From Mexican Amber: a New World Species with Old World Affinities
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PROC. ENTOMOL. SOC. WASH. 110(1), 2008, pp. 116–125 TOXORHYNCHITES (TOXORHYNCHITES) MEXICANUS, N. SP. (DIPTERA: CULICIDAE) FROM MEXICAN AMBER: A NEW WORLD SPECIES WITH OLD WORLD AFFINITIES THOMAS J. ZAVORTINK AND GEORGE O. POINAR,JR. (TJZ) Bohart Museum, Department of Entomology, University of California, One Shields Avenue, Davis, CA 95616, U.S.A. (e-mail: [email protected]); (GOP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e- mail: [email protected]) Abstract.—A new species of fossil mosquito, Toxorhynchites (Toxorhynchites) mexicanus, is described from Mexican amber. This species differs from all extant species of the genus by having a strong bristle posteriorly on the postpronotum and it differs from all species of the subgenus Toxorhynchites by having the unique combination of a short rm index (1.3) and conspicuous caudal tufts of setae on the abdomen. This is the first record of the Old World subgenus Toxorhynchites in the New World. Key Words: fossil, new species, Mexico, Neotropics, disjunct distribution Adult mosquitoes of the genus Tox- the genus can be found in papers by orhynchites Theobald are noted for their Steffan and Evenhuis (1981, 1985) and large size, brilliant coloration, and re- in many of the classic mosquito faunas flexed proboscis. The more than 90 and catalogs of the past century (Bar- extant species and subspecies are placed raud 1934; Belkin 1962; Carpenter and into four subgenera. The genus is pri- LaCasse 1955; Dyar 1928; Edwards marily tropical in distribution, but with 1922, 1932, 1941; Lane 1953; Lee a few species extending into north et al. 1988; Service 1990; Tanaka et al. temperate regions (Japan, southern Si- 1979). beria, southern Canada). Females are Relatively few fossil mosquitoes have non-haematophagous and oviposit in been described from amber. Among phytotelmata. Larvae are predaceous, these are Paleoculicis minutus Poinar, feeding chiefly on other mosquito larvae. Zavortink, Pike, and Johnston, 2000 For a variety of reasons—the lack of from Cretaceous Canadian amber; medical importance, the difficulty of Anopheles (Nyssorhynchus) dominicanus associating the sexes because of strong Zavortink and Poinar, 2000, and Culex sexual dimorphism, the relative unifor- malariager Poinar, 2005 from Tertiary mity of the immature stages and of the Dominican amber; and the alleged mos- male genitalia throughout the genus, and quito Burmaculex antiquus Borkent and the difficulty of obtaining sufficient Grimaldi, 2004 from Cretaceous Bur- numbers of specimens to assess varia- mese amber. A synopsis of all described tion—the genus is poorly known taxo- fossil mosquitoes known at the time is nomically. Further information about given by Poinar et al. (2000). VOLUME 110, NUMBER 1 117 The mosquito Culex loewi Giebel was Sandstone (Lower Miocene) to the La originally described as a fossil species Quinta Formation (Upper Oligocene). supposedly embedded in amber (Giebel They are assigned to the planktonic 1862). Subsequent research has shown, foraminiferal zones represented by Glo- however, that the specimen is in recent bigerina ciperoensis Bolli and Globorota- gum copal and that it belongs to the lia kuglieri Bolli, Loeblich and Tappan, extant species Toxorhynchites brevipalpis an interval within zones N3 and N4 of Theobald. The history of this specimen is the Cenozoic Planktonic Foraminiferal provided in the request to the Interna- Zonal Sequence, which has been dated tional Commission of Zoological No- radiometrically at 22.5 to 26 million menclature to suppress the name T. loewi years (Berggren and Van Couvering and conserve T. brevipalpis (White 1977). 1974). However, since the amber was Among fossils from the Isle of Wight washed into a coastal lagoon from soils provisionally referred to Culex protorhi- beneath resin-producing trees, these nus Cockerell by Edwards (1923) is dates provide only a minimum age for a male abdomen described as follows: it. Further information on Mexican ‘‘The eighth segment is very broad, much amber may be found in Poinar (1992). broader than the hypopygium, and distinctly broader than the seventh seg- MATERIALS AND METHODS ment (thus recalling Megarhinus [5 The piece of amber with the Toxo- Toxorhynchites]).’’ This statement by rhynchites specimen is roughly trapezoi- Edwards is the basis for the overstate- dal in outline, with sides of 40, 15, 20, ment by Mattingly (1962): ‘‘An Oligo- and 25 mm, a thickness of 5 to 10 mm, cene fossil from the Isle of Wight has and a weight of 5.59 g. It is clear orange also been attributed to it [Toxorhynch- yellow in color. In addition to the ites] (Edwards 1923).’’ The true identity Toxorhynchites mosquito, the amber of this fossil has not been determined includes six smaller dipterans, includ- and it has not been formally described. ing two small, damaged culicine mosqui- We take this opportunity to describe toes, and bits of dark brown or black a new species of Toxorhynchites from organic debris. The Toxorhynchites mos- Mexican amber. This is the first genuine quito is nearly complete, with only the fossil of the genus to be described. We left hind leg missing. The position of the place the new species in the subgenus mosquito in the amber precludes a clear Toxorhynchites, thereby establishing an view of portions of the head and thorax, early record of the subgenus in the New and the presence of a thin silvery film World. The tree responsible for pro- over much of the surface of the thorax ducing Mexican amber, the legume obscures the distribution of scales and Hymenaea mexicana Poinar and Brown, other characters on the scutum and 2002, also has Old World affinities, with pleuron. the most closely related living species Adults of Toxorhynchites and several occurring in Africa. other genera of diurnal mosquitoes have The amber with the fossil Toxorhynch- scales with iridescent or metallic colors ites originated from mines or exposed that may vary in intensity and hue amber-bearing veins near the village of according to the angle of observation. Simojovel, Chiapas, Mexico. The amber If scales of the fossil Toxorhynchites, as, there occurs in a sequence of primarily for example, those at the edges of marine calcareous sandstones, siltstones, structures, can be observed by trans- and shales with seams of lignite. These mitted light, then it can be determined deposits extend from the Balumtun whether they are pigmented or not. The 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pigmented ones were probably brown or Edwards 1922, 1941; Steffan and Even- black and probably displayed two or huis 1985). more of the iridescent colors adjacent to Description of holotype female.— each other in the series varying from blue Wing: 5.6 mm. Proboscis: 5.5 mm. Fore- to blue green, green, bronze, deep femur: 3.3 mm. Abdomen: 4.9 mm. Head golden, copper, and purple. The scales (Fig. 2): Integument dark brown. Eyes without pigment may have been matte contiguous above antennae and ventrally white or may have displayed brilliant behind proboscis. Top of head with metallic silver or gold reflections. We broad flat scales, some in front with have described some scales of the fossil dark green iridescence, some in back as dark and some as light, meaning only with purple iridescence; occiput with that they do or do not bear pigment. We erect scales posteriorly. Ocular bristles have described some scales as having present, at least 5 upper, 1 strong lateral green or purple iridescence, but we do near widest part of head on left side; not know what colors they may have underside of head with fine bristles. displayed in life. First, we do not know Clypeus overlying base of palpus later- how fossilization in amber may have ally. Proboscis long, attenuate, curved affected the structural colors of the downward beyond middle; with numer- scales, and second, we do not know the ous subbasal bristles; inserted into cavity complete range of hues the scales may of head capsule, without collarlike ring have displayed since most parts of the at base; labellum long. Palpus short, fossil can be observed in only one about 0.2 length of proboscis, slender, orientation. with 2 obvious palpomeres, the distal 2.5 Observations and photographs were length of proximal. Antenna short, made with a Nikon stereoscopic micro- about 0.6 length of proboscis; pedicel scope SMZ 1500 and Nikon Optiphot small, apparently with scales dorsally; microscope at magnifications up to flagellar bristles sparse; flagellomere 1 1123. Morphological terminology fol- slightly longer than 2, with scales dor- lows McAlpine (1981), Harbach and sally. Thorax: Integument black. Para- Knight (1980), and Steffan and Evenhuis tergite broad. Antepronotal lobes large. (1985). Meron small, its upper margin in line with base of hindcoxa. Scutum with Toxorhynchites (Toxorhynchites) mexi- bristles on anterior promontory, antealar canus Zavortink and Poinar, new species area, and supraalar area; with broad flat scales, some in posterior fossal area dark (Figs. 1–4) with purple iridescence, some in posteri- Diagnosis.—This species differs from or dorsocentral area dark with green all extant species of Toxorhynchites by iridescence. Scutellum evenly rounded, possessing a strong bristle posteriorly on with even row of marginal bristles or the postpronotum. Its short palpus alveoli; with broad flat scales, most comprised of two obvious palpomeres apparently with purple iridescence. Ante- allies it with Old World species of the pronotum with anterior row of numer- genus, from which it can be distinguished ous large and small bristles; with broad by the unique combination of a short rm flat scales, some with purple iridescence. index (1.3; equals length of the basal spur Postpronotum with 1 strong bristle of wing vein R4+5 from the base of vein posteriorly; with broad flat scales, those R4+5 to crossvein r-m divided by the in upper posterior part dark with purple length of crossvein r-m) and conspicuous iridescence.