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Alena Nav—S, 1916 – the Dethroned Genus and Alena (Aztekoraphidia

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Alena Nav—S, 1916 – the Dethroned Genus and Alena (Aztekoraphidia Dtsch. Entomol. Z. 60 (1) 2013, 53–58 / DOI 10.1002/mmnd.201300006 Alena Navs, 1916 – the dethroned genus and Alena (Aztekoraphidia) michoacana sp. n. from Mexico (Neuropterida: Raphidioptera: Raphidiidae) Ulrike Aspck*,1,2 and Horst Aspck3 1 Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, 1010 Vienna, Austria 2 Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Vienna, Austria 3 Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University of Vienna (MUW), Kinderspitalgasse 15, 1090 Vienna, Austria Abstract Received 11 December 2012 The new concept of the genus Alena, in which it is no longer the sister group of all Accepted 13 December 2012 other Raphidiidae but only of the Palaearctic clade, is presented and discussed. A. (Az- Published 22 May 2013 tekoraphidia) michoacana sp. n. is described and differentiated from other species based on the male holotype. In Alena the so-called basal sclerites which connect the gonocoxites 9 with the gonapophyses 9 are homologized with the gonangulum hitherto Key Words only ascribed to female genitalia. The homology is based on sclerites known from Ma- chilidae in both sexes. The hypothesis that the gonapophyses 9 are terminally amalga- New species mated into a complex, which comprises the fused gonocoxites, gonostyli, and gonapo- Phylogeny physes 10, is corroborated by A. michoacana sp. n., in which an intermediate state of Taxonomy this connection is evident. The geographic restriction of Alena species to high altitudes Genital sclerites of Mexico (and in two species – to the southernmost parts of the USA) is interpreted Gonangulum as a radiation following multiple vicariance events after the arrival of a more general- Biogeography ized stem species. Introduction unexpected and surprising result represents a manifold challenge involving not only phylogeny, but also bio- For many years and until recently the genus Alena was geography and particularly the morphology of the geni- unquestionably regarded as the sister to all other extant tal sclerites. Raphidiidae (H. Aspck et al. 1991; U. Aspck & H. Alena is a small genus currently comprising nine Aspck 1996; H. Aspck 1998; H. Aspck et al. 2001; species (U. Aspck & H. Aspck 1996, U. Aspck & H. Aspck & U. Aspck 2007). With respect to the bi- Contreras-Ramos 2004). The discovery of these species zarre appearance and shape of the genital sclerites Ale- spanned about 110 years, Alena australis (Banks, 1895) na is the most eccentric genus of the Raphidiidae – a being the first and Alena horstaspoecki U. A. & Con- fact which supported this assumption. Moreover, the treras-Ramos, 2004, the latest. Snakeflies are among geographic restriction of the distribution to the south- the rarest insects in Mexico, and, in particular, every ernmost parts of North America (predominantly Mexi- record of a specimen of Alena merits attention. Over co) was another supporting argument. the past one hundred years less than 50 imagines of However, a molecular study on the phylogeny of the Alena have been collected. Snakeflies from Mexico are Raphidiidae (Haring et al. 2011) revealed that rather thus extremely seldom in collections, even of the lar- than Alena the distinctly less spectacular genus Agulla, gest institutions. The finding of any new species of which is also confined to the Nearctic and distributed Alena was and will always remain an exciting event. throughout the western parts of North America, is the Thus, the description of Alena (Aztekoraphidia) mi- sister to all other Raphidiidae. In that study, Alena choacana sp. n. presented in this paper is an entomolo- emerged as sister to all Palaearctic Raphidiidae. This gical highlight. * Corresponding author, e-mail: [email protected] # 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 54 Aspo¨ck,U.& Aspo¨ck,H.: Alena and A. michoacana sp. n. from Mexico (Raphidioptera) Results Head “triangular”, black with a fine sculpture, and with a dark brownish median fascia, clypeus and labrum Alena Navs, 1916 ochre brownish. Antennae: scapus, pedicellus and basal third of flagellomeres yellowish, flagellum otherwise Alena Navs, 1916 (original description) (type species Raphidia dis- dark brownish. Pronotum: laterally with broad yellow tincta Banks, 1911): H. Aspck et al. 1991, U. Aspck et al. margin, anterior half brownish, caudal half dark brown- 1994, U. Aspck & H. Aspck 1996, U. Aspck & Contreras-Ra- mos 2004. ish with lighter brownish pattern. Legs: predominantly yellow, third coxae brownish. Wings: membrane hya- The genus comprises three subgenera: line, venation predominantly ochre brownish, subcosta Alena s. str. (with one species: A. distincta (Banks, darker; basal part of Media anterior (Ma) in hind wing 1911)), Mexicoraphidia U. A. & H. A. 1970 (with one as a crossvein. Pterostigma yellow, above the middle of species: A. americana (Carpenter, 1958)), and Aztekor- the first pterostigmal cell, with one vein crossing. Ab- aphidia U. A. & H. A., 1970 (with seven species: A. domen: tergites and sternites blackish brown, the latter caudata (Navs, 1914), A. infundibulata H. A. & U. A. caudally broad yellowish. Genital segments conspicu- & Rausch, 1994, A. schremmeri H. A. & U. A. & ously enlarged and yellowish. Rausch, 1994, A. minuta (Banks, 1903), A. australis Male genital segments (Figs 1, 3–5): Sternite of seg- (Banks, 1895), A. tenochtitlana U. A. & H. A., 1978, ment 8 only slightly shorter than tergite; tergite 9 trape- A. horstaspoecki U. A. & Contreras-Ramos, 2004). ziform, dorsally with median suture; gonocoxites 9 A. distincta and A. minuta have also been found in forming huge plates with a digitiform apex; gonostyli southwestern parts of the USA, all other Alena species large and stout, apically falcate; sternite 9 indistinct; are recorded only from Mexico. distinct large basal sclerites between gonocoxites 9 and gonapophyses 9 (hypovalva); paired gonapophyses 9 Alena (Aztekoraphidia) michoacana sp. n. terminally connected with huge paired sclerites consist- ing of a domed plate with dorsal and ventral processes. Derivatio nominis. The specific epithet is derived from This complex structure is interpreted as the fused gono- Michoacn, that state of Mexico, where the new species coxites, gonapophyses, and gonostyli 10 (called para- was discovered. The name is a noun of feminine gender meres); hypandrium internum huge; ectoproct (tergites in the nominative and an apposition to the name of the 10 þ 11) lacking distinct trichobothria, consisting of a genus. dorsal plate and lateral arms, which are terminally Material studied. Holotype, male: “Carapan, Michoa- rounded. can, Mex. VII-1-63 /Collr: W. A. Foster/Berkeley, v.C.” Differentiation. According to the morphology of the [195103600 N, 1020201000 W, ca. 2000 m], in collection gonocoxites 9 and the gonostyli 9 Alena michoacana of the Californian Academy of Sciences. sp. n. clearly belongs to the subgenus Aztekoraphidia. Morphology (Figs 1, 3–5). A small tender species, From the three Alena species which have the Ma in the length of forewing 8 mm, habitus dominated by yellow hind wing as a longitudinal vein – A. (M.) americana, elements (pterostigma, legs and genital segments). A. (A.) caudata, and A. (A.) horstaspoecki – A.(A.) mi- Figure 1. Habitus of Alena (Aztekoraphidia) michoacana sp. n., holotype, male. Arrow indicates the basal piece of Media anterior in the hind wing. 2. Original labels of Alena (Aztekoraphidia) michoacana sp. n., holotype, male. museum-dez.wiley-vch.de # 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Dtsch. Entomol. Z. 60 (1) 2013, 53–58 55 Figures 3–5. Alena (Aztekoraphidia) mi- choacana sp. n., holotype. Male genital segments, dorsal (3), ventral (4), lateral (5). e – ectoproct; bs – basal sclerite (interpreted as homologous to the female gonangulum); gx9 – gonocoxite 9; hi – hypandrium inter- num; hv – hypovalva (gonapophyses 9); p – fused parameres (complex of fused go- nocoxites, gonapophyses and gonostyli 10); s – gonostylus; S – sternite; T – tergite. choacana sp. n. can be separated eidonomically since assign one of the female specimens to A. michoacana. the Ma in the hind wing is a crossvein. A. (A.) distinc- Clarification would require finding further male speci- ta, A. (M.) americana, and A . (A.) caudata can also be mens, which may not occur in the immediate future. separated eidonomically since the pterostigma has an Distribution. The distribution of A. michoacana sp. n. apical vein. A. (A.) michoacana can be separated from is almost certainly restricted to high altitudes within a all other Alena (Aztekoraphidia) species by the shape small area of southwest Mexico, possibly only in the of the distinctly monstrous “parameres” (fused gono- state of Michoacn. coxites, gonostyli, and gonapophyses 10) which appear as simple acute apices of the gonapophyses 9 (hypoval- Discussion. In connection with the dethronement of va) in A. (A.) minuta, A. tenochtitlana, A. infundibulata Alena as the sister group of all other Raphidiidae and and A. schremmeri or have ventrally protruding bottoms the shift of Agulla to the most basal position in our in A. australis. There is no particularly close relation- molecular analysis (Haring et al. 2011), the morpholo- ship to any of the other Aztekoraphidia species. gical, phylogenetic and biogeographical implications Female unknown. In addition to the male holotype of need to be addressed. A. michoacana sp. n., two female specimens (coll. TAMU, kindly provided by Dr. John Oswald, Texas A&M University, College Station, USA) are available Morphological implications from Michoacn (Fig. 6), each of them representing a different species, and none of them can be clearly as- First: The question arises what is the nature of the so- signed to the male. It would therefore be confusing to called basal sclerites (Figs 4, 5)? In Alena, they connect # 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-dez.wiley-vch.de 56 Aspo¨ck,U.& Aspo¨ck,H.: Alena and A. michoacana sp. n. from Mexico (Raphidioptera) Figure 6. Distribution of Alena in Mexi- co. The map shows not only the records of the described 10 species but also those of the unidentified females.
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