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 , Toxorhynchites (Toxorhynchites) mexicanus, is described from Mexican amber. This species differs from all extant species of the 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 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 ) 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. Pleuron with bristles on caudal tufts of setae on the abdomen (see upper proepisternum (at least 11), spi- VOLUME 110, NUMBER 1 119

Figs. 1–2. Holotype female of Toxorhynchites mexicanus in Mexican amber. 1, Whole mosquito. 2, Head. 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3–4. Holotype female of Toxorhynchites mexicanus in Mexican amber. 3, Wing. 4, Abdomen. VOLUME 110, NUMBER 1 121 racular area (at least 12), posterior scales not obvious, but scales in basal katepisternum (several, 1 large), prealar median triangular patch on sternum IV knob (at least 22, some long, 2.0–2.5 appear darker than others. times length of spiracular bristles), and Type material.—HOLOTYPE female mesepimeron (at least 13 in upper area in amber, from near Simojovel, Chiapas, and caudal row, apparently 2 strong in Mexico. Accession number D-7-188, de- lower area). Pleuron with broad flat posited in the Poinar amber collection scales on anteprocoxal membrane, upper maintained at Oregon State University, proepisternum, katepisternum, and me- Corvallis. sepimeron, possibly also on lower pro- Etymology.—The new species is episternum. Legs: Bristles short, stout. named for its country of origin. Foretarsomeres 4,5 subequal in length. Hindtibia apparently completely dark DISCUSSION scaled. All tarsi apparently completely Toxorhynchites mexicanus clearly be- dark scaled. Claws simple, smaller on longs to the genus Toxorhynchites,as hindleg. Pulvilli absent. Wing (Fig. 3): shown by the following characteristics: Long, slender. Small area of membrane large size; scales of head, thorax, and around crossvein m-cu slightly infus- abdomen broad, flat, and with evidence cated. Scales on anterior veins short, of iridescent colors; proboscis long, broad. Cell R2 short, about 0.36 length attenuate, reflexed, and inserted into of vein R2+3. Vein Rs right-angled at cavity of head capsule, the genae not base. Vein R4+5 right-angled at base and forming a collarlike ring at its base; with long, scaled basal spur bearing posterior margin of scutellum evenly crossvein r-m; rm index about 1.3. rounded; base of meron in line with base Crossvein m-cu slightly angled. Cell of hindcoxa; cell R2 of wing short; cell Cu1 with sclerotized furrow in front of Cu1 of wing with sclerotized furrow in vein Cu2 leading to angled submarginal front of vein Cu2 leading to an angled thickening. Plical vein strongly marked. submarginal thickening; abdomen com- Vein 1A long, ending on margin far pletely covered with scales and with distad of branching of vein Cu. Stem conspicuous caudal tufts of setae. The vein without dorsal bristles. Vein Sc only observed feature of the fossil species without bristles at base ventrally. Alula that deviates significantly from the con- and upper calypter with a few very small dition in extant species of the genus is its marginal setae. Halter: Knob apparently possession of a strong bristle located with pale scales. Abdomen (Fig. 4): In- posteriorly on the postpronotum. tegument light brown. Terga dark scaled Four subgenera of Toxorhynchites are dorsally, most scales apparently with recognized: Afrorhynchus Ribeiro and purple iridescence, but some in middle Toxorhynchites s. s. in the Old World, of tergum II with green iridescence, some and Ankylorhynchus Lutz and Lynch- on tergum VIII with bright green irides- iella Lahille in the New World. The cence; tergum I with very numerous three long-recognized subgenera Ankylo- bristles; laterotergite apparently without rhynchus, Lynchiella, and Toxorhynchites dense vestiture of scales, its lower area differ from each other in structure of the with numerous setae; sides of terga II– female palpus and antenna (Edwards VII with pale scales except at apex of 1932). In Toxorhynchites s. s., the palpus each segment; lateral margin of terga VI, of the female is short, only about 0.20– VII, and VIII with tufts of long brown or 0.25 the length of the proboscis, has only black setae, some of the setae on VIII two obvious palpomeres, and is blunt; in possibly red. Sterna scaled, color of most Lynchiella, the palpus is about 0.67 the 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON length of the proboscis, comprised of important. Because it is embedded in three obvious palpomeres, and blunt; amber, we have only limited knowledge and in Ankylorhynchus, the palpus is of the iridescent and metallic colors of about as long as the proboscis, has three the scales of T. mexicanus and therefore obvious palpomeres, and is pointed and we cannot provide additional diagnostic upturned. Ankylorhynchus also differs characters for the species based on color. from the other subgenera in having the All tarsi of T. mexicanus appear to be antenna of the female subplumose. Rel- completely dark scaled, but we cannot be atively recently, some African species certain that inconspicuous streaks of (the lutescens group of Edwards 1941) whitish or bluish scales similar to those were segregated from Toxorhynchites s. that occur on one side of the tarsus in s. into the subgenus Afrorhynchus on the some extant species of the genus are truly basis of several colorational and male absent. The caudal tufts of setae of claw and genitalic characters (Ribeiro extant species of Toxorhynchites may be 1992, 2005). white, yellow, orange, red, brown, or In T. mexicanus, the palpus is short, black. In T. mexicanus, the setae of these about 0.2 the length of the proboscis, tufts are identical in color to the dark blunt, comprised of two obvious, rela- scales of the right hindtibia that lies tively slender palpomeres, and the an- adjacent to the abdomen, leading us to tenna is sparsely plumose. These diag- believe that the tufts are brown or black. nostic characters place the species into However, some of the setae of the tufts one of the two Old World subgenera, on segment VIII that have become either Afrorhynchus or Toxorhynchites s. detached and lie in the amber near the s. Although we cannot discern most of tip of the abdomen have a decided red the colorational characters that distin- cast, raising the possibility that the tufts guish these subgenera, we place T. on VIII were partly red in life. mexicanus in the nominal subgenus The occurrence of a fossil species of because those parts of the pleural in- Toxorhynchites s. s. in the North Amer- tegument that can be seen are uniformly ican tropics in the mid-Tertiary is an black (rather than yellow or tan), the unexpected disjunct distribution. In gen- hindtibia and hindtarsomere 3 are dark eral, tropical intercontinental disjunc- scaled (rather than one or both having tions can have three explanations: vicar- a conspicuous white patch or ring), the iance, overland dispersal through the caudal tufts of setae on terga VI and VII boreotropics, and long-distance chance are brown or black (rather than yellow, dispersal across an ocean barrier (Givn- orange, or red), and the rm index is 1.3 ish and Renner 2004). We think it highly (instead of greater than 2.0). The fossil unlikely that a mosquito like Toxo- species does not appear to belong to any rhynchites could have crossed the Atlan- of the species groups of Toxorhynchites s. tic by long-distance dispersal. In his s. recognized by Steffan and Evenhuis study of the mosquitoes of the South (1985) and Ribeiro (1991). Pacific, Belkin (1962) noted that the Most taxonomic characters of adult majority of mosquitoes do not seem to Toxorhynchites are colorational, with the be able to cross ocean barriers of even iridescent or metallic colors of the scales a few miles, as shown by their regional of the head and its appendages (palpus differentiation within island groups. The and male flagellomere 1 in particular), exceptions to this generalization are a few the thorax and its appendages (legs widespread South Pacific species of especially, but also the wing), and the Aedes, Coquillettidia, and Culex that abdomen (both terga and sterna) being breed in ground pools in open areas VOLUME 110, NUMBER 1 123 and that may attain high population (1978) reported that in the Samoan densities. According to Belkin, although Islands Toxorhynchites amboinensis (Do- adults of these species may have been leschall) was first collected on Upolu dispersed passively by winds, dispersal of Island 15 years after its introduction to the species may have occurred in the egg Tutuila Island, which lies 72 km to the stage instead. In the case of Aedes, the southeast. Since the mosquito was large- eggs are laid on damp soil at the edges of ly restricted to the lee side of Upolu breeding sites and once embryonated are where, moreover, several harbors, resistant to desiccation. When the soil wharves, and an airport are located, and eggs are dry, it is possible that the and since its immatures were frequently eggs could be picked up by strong winds found in artificial containers (including and blown across ocean barriers. In the tires and large drums), the authors case of Coquillettidia and Culex, the eggs believed the mosquito was more likely are laid in rafts that float on the water introduced through commerce than by surface. It is possible that egg rafts could windblown aerial dispersal. cling to the feathers or legs of waterfowl There are actually two disjunctions of and be carried to distant sites through Toxorhynchites that need to be ex- the active flight of the birds. Adults of plained. The first is the occurrence of Toxorhynchites are never abundant and the genus in the tropics of both the New are largely restricted to sylvan situations and Old Worlds, and the second is the where they rest in protected sites. We do occurrence of the subgenus Toxorhynch- not believe it is likely that an adult ites in Mexico in the mid-Tertiary. As for Toxorhynchites could be picked up by the presence of Toxorhynchites in the winds and be blown across an extensive tropics of both hemispheres, we believe ocean barrier. Eggs of Toxorhynchites that the genus was present in West are deposited in phytotelmata, are not Gondwanaland before its breakup in resistant to desiccation, and hatch within the mid–Cretaceous. Subsequent to the 40 to 60 h (Steffan and Evenhuis 1981). breakup, the subgenera Ankylorhynchus We do not believe there is any possibility and Lynchiella differentiated in South that eggs of Toxorhynchites could be America during that continent’s long blown out of phytotelmata, be trans- period of isolation as an island, and ported long distances by the wind, and Afrorhynchus differentiated in Africa. In be redeposited in a habitat suitable for this regard, it is undoubtedly significant development of the immature stages. that today Ankylorhynchus occurs only The introduction of exotic species of in South America, the majority of species Toxorhynchites to some islands of the of Lynchiella occur only there as well (a South Pacific for biological control of few species extend into or are limited to container-breeding mosquitoes and their North America and the West Indies), subsequent discovery on nearby islands and Afrorhynchus is restricted to Africa. have generated speculation on their We are not the first to invoke vicariance methods of dispersal. Lever (1943) re- as the explanation for the amphi-Atlan- ported that in the Fiji Islands Toxo- tic distribution of Toxorhynchites. Near- rhynchites splendens (Wiedemann) ap- ly 70 years ago, Edwards (1941) stated: peared on Mango Island nine or ‘‘It cannot be a coincidence that the 10 years after its introduction to Vanua- genera and subgenera [of mosquitoes] balavu Island. He believed the had common to Africa and South America crossed the 19 km of open sea between (see p. 450) are those which on morpho- the islands by aerial dispersal aided by logical grounds are regarded as the most the southeast trade wind. Engber et al. primitive. The explanation seems obvi- 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ous, that these groups were present in perhaps as recently as the Pliocene when both regions before the final separation the modern Central American Isthmus of Africa and South America in creta- formed, or was brought about by other ceous times.’’ Edwards’ list on page 450 biotic or abiotic factors independent of includes the genus Megarhinus (5Tox- that event. orhynchites). Toxorhynchites s. s. is largely restricted ACKNOWLEDGMENTS to the Afrotropic and Indomalayan We thank Sandra S. Shanks for ecozones, but with extensions into the suggesting improvements to an earlier eastern Palaearctic and northern Austra- draft of this manuscript and James E. lasian ecozones. The majority of the Pecor for information about species of extant species and species groups occurs Toxorhynchites in the collection of the in southern Asia. We believe the sub- National Museum of Natural History, genus or its progenitors dispersed from Smithsonian Institution. Africa to Eurasia in the late Cretaceous or Paleocene and radiated there. As for LITERATURE CITED the occurrence of Toxorhynchites s. s. in Barraud, P. J. 1934. Family Culicidae. Tribes southern North America in the mid- Megarhinini and Culicini. The fauna of British Tertiary, we postulate that it migrated India, including Ceylon and Burma. Diptera. Vol. V. 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