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Novtates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novtates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2880, pp. 1-23, figs. 1-56 June 10, 1987 Amber Fossil Drosophilidae (Diptera), with Particular Reference to the Hispaniolan Taxa DAVID A. GRIMALDI' ABSTRACT All the known fossils ofDrosophilidae occurring Miomyia io, new genus; Protochymomyza mio- in amber are treated. Neotanygastrella wheeleri, cena, new genus; Scaptomyza dominicana; and new species, is named, redescribed (cf. Wheeler, two Drosophilinae species (A and B) incertae se- 1963) with respect to certain features, and a ho- dis. Relative genealogical relationships for some lotype and paratype are designated for the two drosophiline genera and subgenera are briefly dis- specimens from Chiapas, Mexico. Seven new cussed. Illustrations and photographs of the spec- species are described and two others are recorded imens are included, as well as ideas on the signif- from amber of the Dominican Republic (approx- icance ofsome ofthe fossils for Caribbean historical imately early Miocene in origin): Chymomyza pri- biogeography. A key to amber-fossilized Dro- maeva; Drosophila (D.?) poinari; D. (Hirtodro- sophilidae is provided. sophila) paleothoracis; D. succini (incertae sedis); INTRODUCTION Seven or eight fossilized specimens ofDro- will not be considered here. Two ofthe fossils sophilidae were previously known and all of are of a Neotanygastrella species from Chia- them occur in amber. Four (two males and pas, Mexico (late Oligocene to early Miocene two females) are Electrophortica succini Hen- in origin), which were described by Wheeler nig from the Baltic amber (Eocene to early (1963). Oligocene in age). Hennig (1965) gave very Cockerell (1923) described Drosophila ber- complete and detailed descriptions and draw- ryi from a specimen in amber found in Valle ings of E. succini, and Grimaldi (1987) dis- de Jesus, Santander, Colombia. He noted (p. cussed the placement of the species. This fly 331) that the "material, unfortunately of un- ' Assistant Curator, Department of Entomology, American Museum of Natural History. Copyright © American Museum of Natural History 1987 ISSN 0003-0082 / Price $2.85 _F t~~~~~~~i 2 AMERICAN MUSEUM NOVITATES NO. 2880 certain age, is relatively soft, being easily cut are certainly the youngest in origin (Lambert with an ordinary knife." Langenheim (1969) et al., 1985). In general, though, an early Mio- mentioned two Colombian sites among the cene origin of most Dominican amber is a amber deposits known in the world, at Me- reasonable estimate (Brouwer and Brouwer, dellin and Giron (Antioquia Provincia), of 1982). This dating is based on the fact that unknown Tertiary age, and that Hymenea the amber deposits are intercalated between (Leguminosae) is the probable source. The marine microfossil deposits that were laid location of Cockerell's specimen is unknown down approximately 23 mya, so the date is to me. Softness of the material strongly sug- one of redeposition and is, therefore, a min- gests the fossilized resin to be very recent, imal age. and perhaps it is even copal (1000 or more Langenheim and Beckh (1968) first ana- years old). If found, the matrix of the speci- lyzed the resinous components of various men should be examined for its C'4 decay ambers and found that material from the Do- (Burleigh and Whalley, 1983). The original minican Republic closely matched Hymenea description is actually of sufficient detail to (Leguminosae), which is a genus of trees confirm the specimen's identity as a drosoph- widespread throughout Central and South ilid, but an identification beyond this is im- America and the Greater Antilles. Lambert possible at present. Loew (1850) mentioned, et al. (1985) found, however, three major but did not describe, a "Drosophila" from classes of resinous components among sam- Baltic amber. This was the fly later described ples from nine Dominican amber localities. by Willi Hennig, Electrophortica. Bachofen- They concluded that ifHymenea is the source Echt (1949) listed three known amber fossil of the amber then the fossil resin is more drosophilids, no doubt referring to Loew's variable than samples from extant Hymenea. specimen, to the other specimens later de- Also, relative dating of the amber was done scribed by Hennig, and to Cockerell's spec- based on the NMR intensity ofthe exomethy- imen. lene carbon in the sample. Other than the This study is an attempt to add some dates provided by Brouwer and Brouwer chronological data to emerging hypotheses (1982), the actual dates of origin for various on the phylogenetic relationships and histor- Dominican ambers are rather hypothetical. ical biogeography ofthe Drosophilidae. Gri- maldi (1987) hypothesized that some of the Antillean drosophilid lineages are relicts; that METHODS, MATERIALS, AND is, their endemicity is imposed by extinction. ACKNOWLEDGMENTS Specimens from the rich amber deposits of All amber pieces from the Dominican Re- the Dominican Republic (Schlee and Glock- public had been previously tumble-polished. ner, 1978; Baroni-Urbani and Saunders, For closer observation ofsome details, amber 1982) and elsewhere provide information on was ground down close to the specimen using minimal age and perhaps on extinctions (e.g., a lapidary wheel and various-size grinding Krishna and Emerson, 1983; Wilson, 1985) grits (Buhler 240-1000) with water; amber for some groups ofthe Antillean pomace flies. was polished with a white alumina powder The former aspect would be particularly use- (#2 Buhler-micropolish 2A) plus water. The ful for comparison to the postulated dates of fossils were examined using incident and re- geological origin and position of Caribbean flected light from incandescent and fiber op- land masses. tics lamps, generally at 25-50 x magnifica- Amber of the Dominican Republic origi- tion using a Zeiss SV-8 stereoscope with nates from at least 10 major mining sites lo- attached camera lucida. Specimens were po- cated throughout the eastern three-quarters sitioned by placing them on cotton that was ofthe country at various altitudes. As a result, molded to accommodate the shape of each geological processes such as sedimentation amber piece, generally in an oil medium with have had different effects upon each site, and a refractive index equal to that ofthe amber. the calculated ages of the amber deposits can Photography employed the use of an Olym- vary among sites. Virtually transparent pieces pus OM-2N camera with auto-bellows and a of amber, which come from mines at Cotui, Zuiko macrolens (f6, about 12 x magnifica- 1987 GRIMALDI: AMBER DROSOPHILIDS 3 tion) and a Zeiss Tessovar (1 sec exposure), thoracic leg with femur, tibia, and proximal and Kodak Pan-X film. The standard mea- 3/4 of basitarsus darkened; remaining seg- surements made, where possible, were total ments of leg light. Abdomen dark brown to length ofbody (TL), thorax length (ThL), wing black. Oviscape without peg ovisensilla (male length (WL), head width (HW), 4-vein index unknown). (4-V), and costal index ofwing (CI). Protocol DESCRPTrION: Head light yellow. Arista with for these measurements is given in Grimaldi 3 dorsal and 2 ventral branches and 5-6 short (1986). Measurements were made at 50x medial ones. Pedicel with 2 long setae and magnification with an ocular micrometer. several shorter ones. Ocellar setae extended Dr. David Lindberg ofthe Museum ofPa- about to facial margin. Face flat; without ca- leontology, University of California, Berke- mna. One pair ofvibrissae present; ends near- ley, kindly loaned the two hypotypes ofNeo- ly touch; subvibrissae tiny. Clypeus and palps tanygastrella described by Marshall Wheeler. retracted within oral cavity. Eyes light red, Mr. Jacob Brodzinsky allowed me to sort bare of interfacetal setulae; collapsed in ho- through his collection forthe purchase ofsome lotype. Inner vertical setae c. equal in length specimens. Dr. George 0. Poinar, Depart- to outer verticals and anteromedial to them. ment of Entomology and Parasitology, Uni- Proclinate orbital about midway between an- versity of California, Berkeley, generously terior and posterior reclinates and c. equal in loaned several specimens from his personal length or slightly shorter. collection. Dr. Wayne N. Mathis, Depart- Acrostichal setulae in 4 even rows. Dor- ment of Entomology, National Museum of socentrals thin and straight, parallel. Two hu- Natural History, sorted through the collec- meral setae present. Anterior scutellar setae tion at the Smithsonian, from which five am- slightly divergent; apical scutellars conver- ber pieces were borrowed. Drs. Gerhard gent. Postnotum prominent. Halter lighter Bachli, Ian Bock, Walter Hackman, Wayne than evenly colored, yellow thorax. Wing Mathis, Toyohi Okada, George Poinar, Nor- completely hyaline (wings are folded in ho- man Platnick, Randall Schuh, and Marshall lotype in way to prevent accurate measure- Wheeler provided comments on the manu- ments). Meso- and metathoracic legs mostly script, which Ms. Carol Ievolella kindly typed. yellow. Prothoracic leg with yellow coxa and Ms. Susan Klofak helped with the amber trochanter; tarsal segments 2-5 lighter, nearly preparations. white. Forefemur, tibia, and proximal 3/4 of Abbreviations for collections in which basitarsus black-brown. Forecoxa elongate, specimens are deposited are the following: length c. 2/3 that of femur. Abdomen slender, AMNH, American Museum of Natural His- laterally compressed in holotype. Tergites tory (New York); GOP, George 0.
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