March, 30, 2016 Vol. 8, No 1

March, 30, 2016 Vol. 8, No 1

The European Entomologist, Vol. 8, No. 1 New species of Endoclita C. &. R. Felder, 1874 from Sumatra, Indonesia (Lepidoptera: Hepialidae) 1 2 1 2 Abstract Endoclita fahringeri sp. n. is described from a unique high elevation female from 3,000 m in the Leuser Mountains of northern Sumatra, Indonesia. Considerable loss of forewing scales precludes complete characterization of external appearance, but several distinct Endoclita spp. by forewing and hindwing shape and ornamentation. Female genitalia are compared with the limited descriptions available for other species. The conservation importance of this species is discussed with respect to biogeography and future economic pressures on the regional ecosystem. The female holotype will be deposited in the Museum Zoologicum Bogoriensis, Cibinong, Java, Indonesia. Dorsal images of the holotypes of raapi Keywords: biogeography, conservation, taxonomy, tectonics. Introduction The genus Endoclita widespread genera of the Hepialidae with about 60 recognized species and a geographic range encompassing the Russian Far East, India, Sri Lanka, China and Southeast Asia Endoclita larvae are almost certainly all callus feeding stem borers, although Nair ). The generic limits and monophyly of Endoclita and its sister group relationships are unresolved. The genus is almost by default a group of species not conforming to the Asian genera Palpifer Parahepialiscus Thitarodes Parathitarodes Phymatopus boring genera Aenetus Herrich-Schäffer, 1855 of Australasia and Phassus Walker, 1856 of Mexico/Central America that may prove to be phylogenetically informative in the future (Grehan & Rawlins 2003).

18 The European Entomologist, Vol. 8, No. 1 With the exception of the yellow-green Endoclita viridis Endoclita moths are various shades of brown with darker brown to black markings. Some species have distinctive wing patterns, but published illustrations are often of relatively poor quality and genital dissections either lack illustrations or are limited to generalized diagrams of externally visible features. Many species were reviewed or proposed in the remains problematic, if not impossible, due to the lack of detailed published information on type specimens. This limitation has been offset to some extent by the willingness of several institutions to provide high quality digital images for public access (see http:// www.johngrehan.net/index.php/hepialidae/endoclita). a specimen of Endoclita he collected in the Leuser mountain range in northern Sumatra aroura eastern Sumatra, and of the type specimens of these species by the Natural History Museum, London, and the Naturalis Biodiversity Center, Leiden, it was possible to recognize that the Leuser specimen showed species level differences in forewing pattern and wing shape. In this paper, we describe the external characteristics of Endoclita fahringeri sp. n. and comment on its potential relationships with other Endoclita spp. and the features that Endoclita as currently known or possible subclades within the genus. In addition, brief comments about biogeography are presented for the newly described species and Endoclita. Materials & Methods internal genitalia were dissected. Terminology follows that of Mielke & Casagrande (2013).

The European Entomologist, Vol. 8, No. 1 Abbreviations HT (holotype), FW (forewing), HW (hindwing), Taxonomy section Endoclita fahringeri sp. n. Holotype Endoclita fahringeri Zoologicum Bogoriensis, Cibinong, Java, Indonesia. Figs. 1a, 1b. Etymology. Endoclita fahringeri sp. n. is named after the collector, Alfred Fahringer, who led the expedition to explore the Leuser, and encouraged observation of the insect fauna by members of his group that led to collection of the specimen and brown scales (Fig. 2b). Eyes prominent (Fig. 3a). Mouthparts: prelabium pentagonal pale greyish-brown. FW costal margin slightly convex and widely separated from Sc almost straight from Cu1a to A veins, then posterior margin with shallow curve to base. distally terminating beyond jugum. FW dorsal ground colour pale greyish brown with small, irregular, dark ovoid or sub-elliptical patches of dark brown with pale yellowish

20 The European Entomologist, Vol. 8, No. 1 humeral vein between costal margin and Sc, concave with anterior surface free of scales and posterior surface with short dark brown pilose scales (Fig. 5c). HW dorsally and ventrally dark greyish-brown, except for ornamentation along costal and outer margins surface of tarsal segments with short scales resistant to removal. Abdomen greyish- narrower than tergum VII (Fig. 8a). genitalia which forms a laterally broad sclerotized concavity and dorsally, projected dorsally and wall of the concavity reaches the antrum. Ductus and corpus bursae of similar length. Geographical distribution: inferred to be widespread at higher elevations in the Leuser Mountains. Remarks: Endoclita ssp. remain unresolved at this time due to lack of comparative morphological descriptions and illustrations of other species. Diagnosis: Distinguished from all other Endoclita ssp. by its elongated sub-triangular FW with of ovoid sub-elliptical dark patches with pale borders visible between M 2 -CuA 2 where scales remain intact. Comparison of the external features of sp. n. with those of all the other four named species present on the island of Sumatra supports sp. n. status as a distinct species. Each of the previously known species contrasts with

The European Entomologist, Vol. 8, No. 1 21 sp. n. in FW ornamentation and wing shape. The wing patterns of some types are also in poor condition, although still distinct. (Fig. 11), (Fig. 12), and taranu (Fig. 13) have a falcate FW apex in contrast to sp. n. The subtriangular shape of the HW in is most like that of sp. n., but the is similar to that of sp. n., but without the concave outline of the costal margin and the HW lacks the triangular shape of sp. n. The presence of the and also precludes sp. n. The female genitalia of Endoclita ssp. are poorly illustrated for most species to identify similarities and differences with fahringeri sp. n. Most, if not all, species of Endoclita share with sp. n. the presence of three dorsally projecting lobes where the central lobe is usually strongly signifer (Walker, 1856) of northeastern India (Fig. 15) and Japan (Fig. 16). Features of general systematics interest: Endoclita with following features: antenna with about 22 2 HW without Sc1 and Rs branching well before midwing. With the possible exception of vein 2V enclosing a cell (see below), each of these features is also found in other Hepialidae so none of them individually diagnoses Endoclita. The presence of Sc1 appears to be relatively rare in Hepialidae although the vein can be very faint and easily overlooked. It is present in other families of Lepidoptera and is Endoclita appears to represent a derived condition within Hepialidae as it is absent from other exoporian molecular phylogeny of these moths (Regier et al. 2015). The illustration of basal venation in cells in sp. n. suggests that this arrangement will be present in many, if not all, Endoclita spp. If present in only some species, this feature may indicate a subclade of Endoclita of these basal cells and veins to all Endoclita they would potentially represent one or

22 The European Entomologist, Vol. 8, No. 1 more shared derived character states for this genus, provided also that this feature is Two features in E. fahringeri sp. n. that came to our attention do not appear to have previously received general comment in the literature: the costal pocket of the ventral FW (Fig. 5c) and the absence of scales over most of the dorsal surface of scutum III second, after cursory examination of several hepialid genera, suggests that a lack of scales over part, or all, of this region is widespread. Further observations of its condition in the Hepialidae and outgroups will be necessary for adequate interpretation of its sp. n. scales are missing from more than half of the dorsal surface from the anterior margin of the scutum III and the exposed surface becomes dorsally visible when the wings are spread. This condition is present in all Endoclita spp. for which detailed photos are available. After examining specimens of most genera we conclude that the absence of scales from the dorsal surface of the scutum III probably applies to all Hepialidae, but the surface is not dorsally visible when posterior mesothoracic scales are long enough to cover the metathoracic scutum. Further observations on scutum III exposure in Hepialidae and outgroups are necessary to assess any potential phylogenetic information for this feature. Conservation Status: The occurrence of sp. n. at high elevation is of evolutionary and ecological interest. The species occurs in a region of tectonic uplift from Mesozoic time and includes As areas of low topography are continually being uplifted to higher elevations, the plants and animals of the lowlands are also brought into higher elevations. For those organisms that survive in the higher elevations they may over time diverge from their Analysis of the temporal context for uplift and isolation of E. fahringeri sp. n. will require further resolution of its precise distribution and its sister group relationships with other Endoclita species. Because Endoclita larvae require live tree hosts in a forest environment, their survival depends on the continuity of forest ecosystems, whether in their original condition or sp. n. was collected thick peat with a ground cover of orchids, low bushes, mosses, lichens, and liverworts, fahringeri sp. n. specimens from historical collections in lowland habitats of Sumatra provides a strong indication that the species occupies higher elevation habitats, even

The European Entomologist, Vol. 8, No. 1 23 if not necessarily strictly at the elevation of Bipak Ridge. The Leuser Ecosystem of approaches to ecosystem management. The main reason for this destruction of forest despite conservation measures lies in the strong political power of the logging conservation. This situation prevents the most sustainable approach from occurring (van sp. n. remains contingent upon the future extent of environmental threats to the Leuser ecosystem in general. Acknowledgements his willingness to provide the specimen for description. We are very thankful to John Rawlins (Carnegie Museum of Natural History, Pittsburgh) for his support for this project and providing research facilities and resources, and to Jane Hyland (Carnegie Museum of Natural History, Pittsburgh) for photographing the type specimen. We are also deeply grateful to Rob de Vos (Naturalis Biodiversity Center, Leiden), Alessandro Giusti and Geoff Martin (Natural History Museum, London) and to Thomas Witt (Witt Museum, München) for providing photos of Endoclita spp. for comparison. Finally, we thank Dr. Ian Kitching (Natural History Museum, London) and Dr. Rodolphe Rougerie (Muséum nationale d Histoire naturelle de Paris, Paris) for their valuable suggestions and corrections. Fig. 1: Endoclita fahringeri sp. n Photo by Jane Hyland.

The European Entomologist, Vol. 8, No. 1 Fig. 2: Endoclita fahringeri sp. n Photo by John Grehan. Fig. 3: Endoclita fahringeri sp. n., HT m = mandible, lp = labial palp, mp = maxillary palp.) - unscaled. Photo 3a by Jane Hyland, 3b by John Grehan. Fig. 4: Endoclita fahringeri sp. n., HT Photo by John Grehan.

The European Entomologist, Vol. 8, No. 1 25 Fig. 5: Endoclita fahringeri sp. n position of Sc1 marked with arrows (5b), ventral FW costal pocket with anterior surface free of scales, posterior half with short, pilose scales (5c), FW distal position of Sc and R at costal margin (5d) - unscaled. Photo by John Grehan.

26 The European Entomologist, Vol. 8, No. 1 Fig. 6: Endoclita fahringeri sp. n unscaled. Photo by John Grehan.

The European Entomologist, Vol. 8, No. 1 Fig. 7: Endoclita fahringeri sp. n Photo by John Grehan. Fig. 8: Endoclita fahringeri sp. n genitalia (8a), terminal segments with orange-brown trilobed lamella antevaginalis and darkly sclerotized antrum (8b) - unscaled. Photo 8a by John Grehan, 8b by Jane Hyland.

28 The European Entomologist, Vol. 8, No. 1 Fig. 9: Endoclita fahringeri sp. n unscaled. Photo by John Grehan. Fig. 10: Endoclita fahringeri sp. n. distribution within Sumatra, Indonesia.

The European Entomologist, Vol. 8, No. 1 Fig. 11: Endoclita niger Photo by Rob de Vos. Fig. 12: Endoclita raapi Photo by Alessandro Giusti.

30 The European Entomologist, Vol. 8, No. 1 Fig. 13: Endoclita taranu Photo by Alessandro Giusti. Fig. 14: Endoclita aroura Photo by Alessandro Giusti.

The European Entomologist, Vol. 8, No. 1 31 Fig. 15: Endoclita nr signifer Collection, JRG 235, Buffalo, USA). Photo by John Grehan.

32 The European Entomologist, Vol. 8, No. 1 Fig. 16: Endoclita signifer Germany). Photo by John Grehan.

The European Entomologist, Vol. 8, No. 1 33 Fig. 17-18: Type locality of Endoclita fahringeri sp. n. Alfred Fahringer and his team along Bipak Ridge trail. Photos by Alfred Fahringer.

The European Entomologist, Vol. 8, No. 1 References Acta Entomologica Sinica 28 Fauna Sinica. Insecta Vol. 38. Lepidoptera: Hepialidae, Dacnonypha (Lepidoptera) based on three new species from southern Australia, with notes on the Agathiphagidae. Journal of the Australian Entomological Society 12: 11-23. (Lepidoptera). 9 the monophyly of the Latin American genera Walker, Druceiella Viette, Viette and Trichophassus Le Cerf (Lepidoptera: Hepialidae). 39 Grehan, J. R. & Rawlins, J. E. (2003): Larval description of a new world ghost moth, Phassus sp., and the evolutionary biogeography of wood-boring Hepialidae (Lepidoptera: Exoporia: Hepialoidea). Proceedings of the Entomological Society of Washington 105 Indonesia,. In: Gillespie, R. & Clague, D. (eds.). Encyclopedia of Islands. Berkeley, Tectonophysics 570-571. Cambridge, Cambridge Phassus signifer and its some ecological notes. Transactions of the Tottori Society of Agricultural Science 13 Liang, J. Y. & Lee, M. J. (2011): Status of forest trees infested with Endoclita sinensis (Lepidoptera: Hepialidae). 25: 203-210. Butler. Kontyu 34 Mielke, C. G. C. & Casagrande, M. M. (2013): A new Walker, 1856 from southern Brazil (Lepidoptera, Hepialidae). (N.F.) 34 (Lepidopera: Hepialidae). Entomonograph 12 Nielsen, E. S., & Robinson, G. S. & Wagner, D. L. (2000): Ghost-moths of the world: a global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera). 34 (Butler). Journal of the Sericultural Science of Japan 48

The European Entomologist, Vol. 8, No. 1 35 Regier, J. C., Mitter, C., Kristensen, N. P., Davis, D. R., Van Nieukerken, E. J., Rota, J., Simonsen, T. J., Mitter, K. T., Kawahara, A. Y., Yen, S. H., Cummings, M. P. & Zwick, A. (2015): A molecular phylogeny morphology and life-history evolution. Systematic Entomology 40: 7 South Australian Museum 7: 151-168. South Australia Museum 13 of the Teak sapling borer, (Moore) (Lepidoptera: Hepialidae) attacking coffee. International Journal of Advanced Life Sciences 7 Endoclita Felder & Rogenhofer (Lepidoptera: Hepialidae) from Taiwan. Tinea Supplement 12: 103-108 Park on Sumatra, Indonesia. Ecological Economics 44 species (Lepidoptera: Hepialidae). Entomological Journal of East China 1: 10-16. Endoclita signifer (Lepidoptera: Hepialidae) as a new pest on Eucalyptus. Journal of Economic Entomology 106 Endoclyta signifer Walker (Lepidoptera: Hepialidae), a new wood borer on Eucalyptus. 26 (1): Endoclita hosei Tindale (Lepidoptera: Hepialidae) attacking eucalyptus in Sabah, with descriptions of the immature and imaginal stages. 21 10