Diptera: Culicidae) from Dominican Amber
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SZAQYSTEMATICS Anopheles (Nyssorhynchus) dominicanus sp. n. (Diptera: Culicidae) from Dominican Amber 1 THOMAS J. ZAVORTINK AND GEORGE O. POINAR, JR. Department of Biology, University of San Francisco, San Francisco, CA 94117Ð1080 Ann. Entomol. Soc. Am. 93(6): 1230Ð1235 (2000) ABSTRACT The female and egg of Anopheles dominicanus sp. n. are described from Dominican amber. The species is placed tentatively into the subgenus Nyssorhynchus on the basis of the following wing characters: a preapical pale costal spot; an accessory sector pale spot that is not joined to a sector pale spot; pale scales at the apex of vein R4ϩ5; basal, median, and apical pale spots on vein A; and the termination of vein Sc at the proximal end of the subcostal pale spot. This newly described mosquito from Dominican amber establishes an ancient lineage of Anopheles in the New World. KEY WORDS Anopheles dominicanus, new species, Culicidae, Diptera, mosquitoes, Dominican amber AS A RESULT of their small size and fragile constitution, Materials and Methods fossil mosquitoes are rare (see Poinar et al. [2000] for The amber with the new anopheline is clear yellow, a list and critical examination of known fossil Culici- with numerous small cloudy droplets, and with many dae). The current study describes the female and egg minute irregular or ovoid particles, some of which are of a new species of spotted-wing Anopheles in amber joined into chains. It includes the adult female Anoph- from the Dominican Republic. This is the Þrst de- eles; two adjacent eggs laid by it; three long, slender, scription of a mosquito from Dominican amber and nonculicid insect legs; a plant bud; and a small, dark, the Þrst fossil anopheline from the New World. hemispheric object covered with fungal hyphae. The The fossil described here is unquestionably an piece of amber is irregularly oval in outline, Ϸ9.5 by Anopheles. Its subgeneric placement is less certain, 11.0 mm, with a portion of its perimeter polished ßat. however, in part because the subgeneric classiÞcation The piece has been polished to a thickness varying of Anopheles is based to a great extent on male char- from Ϸ1.5 to 2.0 mm, and its weight is 0.18 g. The acters, and in part because many taxonomic characters amber has been examined with both stereoscopic of femalesÑdevelopment of the buccopharyngeal ar- (with magniÞcations up to 140 diameters) and com- mature, presence or absence of pruinose markings on pound (with magniÞcations up to 250 diameters) mi- the thorax, presence or absence of scales or scale-tufts croscopes using both transmitted and reßected light. on various parts of the body, and ornamentation of the Measurements were made at various magniÞcations legsÑare missing, not discernable, or not known with with an ocular reticule and the original line drawings certainty. We have had to rely heavily on character- were prepared with the aid of the reticule. Photo- istics of the wing and egg in our attempts to relate the graphs were taken with a Nikon Orthophot micro- fossil Anopheles to modern species. The fossil appears scope. The drawing of the wing (Fig. 2) is diagram- to be most similar to species in the subgenus Nysso- matic and intended only to show the positions of the rhynchus, so we place it there tentatively. pale spots. Morphological terminology follows Mc- The amber containing the mosquito originated from Alpine et al. (1981) in large part, Wilkerson and Pey- the northern mountain ranges of the Dominican Re- ton (1990) for wing spots, and Hinton (1968) and public. Amber mines there are in the El Mamey For- Linley (1992) for egg characters. mation (Upper Eocene), which is a shale-sandstone interspersed with a conglomerate of well-rounded pebbles (Eberle et al. 1980). The exact age of Domin- Results ican amber is unknown, and estimates based on var- ious analyses of microfossils and the amber provide a The fossil Anopheles is damaged, and many charac- range from 15Ð20 million years (Iturralde-Vincent and ters used in anopheline taxonomy are not present. The MacPhee 1996) to 30Ð45 million years (Cepek in proboscis and palpi extend to the edge of the amber, Schlee 1990). and the distal parts are missing. Only the right foreleg is complete, and both midlegs and hindlegs are broken off and missing. Various parts of the mosquitoÑthe 1 Department of Entomology, Oregon State University, Corvallis, head, antennal pedicels, antepronota, proepisterna, all OR 97331Ð2907. coxae, halters, and cerciÑare ßattened, apparently 0013-8746/00/1230Ð1235$02.00/0 ᭧ 2000 Entomological Society of America November 2000 ZAVORTINK AND POINAR:NEW Anopheles FROM DOMINICAN AMBER 1231 having collapsed during fossilization. The thorax is without scales; ßagellomere 5 about six times longer cracked and distorted. The abdomen is ßattened, the than broad; median ßagellomeres with eight bristles in cuticle of the tergites and sternites is wrinkled, and the basal whorl and 1Ð3 short setae distally; each of ßag- pleural membrane is extruded on the left side of seg- ellomeres 10Ð13 slightly longer than preceding one; ments IIÐV. The head, thorax, and abdomen are largely apical ßagellomere about 11 times longer than broad, denuded. The presence or absence of scales on the pointed. Thorax. Scutal integument dark brown; thorax and abdomen has been particularly difÞcult to darker areas not evident. Long rows of large setal determine, and any pruinose pattern that may have alveoli in acrostichal, dorsocentral, and supraalar ar- been present on the scutum or pleuron has been oblit- eas; additional large setal alveoli along lateral and erated during fossilization. One of the eggs is cracked posterior edges of scutal fossa. Left humeral area with and shriveled; both eggs are unsclerotized and visible three long, narrow, dark erect scales; no other scales only in lateral aspect. evident. Scutellum rounded; six large setal alveoli vis- ible along right half of posterior border, the lateral alveoli closer together; without smaller setae or alveoli Anopheles dominicanus Zavortink & Poinar, sp. n. dorsad of row of large alveoli; scales not evident. Pleu- (Figs. 1–5) ral integument dark brown. Left antepronotum with Diagnosis. A spotted-wing Anopheles, the wing pat- several setal alveoli and six long, narrow, dark erect tern most similar to that of subgenus Nyssorhynchus, scales above, at least Þve setae or alveoli below; pro- the following wing characters diagnostic alone or in episternum with one long seta; spiracular setae two; combination: accessory sector pale spot absent on prealar knob with several setal alveoli; upper katepis- veins C and Sc, present on vein R; veins Rs, R2ϩ3, and ternum with two setal alveoli; lower katepisternum R3 primarily dark scaled; vein R2 dark at apex; vein with one long and one shorter seta; upper R4ϩ5 with short median pale spot; vein M1 without mesepimeron with at least one long and one shorter pale scales beyond fork; vein CuA dark at base and its seta. Legs. Scales not evident on coxae. Integument of branches primarily dark scaled; basal pale fringe spot coxae and trochanters dark brown. Femur of foreleg absent. Flagellomeres uniformly brown, relatively not swollen basally. Foreleg partly denuded, appar- long and slender, ßagellomere 5 about six times longer ently dark scaled, apex of tibia possibly with narrow than broad, ßagellomere 13 about 11 times longer than pale ring. Tarsal claws of foreleg small, subequal, sim- broad. Antepronotum with long, narrow, erect scales. ple. Wing (Fig. 2). Both wings present; basal portion Pleuron with one strong proepisternal seta, two spi- of costa and radius of both partly to completely de- racular setae, and two lower katepisternal setae. Fe- nuded. Fringe missing from behind M1 to before A on mur of foreleg not swollen basally. Egg apparently right wing and from behind M2 to before A on left without deÞnite polygonal or blister-like outer chori- wing. Scales on costa in subbasal dark spot with six onic cells; ßoats dorsolateral, broad, Ϸ0.5 length of striations, those on vein R in subbasal dark spot with egg; frill deep, extending to ends of egg; lobed tuber- four striations. Costa with basal, humeral, sector, sub- cles apparently present. costal, and preapical pale spots; humeral pale spot 0.7 Female(Figs. 1Ð3). Length of scutum and scutel- length of subbasal dark spot; sector pale spot 0.2 length lum: 0.86 mm. Forefemur: 1.58 mm. Wing: 2.80 mm. of sector dark spot; subcostal pale spot 0.33 preapical Abdomen: 2.2 mm. Head. Flattened dorsoventrally, dark spot. Stem vein denuded, but probably bore dark the left compound eye extruded and ruptured; largely scales as vein is darkly pigmented. Vein R-R1 denuded denuded. Integument dark brown. Interocular space basally, with sector, accessory sector, subcostal, and apparently concave, its width equal to width of 3Ð4 preapical pale spots. Vein Rs dark scaled at base; with facets. Frontal tuft missing. Dorsal surface of head distinct basal spur with one scale. Vein Rs-R2ϩ3 with with one long, dark erect scale left of center on vertex, small pale spots at base of R4ϩ5, over fork into R2 and one long, slender, dark erect scale caudolaterally on R3, before apex on R2, and at apex of R3. Vein R4ϩ5 left side, and two long setae and one long, slender, dark without basal spur; with scales of dorsal surface erect scale caudolaterally on right side. Clypeus prom- spreading; small pale spots at base and apex and longer inent, bare. Proboscis and palpi extending to edge of pale spot basad of level of forks in R2ϩ3 and M. Stem amber, distal portions missing. Proboscis dark scaled, of vein M relatively densely scaled. Vein M with small scales decumbent; only one basal bristle evident. One pale spot at crossveins, over fork, and at apex of M2.