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A New Tube-Nosed Fruit Bat from New Guinea, Nyctimene Wrightae Sp

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A New Tube-Nosed Fruit Bat from New Guinea, Nyctimene Wrightae Sp © The Author, 2017. Journal compilation © Australian Museum, Sydney, 2017 Records of the Australian Museum (2017) Vol. 69, issue number 2, pp. 73–100. ISSN 0067-1975 (print), ISSN 2201-4349 (online) https://doi.org/10.3853/j.2201-4349.69.2017.1654 A new Tube-nosed Fruit Bat from New Guinea, Nyctimene wrightae sp. nov., A Re-diagnosis of N. certans and N. cyclotis (Pteropodidae: Chiroptera), and a Review of their Conservation Status Nancy Irwin School of Biological Sciences and Veterinary Science Department The University of Queensland, St Lucia Qld 4072, Australia Current address: Department of Biology, University of York, Wentworth Way, York, YO10 5DD, United Kingdom [email protected] Abstract. The tube-nosed fruit bat genus Nyctimene comprises 18 species found in the Philippines, Wallacea, Melanesia and the Solomon Islands but species taxonomy has remained problematic. A review of the cyclotis group, consisting of N. cyclotis and N. certans from New Guinea is presented, using morphological and genetic data. Historically, the taxonomy of the cyclotis group has been severely impeded by the lack of illustrations or photographs of the majority of the types. This led authors to differ in their treatment of these species, suggesting that the distinct taxa N. cyclotis and N. certans might be conspecific. Here, N. cyclotis and N. certans are recognized as full species and are re-diagnosed. I describe Nyctimene wrightae sp. nov., a widespread New Guinean species which is tentatively placed in the cyclotis group. These three species and the sympatric N. a. papuanus, of similar body-size, are morphologically distinguished using discriminant function analyses and non-metric characters. An identification key is provided. Species distribution and conservation status are evaluated. The IUCN threat status recommended for each species is: N. wrightae sp. nov. Least Concern; N. certans (known from < 200 specimens) with unknown population size and trends, Data Deficient; and N. cyclotis, known from only two male specimens, Vulnerable. Further research is required on the basic ecology of all of these species, which remains virtually unknown. Keywords. Nyctimene; taxonomy; revision; Papua New Guinea; Indonesia; cryptic species; conservation; IUCN threat category; Happy tube-nosed fruit bat Irwin, Nancy. 2017. A new tube-nosed fruit bat from New Guinea, Nyctimene wrightae sp. nov., a re-diagnosis of N. certans and N. cyclotis (Pteropodidae: Chiroptera), and a review of their conservation status. Records of the Australian Museum 69(2): 73–100. https://doi.org/10.3853/j.2201-4349.69.2017.1654 74 Records of the Australian Museum (2017) Vol. 69 The tube-nosed fruit bats Nyctimeninae (Miller, 1907) are a difficult when specimen numbers (e.g., Flannery, 1995a) or distinctive subfamily of Pteropodidae, currently represented measurements are omitted (e.g., Smith & Hood, 1983) and by 18 species and two genera, Nyctimene (16 spp.) and when taxonomic rank is changed without formal justification, Paranyctimene (2 spp.) (Simmons, 2005). The subfamily e.g., N. a. papuanus Andersen, 1910, N. bougainville and N. is distributed throughout the rainforests of Wallacea draconilla Thomas, 1922a, were all demoted to subspecies of (including the Philippines), New Guinea and its islands, to N. albiventer by Laurie & Hill (1954). Intraspecific variation southeastern Australia and the Solomon Islands (Flannery, remains to be determined for some taxa that were based 1995b; Simmons, 2005). The subfamily is characterized by either on only the holotype (e.g., N. malaitensis Phillips, protruding tubular nostrils, the absence of lower incisors and 1968; Paranyctimene tenax tenax Bergmans, 2001), or on a wing spotting. Individuals range in size from 45 to 85 mm very limited number of specimens (e.g., N. masalai Smith in forearm length. They have a distinctive dentition, with no & Hood, 1983) and for which additional material has not lower incisors, no upper and lower second premolars and no yet been determined. upper second molars. Sexual dimorphism in colour and size has been reported All authors during the last 40 years have found the for some species of Nyctimene, for example, N. a. papuanus, classification of Nyctimeninae problematic. No taxonomic N. major (Dobson, 1877) and N. cephalotes (Pallas, 1767) review has looked at the entire subfamily since Andersen’s (Kitchener et al., 1993, 1995; Andersen, 1912a), but not for revision over a century ago, which was based on only 61 other taxa such as N. certans Andersen, 1912b, N. cyclotis specimens in 13 species (Andersen, 1912a; Table 1). At Andersen, 1910 (Peterson, 1991) and N. keasti Kitchener, in present, 21 names are assigned to Nyctimeninae, though Kitchener et al. 1993 (Kitchener et al., 1995). Additionally, authors vary widely in the species they recognize, in some many of the types are damaged, have worn dentition (Smith cases with the same specimen being assigned to different & Hood, 1983; Peterson, 1991) or have lost described species (e.g., N. bougainville Troughton, 1936 / N. albiventer diagnostic alpha characters (e.g., in the holotype of N. minor Philips, 1968 / N. albiventer albiventer (Gray, 1863) cyclotis, the right P3 and P4 have fallen out of the skull and / N. vizcaccia Thomas, 1914). Recent appraisals of species are lost). High individual variation (Smith & Hood, 1983), have concentrated on island populations (Kitchener et al., variation in cranial and dental structures caused by aging 1993, 1995; Flannery & White, 1991), leaving mainland (Heaney & Peterson, 1984) and diagnoses that rely on size taxa in dire need of revision (Mickleburgh et al., 1992). and colour patterns (Smith & Hood, 1983; Bonaccorso, 1998) Contributing directly to the taxonomic confusion is the have made Nyctimeninae one of the most taxonomically absence of illustration for most species described before confused taxa in Chiroptera. the 1970s, or the exaggeration of characters depicted In their electrophoretic study of Australian and Papua in illustration, e.g., Andersen’s (1912a) treatment of N. New Guinean Nyctimene and Paranyctimene Donnellan et al. aello Thomas, 1900 versus N. major scitulus Andersen, (1995) did not resolve species boundaries despite sampling 1910 (Tate, 1942). Taxonomic clarity is also made more more than 30 allozymes from a large number of individuals Table 1. A summary of different taxonomic arrangements ofNyctimene taxa by selected authors subsequent to Andersen. Andersen, 1912a Andersen, 1912a Heaney & Peterson, 1984 Bergmans, 2001 Bergmans, 2001 supra-specific groups supra-specific groups N. albiventer N. albiventer albiventer N. albiventer N. papuanus (N. a. papuanus) (N. a. papuanus) N. minutus N. minutus N. minutus N. varius N. malaitensis N. malaitensis N. draconilla N. draconilla N. bougainville cyclotis N. cyclotis N. cyclotis cyclotis N. cyclotis N. certans aello N. aello N. aello aello N. aello N. celaeno (N. a celaeno) cephalotes N. cephalotes N. cephalotes cephalotes N. cephalotes N. major N. major N. major N. geminus (N. m. geminus) (N. m. geminus) N. lullulae (N. m. lullulae) (N. m. lullulae) N. scitulus (N. m. scitulus) (N. m. scitulus) N. robinsoni N. robinsoni N. robinsoni N. masalai N. masalai N. rabori N. rabori N. santacrucis N. santacrucis N. vizcaccia N. vizcaccia N. keasti Paranyctimene P. raptor P. tenax Irwin: New Guinea tube-nosed fruit bat 75 (n = 178). They delineated five species from New Guinea but Although Andersen (1912a) had only one specimen at his were unable to determine species status for any of the species disposal, he recognized that N. cyclotis was so distinctive from the islands. from the other Nyctimene species to warrant a separate group, Poor taxonomic resolution and lack of information on the cyclotis group. He defined the cyclotis group as having 1 status were deemed to be the most important threats to this reduced M and M1, broadly rounded ears and mottled dorsal subfamily by the IUCN/SSC Chiroptera group (Mickleburgh fur with darker tips to the hairs (Andersen, 1912a). Nyctimene et al., 1992). One species is currently IUCN red-listed as certans was originally described as a species on the basis of Endangered: N. rabori Heaney & Peterson, 1984 (Ong et heavier dentition, darker fur and greater size than N. cyclotis al., 2008); two species as Vulnerable: N. minutus Andersen, (Andersen, 1912b). The derivation of its name from the 1910 (Hutson et al., 2008a) and N. keasti (Helgen & Hutson, Latin to mean indisputable or reliable, perhaps refers to its 2008); and five species as Data Deficient: N. cyclotis supposed distinctiveness as a species (Flannery, 1995b). Yet, (Huston et al., 2008b), N. draconilla (Huston et al., 2008c), the taxonomic status of N. certans has been anything but N. malaitensis (Hamilton, 2008), N. masalai (Helgen & certain in the last 50 years. In his description of N. certans, Bonaccorso, 2008) and N. sanctacrucis Troughton, 1931 Andersen (1912b) and later in an appendix to his Catalogue (Leary et al., 2008). The IUCN/SSC Chiroptera group of Chiroptera (Andersen, 1912a), emphasized that N. certans recommended surveys of all species of tube-nosed fruit was an eastern representative of N. cyclotis. He pointed out bats throughout their range and proposed several specific that both the molariform teeth and ears were as in N. cyclotis. research projects in Papua New Guinea (Mickleburgh et al., Andersen did not formally place N. certans in the cyclotis 1992). However, more recent assessments have reduced the group but its close relationship to
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