KJELL ARNE JOHANSON
Swedish Museum of Natural History
THE SISTER SPECIES OF JAMAICAN HELICOPSYCHE KINGSTONA SP. N., IS MEXICAN H. VILLEGASI DENNING & BLICKLE (TRICHOPTERA, HELICOPSYCHIDAE)
Johanson, K. A., 2003. The sister species of Jamaican Helicopsyche kingstona sp. n., is Mexican H. villegasi Denning & Blickle (Trichoptera, Helicopsychidae). – Tijdschrift voor Entomolo- gie 146: 33-37, figs. 1-9. [ISSN 0040-7496]. Published 1 June 2003. Helicopsych kingstona sp. n., is described from Jamaica. Its sister species, H. villegasi Denning & Blickle is found in Mexico, suggesting past exchange of biota between Jamaica and Mexico. Correspondence: Kjell Arne Johanson, Swedish Museum of Natural History, Box 50007, S- 104 05 Stockholm, Sweden. E-mail: [email protected]. Keywords. – Trichoptera; Helicopsychidae; Helicopsyche kingstona; new species; Jamaica; ville- gasi; Mexico; sisterspecies.
Eight of microcaddisflies (Hydroptilidae) and one (1998) inferred that the Helicopsyche of the Greater long-horned caddisfly (Leptoceridae) share a distribu- Antilles are polyphyletic and originated independent- tion that includes both the Greater Antilles and the ly on the islands several times. mainland (Flint et al. 1999), indicating a recent dis- In this paper, a new species, Helicopsyche kingstona, is persal of biota between the two regions. Six of these described from Jamaica. Together with Helicopsyche species are recorded from Jamaica, including Oecetis kingstona sp. n., three other Helicopsyche species, H. inconspicua (Walker), which is widespread from Cana- molesta Botosaneanu, H. ochthephila Flint, and H. um- da to Peru, including several islands in the Greater An- bonata Hagen, are known from Jamaica, all endemics. tilles; Oxyethira simulatrix Flint, also recorded from Specimens of H. cubana Kingsolver were reported Costa Rica; O. janella Denning, also collected from from Jamaica by Flint (1968), but Botosaneanu (2002) the USA, Costa Rica, and several Greater Antillean is- considered these specimens to belong to H. molesta. lands; O. arizona Ross, with a similar distribution as The far richer Mexican Helicopsyche fauna includes 15 O. janella, but also known from Mexico; and Or- species, of which eight are endemic. One species, H. thotrichia cristata Morton, known also from Canada, villegasi Denning & Blickle was described from Za- Cuba, and the USA . catecas, 4 km W of Nochistlan, but has never been col- The American Helicopsyche fauna is divided into lected again. Helicopsyche kingstona is inferred to be the two subgenera, Cochliopsyche Müller, 1885 (four sister species of H. villegasi as indicated by two synapo- species) and Feropsyche Johanson, 1998 (72 species) morphies. This indicates a past faunal exchange of He- (Johanson 2002). Since no species of Helicopsychidae licopsychidae between Mexico and Jamaica. are in common between the Greater Antilles and the mainland, no recent dispersal between the two areas Helicopsyche kingstona sp. n. appears to have occurred. However, no comprehen- (figs. 1-8) sive phylogenetic or biogeographic analysis of Ameri- can Helicopsychidae is presently available such that no Type material. – Male holotype: Jamaica: 5 mi di- past biogeographical relationship between the faunas rectly W of Kingston, TRC, ACS [Sears Gardner of the two area has been hypothesized. Johanson Glaser] (University of California, Davis, in alcohol).
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Figs. 1-8. Helicopsyche kingstona. sp.n., male holotype. – 1, Right wings; 2, VIth sternal process, lateral view; 3, VIth sternal process, ventral view; 4, genitalia, lateral view; 5, gonocoxite, median view; 6, genitalia, dorsal view; 7, genitalia, ventral view; 8, phallus, lateral view.
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Diagnosis 4), slender along its length, slightly curved ventrad, ta- Both H. villegasi Denning & Blickle and H. pering slightly toward truncated apex; basal incision kingstona are large species, having the forewing longer absent; in dorsal view (fig. 6), with slightly converging than 6 mm. In the male genitalia, the two species share lateral margins, apically rounded with small central a uniquely bilobed posterior margin of sternite IX and notch; with about 11 moderately long megasetae in a prominent subapical median tooth on the gonocox- longitudinal rows starting proximally on segment; 5 ite. The shape of the basimesal lobe of the gonocoxite small megasetae on apex. Superior appendage (fig. 4), and the numerous megasetae along the median mar- club shaped, oriented posteriad. Primary branch of gin in ventral view is shared also with H. turbida gonocoxite, in lateral view (fig. 4), approximately rec- Navás. Also, the gonocoxite of H. kingstona is slightly tangular; dorsal margin undulate; apex curved ven- wider than in H. villegasi.Helicopsyche kingstona can be tromesad, visible in lateral view; gonocoxite medially separated from other species by its longer antennal nearly 3ϫ broader than height of central part of Xth scape and more slender median process of abdominal tergum (figs. 4, 5); anterodorsal margin slightly sig- sternum VI. moid, smooth; posteroventral margin slightly undu- late, ventral part concave, smooth. Basimesal lobe (fig. Etymology 5) broad, rhomboid, apex visible in lateral view (fig. 4), H.kingstona, named after the type locality. To be with numerous megasetae on apex and dorsomedial treated as a noun in apposition. face (fig. 5); in ventral view (fig. 7), produced medially into rounded plate bearing numerous, marginal Description megasetae; median margin almost straight, with about Male – Head: Antennal scape length about 1.75ϫ 2 small, marginal setae. Basal plate stout (figs. 4, 5), diameter of eye, scapes parallel and very close along with slightly convex dorsal and ventral margins; in ven- their lengths. Maxillary palp segments equally long, tral view (fig. 7), narrowing anteriorly, apex truncate. each as long as eye diameter. Cephalic warts very large, Phallus (fig. 8): slender; strongly bent ventrad sub- rounded, strongly convex. Postantennal warts small, basally; equally thick along its length, except anterior club-shaped, located immediately behind scapes. Fore- one-sixth about two-times thicker than central part; leg anterior tibial spur about 2ϫ longer than posterior phallbase wide; endotheca not produced into dorsal tibial spur. Wings (fig. 1): Fore wing 6.3 mm, slender; lobe; sperm channel posteriorly thick, anterior one- fork 1 slightly longer than one-fourth wing length; fork sixth significantly more slender; posteroventral part 2 and discal cell about two-fifths wing length; crossvein weakly sclerotized. R-M long; M1 about one and one-half times longer than stalk of M1+2; M2 one-fifth of wing length; DISCUSSION crossvein M-Cu1 originates from M3+4 shortly after bifurcation of M and meets Cu1 at a distance before Helicopsyche villegasi and H. kingstona both have a bifurcation of Cu1 equal to one half length of Cu1a; uniquely derived bi-lobed posterior margin of sternite Cu2 ends in wing margin with crossvein connection to IX (1 in fig. 9) and a prominent subapical median tooth Cu1. Hind wing 5.1 mm, with 27 hamuli; fork 1 L- on the gonocoxite (2 in fig. 9). These characters are shaped; M divides distally to bifurcation of Rs; proposed as synapomorphies unambiguously support- crossvein R-M present; crossvein Cu1-M absent; apex ing the sister species relationship of the two species (fig. slightly hyperbolic. Vth sternal process (figs. 2, 3) ori- 9). Other striking features include the basimesal lobe of ented posteroventrad, almost straight; in lateral view the gonocoxite which is present as a median plate hav- slightly pointed, about 0.35 times segment length, ing a rounded posteromedian corner (3 in fig. 9) with does not reach segment VII; smooth; in ventral view marginal megasetae (4 in fig. 9); the general shape of (fig. 3), lateral margins parallel-sided, apex oval, cov- the primary branch of the gonocoxite; the number and ered by ventral lamellae. Genitalia (figs 4-8): Segment shape of Xth tergal setae. Also, the Argentine species H. IX, in lateral view (fig. 4), with anterior lobe located turbida Navás has the same characteristically shaped dorsally on segment, triangular, oriented anteriad, an- basimesal lobe and marginal megasetae of the gonocox- terodorsal margin almost straight; anteroventral mar- ite, but has a narrower primary branch and fewer, but gin shallowly concave; in dorsal view (fig. 6), with in- stronger setae on Xth tergum. The former characters ner margin widely elliptical; in ventral view (fig. 7), are synapomorphies for H. turbida + H. villegasi + H. with central, bifurcated process on posterior margin; kingstona (fig. 9). The latter two characters are autapo- lateral apodeme anteriorly almost straight, posteriorly morphies of H. turbida. curving ventrad, fades out before anterior margin be- Since H. turbida, the sister species of H. villegasi + low anterior lobe apex; tergal transverse apodeme ab- H. kingstona, occurs in Argentina, the ancestor of the sent; slightly developed sternal transverse apodeme group of three species was widely distributed on the along posterior margin. Segment X, in lateral view (fig. mainland. Subsequently, secondary dispersal to the
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Fig. 9. Phylogenetic hypothesis of H. kingstona, H. villegasi, H. turbida and other Helicopsyche subgenus Feropsyche. Numbers indicate synapomorphies discussed in text.
Greater Antilles occurred, followed by vicariant speci- (MacPhee & Iturralde-Vinent 2000). Much more re- ation. Since Jamaica was more or less submerged until cent landbridges were formed during the Plio-Pleis- early Miocene (Robinson 1994), reception of fauna tocene glacial cycles when the level of the sea repeat- from the mainland was probably not possible earlier, edly rose and fell, facilitating dispersal to the Greater giving a maximum age of H. villegasi and H. kingstona Antilles and resulting in subsequent vicariant specia- of less than 24 million years (MY). If more Greater An- tion. Alternatively, if the geographical separation of tilles islands were involved in the vicariance event of Jamaica and Mexico was already present, the ancestor the ancestor which affected the Greater Antilles and could have dispersed over an existing sea barrier be- Mexico populations, and with secondary dispersal to tween the mainland and Jamaica. The ancestral Jamaica, the maximum age could be as high as 36 MY. species could have reached the Antilles via aerial At that time northern South America, Cuba, Hispan- planktonic dispersal. If so, it would be impossible to iola and Puerto Rico formed a continuous land bridge place an exact date on any speciation event.
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