lateral view of left surstylus, and (c) right lateral view ofright surstylus of (36) Dorylomorpha (DOfylomyia) beckeri Aczel, (Holarctic, Albrecht, 1990, fig. 47) and of (3 7) D. (Pipun culina) borealis Walhgren, (Holarctic, Albrecht, 1990, fi g. 64).

Synopsis of the fauna studied, and only 14 genera and approximately 90 species are known. Pipunculids are so poorly sampled that many new The following synopses are based on De Meyer (1996), species can be expected, and it is likely that there are well Rafael & Menezes (1999), De Meyer & Skevington (2000), over 200 species. As stated above, pipunculids are found in and unpublished material from the Canadian National almost all terrestrial habitats, including canopy, lowland rain Collection of (CNC), the National Museum of forest, dry forests, open areas, riparian regions, and woodland Natural History (USNM), and the Instituto Nacional de edges, being more diverse in wet environments and along Biodiversidad (INBio). ecotones. Of the 1401 valid pipunculid species, 270 occur in the Neotropical Region (De Meyer, 1996; De Meyer & Skeving­ Allomethus Hardy. This genus contains four described ton, 2000). The Central American Pipunculidae are poorly species. A species formerly known only from Brazil PIPUNCULlDAE (BIG-HEADED FLIES) 54 801

(A. catharinensis Rafael) and three undescribed species have the relevant species can be formally transferred into these recently been collected in Costa Rica (based on material from genera. Of nine species of Eudorylas recorded from Costa INBio). Rica, six were described by Rafael & Menezes (1999) (one as Metadorylas). Mexican and Central American species Basileunculus Rafael. All three recognized species in this were revised by Rafael (1993) but not keyed. The Neotropi­ Neotropical genus (Rafael, 1987a) have been recorded in cal species of Eudorylas (=Metadorylas sensu Rafael) were Central America and the West Indies. revised and keyed by Rafael (1 990a).

Cephalops Fallen. Eight species have been identified (six Jassidophaga Aczel. A male and a female of this ge­ described) plus at least two undescribed species from Costa nus from the Durango region of Mexico are in the CNC. Rica that are known only from females. Neotropical species One specimen has associated elevational data (collected at can be keyed using Rafael (1990c). 2743 m). Based on these Mexican specimens, this genus Cephalosphaera Enderlein. Nine described species could reasonably be expected to occur in the mountains else­ occur in Central America, Mexico, and the West Indies. where in Central America. Keys to the Neotropical (Rafael, 1992), the West Indian Microcephalops De Meyer. Microcephalops latifrons (Rafael, 1996), and the Costa Rican species (Rafael & (Hardy) from the West Indies and M. williamsi (Hardy), Menezes, 1999) are available. which is widespread across the Neotropical Region, are the Chalarus Walker. Five species have been identified (four only described species. Two unidentified species (based on are described) plus at least two more from Costa Rica that females) have also been recorded in Costa Rica. The Neotro­ are known only from females. Neotropical species can be pical Region species were revised by Rafael (1990c) under keyed using Rafael (l990d). Cephalops, group latifrons.

Dorylomorpha Aczel. This widespread genus has not pre­ Nephrocerus Zetterstedt. Two female specimens in viously been recorded in Central America. However, a male the CNC were collected in San Cristobal de Las Casas of a new species of Dorylomorpha (Dorylomyia) is in the (2160 m, Chiapas, Mexico), and an additional female was USNM, and a female of a new species (possibly the same) collected near El Salto (2499 m, Durango, Mexico). As with is in INBio. Albrecht (1990) provides keys to the world Jassidophaga, this genus should be searched for elsewhere species. in the mountains of Central America.

Elmohardyia Rafael. All seven Central American species Pipunculus Latreille (= Parapipunculus Rafael). Ten spe­ occur in Costa Rica and can be identified usi ng the key in cies have been identified from Central America and Mexico Rafael & Menezes (1999). Eleven undescribed species were (nine described). Skevington & Marshall (1998) provided a discovered in recent collections made by INBio. key to all New World species.

Eudorylas Aczel (= Metadorylas Rafael as synonymized Tomosvaryella Aczel. This is the second largest genus recently by Skevington & Yeates, 2001). This is the larg­ in thi s region with 15 recorded species. The species from est genus in the family and certainly the largest in Central Mexico, Central America, and the West Indies were revised America. Many undescribed Costa Rican species exist in and keyed by Ale-Rocha & Rafael (1995). The West Indian collections. Recent work on the phylogeny of the Eudorylini species were keyed by Rafael (1996). has created a firm footing for the classification of the tribe in all but the Neotropical Region (Skevington & Yeates, 2001). Verrallia Mik. One Central American specimen is known There appears to have been a rapid radiation of members of from Chichicastenango, Guatemala. Previously, Verrallia this tribe within the Neotropical Region, leaving an appar­ was not recorded south of 37°N in the New World. We are ently heterogeneous mix of species that are currently all con­ now aware of specimens from the southwestern USA (New fined to the genus Eudorylas. Efforts to place this fauna into Mexico), Mexico, and Guatemala. Only two Nearctic species a world context will continue. It is apparent that the genera are known, but the genus is in need of revision in the New Claraeola Aczel and Dasydorylas Skevington occur in the World. The Guatemalan specimen likely belongs to an un­ New World tropics, but considerable work remains before described species. 802 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME 2

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Albrecht, A. 1990. Revision, phylogeny and classification of the genus Rafael, J.A. 1987a. Basileunculus, urn genero novo de Pipunculi­ Dorylomorpha (Diptera, Pipunculidae). Acta Zoologica Fennica dae (Diptera) da regiao Neotropical, corn chave para as espe­ 188: 1-240. cies. Acta Amazonica 16- 17 : 627-634. Ale-Rocha, R. & lA. Rafael. 1995. Tomosvaryella Aczel do Mexico, Rafael, l .A. 1987b. Two new genera of Pipunculidae (Diptera) America Central e Antilhas (Diptera, Pipunculidae). Revista Brasi­ from the New World: Metadorylas, gen. n. & Elmohardy­ leira de Zoologia 12: 407--427. ia gen. n., with new synonyms, designation of lectotypes and Benton, F.P. 1975. Latval taxonomy and bionomics of some British revalidation of a species. Revista Brasileira de Entomologia Pipunculidae. Ph.D. thesis, University of London, 208 pp. 31: 35- 39. Coe, R.L. 1966. Diptera Pipunculidae. Handbooksfor the Identification Rafael, 1.A. 1990a. 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Vega, F.E., P. Barbosa, & A.P. Panduro. 1991. EudOlylas (Meta­ of the corn leafhopper (Hemiptera: Cicadellidae) dorylas) sp. (Diptera: Pipunculidae): a previously unreported in Argentina, with a table of pipunculid- host associations parasitoid of Dalbulus maidis (Delong and Wolcott) and Dal­ in the Neotropical Region. Neotropical Entomology 38: bulus elimatus (Ball ) (Homoptera: Cicadellidae). The Canadian 152- 154. Entomologist 123 : 241-242. Waloff, N. & M.A. lervis. 1987. Communities of Virla, E.G. & lA. Rafael. 1996. Datos bion6micos preliminares y associated with leafhoppers and planthoppers in Europe, descripcion de la hembra de Cephalops penepauculus (Hardy) pp. 28 1- 402. In Macfayden, A. & E.D. Ford (editors). Ad­ (Diptera: Pipunculidae) um parasitoide de Delphacidae (Homop­ vances in Ecological Research. vol. 17. Academic Press, tera : Auchenorryncha) em Argentina. Revista de Investigacion London, 382 pp. Centro de Investigaciones para la Regulacion de Poblaciones Williams, F.X. 1919. Some observations of Pipunculus flies de Organismos Nocivos, Tucuman 10: 33-36. which parasitize the cane leathopper, at Pahala, Hawaii, Virla, E.G., G. Moya-Raygoza, & l .A. Rafael. 2009. First Record Feb. II- April 25, 1918. Proceedings of the Hawaiian Ento­ of Eudorylas schreiteri (Shannon) (Diptera: Pipunculidae) as a mological Society 4: 68-71.