AMERICAN MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3127, 8 pp., 13 figures, 1 table April 6, 1995

A Remarkable New Species of Ogcodes (Diptera: Acroceridae) in Dominican Amber

DAVID GRIMALDI'

ABSTRACT Four specimens of a distinctive new species of and tergal mounds (structure of the antennae and acrocerine fly are described as Ogcodes exotica, wing venation unquestionably places it in the cos- preserved in Oligo-Miocene amber ofthe Domin- mopolitan genus Ogcodes). However, features of ican Republic. The fossil species appears most its cervical region are found in the African genera closely related to several Old World species of Meruia and Sabroskya. Ogcodes, based on distinctive, clavate hind tibiae

INTRODUCTION The family of obligate internal parasitoids grasshoppers and beetles). The bee flies (fam- of spiders, the Acroceridae, is one of con- ily Bombyliidae) have the same life history. trasts. Although the family has only 500 Nemestrinids were described by Rohden- known species and 50 genera, its morphology dorf (1968), in Jurassic sediments of Russia, is exceptionally diverse. It is undoubtedly a so this would imply that the acrocerids are monophyletic lineage, based not only on the equally ancient, based on sister-group dating. distinctive life history, but also on a suite of In fact, an acrocerid has been described from morphological characters: adult flagello- the same Jurassic beds of Karatau (Ussat- meres fused into a single flagellum; lower ca- chov, 1968), but the preservation of Archo- lypter extremely large; and a wing membrane cyrtus gibbosus is not complete, especially for with very fine pleating. Woodley (1989) the head. The venation ofArchocyrtus is most grouped the acrocerids with the nemestrinids similar to that of certain acrocerines, es- on the basis of hypermetamorphosis. Ne- pecially Acrocera, but there are some features mestrinid larvae are also parasitoids (but on unlike any found in living members of the ' Associate Curator and Chairman, Department of Entomology, American Museum of Natural History.

Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $1.20 2 AMERICAN MUSEUM NOVITATES NO. 3127 family. Huge gaps exist in the fossil record: that established in the Manual of Nearctic the next oldest fossils are in the Baltic amber Diptera; wing vein nomenclature follows that from the upper Eocene-lower Oligocene, ca. of Hennig (1968). Length of the wing was 40 million years old (Hennig, 1966, 1968). taken from the very base of the wing to the In a family of such apparent antiquity, a tip. fossil from the Oligo-Miocene (25-30 million years old) contributes little toward under- ACKNOWLEDGMENTS standing basal relationships. Rather, the im- portance of such a young fossil provides in- formation on the origins and stasis ofspecies- Thanks are extended to Dieter Schlee and level taxa, and biogeography. Ettore Morone, for loaning specimens of 0. Amber from the Dominican Republic lies exotica in their care; to Norman Woodley in marine sediments northeast of Santiago, (USDA/SEL) and Evert Schlinger for original in discussions on the placement of this bizarre extensively uplifted terrain. The only mine species; and to Manuel Perez and Jake Brod- to have been stratigraphically dated (with zinsky, for allowing me to sort through their benthic Foraminifera) is the Palo Alto mine, amber fossil caches. NSF grant BSR 9020102 dated as 23 million years old (Baroni-Urbani funded the field and laboratory work on Dip- and Saunders, 1980). Since the amber obvi- tera living in the Caribbean and those fos- ously does not lie in its original milieu, this silized in Dominican amber. David Barra- is a minimum age, although it can safely be clough kindly lent the types ofSabroskya. He, assumed that the age is hardly greater than Norman Woodley, and Evert Schlinger pro- this. vided helpful critiques of the manuscript. MATERIALS AND METHODS DESCRIPTION Most Dominican amber specimens avail- Ogcodes exotica, new species able for study to the scientific community are Figures 1-4, 10-13 purchased from dealers, who themselves ac- quire the specimens from the campesinos who DiAGNosIs: Thorax relatively small and not mine the material. With the exception of a extremely arched; distinct cervical region few dealers, virtually all of the amber pieces present; abdomen slender (not dorsoventral- become mixed, so that it is nearly impossible ly flattened), abdominal tergites 2-5 each with to trace the source of an amber piece to one a central mound, bearing tuft of fine, erect of dozens of mines. The exception is that the setae (e.g., figs. 2, 4); venation typical of light, soft "copal" from the eastern mines Ogcodes, with crossvein m-cu present; hind (Bayaguana, Cotui) is easily distinguished tibia clavate, with apical half enlarged to 3- from the darker, heavily polymerized mate- 4 x diameter ofproximal end. Proboscis ves- rial (true amber) from the hills surrounding tigial. Known from four males. Santiago. Chemical fingerprinting, using py- DESCRIPTION: Measurements: see table 1. rolysis-gas chromatography, has not revealed Head: Ocellar triangle slightly raised, me- any consistent differences among amber from dian ocellus lost. Dorsal margins ofeyes con- the different mines. Thus, the best and most tiguous to level of antennae; hind margin of conservative estimate is that any true Do- eye with slight indentation, dorsal and ven- minican amber is approximately 25 million tral facets undifferentiated in size. Proboscis years old. Certainly, published claims (e.g., vestigial. Antenna with long, thin flagellum Scarbrough and Poinar, 1992) that particular having slightly swollen apex; apical setulae fossils sold commercially originate from the unapparent; antennae situated ventrally, al- La Toca mines (based simply on color) and most near oral margin. are, therefore, Eocene in age, have virtually Thorax: Distinct cervical region present, no basis. Methods for preparation, micro- this area not hidden behind the head; posi- scopic examination, and photography of the tioned between enlarged postpronotal lobes. amber specimens are given in Grimaldi Postpronotal lobes widely separated. Meso- (1993). Morphological terminology follows notum not strongly arched; with black me- 1995 GRIMALDI: OGCODES IN DOMINICAN AMBER

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Fig. 4. Habitus illustration of holotype, AMNH DR- 11-8. dian stripe and several rows of fine, erect ac- crossvein m-cu present, well developed. Vein rostichal setae that increase in length poste- Cu meeting anal vein before wing margin, riad. Posterior rim of scutellum with similar forming a long triangular cell, cup. fine, erect setae. Legs slender. Hind tibia cla- Abdomen: Rather long and slender, with vate, the diameter of the apical half 3-4 x tergites 2-5 each raised with central mound that ofthe proximal half; with numerous fine, bearing tuft of long, fine, erect setae. Apical stiffsetulae on short, tubercular bases. Claws tergite also with tuft oflong, fine setae. Color and pulvilli large. Wing with fine pleating pattern seen on Stuttgart specimen, as shown typical of acrocerids; costal vein extended to in figure 11. Genitalia observed on two spec- tip of wing, slight swelling where vein R1 + imens, as figured (figs. 12, 13): Distiphallus meets costal vein. Vein R4+s with slight kink, arrowhead-shaped. not quite reaching posterior margin of wing. TXPEs: Holotype: AMNH DR- 1 1-8, a male Crossvein r-m slanted strongly anteriad; in- in a drop-shaped piece of clear yellow amber complete, not reaching vein M3. Vein M2 in- from Dominican Republic, 16 x 10 mm (figs. complete: not reaching wing margin, present 3, 4). Paratypes: AMNH DR-8-88 (fig. 1), in only as a fold at level ofcrossvein r-m. Apex a large chunk of yellow amber, with several of vein M3 not reaching margin of wing; unpolished surfaces; Staatliches Museum fur 1995 GRIMALDI: OGCODES IN DOMINICAN AMBER 5

M~~~M M2~~M 7

4+ M3 Cu 8 Figs. 5-8. Comparative structures on extant Acroceridae. 5-7: Sabroskya palpalis (holotype). 5, 6. Head and anterior portion of thorax (lateral view, 5; dorsal view, 6). ppl, postpronotal lobe. 7. Wing. 8. Wing, Ogcodes clavatus. Drawing based on a specimen in the Smithsonian Institution from Naivasha, Kenya. The dotted r-m crossvein was not observed in this specimen but it was reported as present in a specimen from Pretoria, South Africa (Schlinger, 1960a: fig. 11). A trace of r-m was reported (Schlinger 1960b: pl. 3, fig. 10), in probably the same specimen as this one from Naivasha. 6 AMERICAN MUSEUM NOVITATES NO. 3127

. m 0.5 mm

Figs. 9-13. Features of Ogcodes exotica (figs. 10-13) and 0. clavatus (fig. 9). 9, 10: Hind tibiae. 11. Dorsal view of Stuttgart paratype (DO-5374-H), showing color pattern and position of tergal mounds. 12, 13: Male genitalia. 12. Morone collection, no. 853. 13. Holotype, AMNH DR- 11-8.

Naturkunde, Stuttgart DO-5374-H (17 x 9 following characters: antennal flagellum with mm); Morone Collection no. 853(private), in a slight apical swelling (plesiomorphically a hemispherical piece 12 x 14 mm. there is no swelling); antenna located ven- ETYMOLOGY: From the Greek, exotikos, trally, near the oral margin (a feature also meaning from the outside, alien, foreign, and occurring in Sabroskya, Meruia, and some exotic; in reference to its apomorphic features other acrocerines); mouthparts (proboscis and and apparent Old World relationships. palps) absent or vestigial; eyes bare (dense pilosity is plesimorphic and widespread RELATIONSHIPS among Acrocerinae); venation highly re- The fossil species is unequivocally placed duced, such that crossvein r-m is incomplete into the genus Ogcodes, on the basis of the or entirely absent, and vein M2 is incomplete 1995 GRIMALDI: OGCODES IN DOMINICAN AMBER 7

(not reaching the wing margin or to level of TABLE 1 crossvein r-m) (e.g., fig. 8). Measurements (in mm) of Ogcodes exotica In fact, several apomorphic features of 0. Specimens exotica link it with a few Old World species. Total Thorax Wing The clavate hind tibia is found in 0. clavatus length length length Becker (from South and East Africa) and 0. respersus Seguy (from northern China). I was AMNH DR-11-8 5.08 1.55 3.73 AMNH DR-8-88 5.05 1.50 3.75 able to compare the hind tibia of the fossil M-853 5.33 1.68 4.33 only with clavatus (based on a specimen in 6.21 1.96 4.65 the Smithsonian Institution from Naivasha, DO-5374-H Kenya [H.J.A. Turner, coll., 4-40, C.I.E. 12838]). The hind tibia in clavatus is not as abruptly swollen as in exotica (cf. figs. 9, 10), tinctive character has arisen several times in and clavatus also has a cervical region more the family. Likewise, the widely separated extended than in other Ogcodes (but not to postpronotal lobes and protrudent pronotal the extent seen in exotica). Ogcodes clavatus, collar seen in the fossil are certainly conver- however, has the stout abdomen typical of gent with what is seen in Sabroskya (e.g., pal- virtually all Ogcodes, and it lacks tergal palis, figs. 5, 6) and Meruia. In virtually every mounds. Only three Ogcodes species, in fact, other respect, such as eye pile, well-devel- have mounds on the tergites: 0. hirtus Sack oped mouthparts (fig. 5), and the complete (from Iran) with "bituberculate swellings on wing venation (fig. 7), Sabroskya and Meruia tergites II-IV" [Schlinger, 1960b: 271]), 0. are plesimorphic with respect to the fossil and guttatus Costa (widespread from South Af- other Ogcodes. rica to Italy and India), and 0. orientalis The Acroceridae are notorious for rarely Schlinger (from Cambodia) (with tergites II being encountered. Thus, it is uncertain if and III swollen in the middle). From the cur- species of Ogcodes similar to the fossil will sory accounts of 0. hirtus and 0. orientalis, be found living in the Western Hemisphere. the tergal mounds are fewer than in 0. exo- At present, the only definitive relationships tica, and they are certainly not bituberculate of the fossil are with Oriental and African in the fossil. Tergal mounds are common in species. This is a biogeographic connection some philopotine acrocerids, such as Oligo- that is slowly emerging with the study of or- neura. This subfamily has a suite of features ganisms fossilized in Dominican amber. plesiomorphic to the Acrocerinae, so this dis-

REFERENCES Baroni-Urbani, C., and J. B. Saunders Rohdendorf, B. B. 1980. The fauna of the Dominican Republic 1968. New Mesozoic nemestrinids (Diptera, amber: the present status ofknowledge. Nemestrinidae), In B. B. Rohdendorf Proc. Ninth Caribbean Geological Conf. (ed.), Jurassic insects of Karatau, pp. (Santo Domingo, Aug. 1978), pp. 213- 180-189. Moscow: Academiya Nauk 223. SSSR. Grimaldi, D. Scarbrough, A. G., and G. 0. Poinar, Jr. 1993. The care and study of fossiliferous am- 1992. Upper Eocene robber flies of the genus ber. Curator 36: 31-49. Ommatius (Diptera: Asilidae) in Do- Hennig, W. minican amber. Insecta Mundi 6: 13- 1966. Spinnenparasiten der Familie Acrocer- 18. idae im Baltischen Bernstein. Stuttgart- Schlinger, E. er Beitr. Naturkd. 165: 1-21. 1960a. A review of the South African Acrocer- 1968. Ein weiterer Vertreter der Familie Ac- idae (Diptera). Ann. Natal Mus. 14: 459- roceridae im Baltischen Bernstein (Dip- 504. tera: Brachycera). Stuttgarter Beitr. Na- 1960b. A revision of the genus Ogcodes La- turkd. 185: 1-6. treille, with particular reference to spe- 8 AMERICAN MUSEUM NOVITATES NO. 3127

cies of the Western Hemisphere. Proc. Woodley, N. E. U.S. Natl. Mus. 3429: 227-336. 1989. Phylogeny and classification ofthe "Or- Ussatchov, D. A. thorrhaphous" Brachycera. In J. F. 1968. New Jurassic Asilomorpha (Diptera) McAlpine (ed.), Manual of Nearctic fauna from Karatau. Entomol. Rev. 47: Diptera, 3: 1371-1395. Research Branch 617-628. Agriculture Canada Monograph no. 32.

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