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Systematic Review of Nyctimene Cephalotes and N

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Systematic Review of Nyctimene Cephalotes and N Records afthe Westem Australian Museum 17: 125""142 (1995). Systematic review of Nyctimene cepJzalotes and N. albiventer (Chiroptera: Pteropodidae) in the Maluku and Sulawesi regions, Indonesia l l 2 D.J. Kitchener , W.e. Packer and A. Suyanto 1 Western Australian Museum, Francis Street, Perth, Western Australia 6000 2 Museum Zoologicum Bogoriense, L1PI, Jalan Ir. H. Juanda 9, Bogor, Indonesia, 16122 Abstract -A univariate and multivariate statistical study of the morphology of island populations of Nyctimene in the Maluku region, Indonesia, distinguishes Nyctimene keasti Kitchener, 1993, from the Tanimbar and Kai Islands, as a species. The form from the Tanimbar islands is described as a new subspecies of N. keasti; it is not associated with N. cepha/otes as previously considered in lilt. Nyctimene cepha/otes is restricted to Sulawesi and the Maluku region. The Sulawesi population of N. cepha/otes is described as a distinct new subspecies. The Aru population of N. a/biventer is somewhat differentiated from the Papua New Guinea population of N. a. papuanus; its subspecific status is not determined. Close island populations of N. c. cepha/otes, N. k. keasti and N. k. subsp. novo show considerable morphological differentiation. INTRODUCTION terrestrial vertebrate fauna of South Sulawesi, Kitchener et al. (1993) examined morphological Maluku and Nusa Tenggara, Indonesia. On these variation among populations of Nyctimene expeditions, series of Nyctimene that would be albiventer (Gray, 1863) in the Maluku region. They attributed to N. cephalotes by earlier authors, were showed that the population on Am was collected from the following islands: Seram; morphologically similar to the Papua New Guinea Ambon; Bum; Yamdena and Selam, Tanimbar N. a. papuanus K. Andersen, 1910; while those on group; and South Sulawesi. This paper reports on the Kai and Banda Neira Islands (N. a. keasti an examination of these extensive modern Kitchener, 1993) were distinct from both the Am collections which allow both a re-appraisal of form and the nominate subspecies on Halmahera morphological variation among island populations and Ternate Islands. of N. cephalotes and N. albiventer and a The discovery of N. a. keasti confuses somewhat reassessment of their taxonomy in the study the previous diagnostic distinction between N. region. albiventer and the closely allied N. cephalotes (pallas, 1767). This distinction had been based on the larger size of N. cephalotes and the fact that the females had a dorsum that was much paler than the males, MATERIALS AND METHODS whereas in N. albiventer there was no such colour A total of 142 adult specimens (listed in distinction between the sexes (Anderson 1912). A specimens examined section) was examined. These number of Nyctimene a. keasti specimens had were from Banda Neira Neira I. (2); Dullah I., Kai forearm lengths overlapping those of both N. a. Kecil (7); Kai Besar I. (5); Wokam I., Am islands albiventer and N. cephalotes. Further, female N. a. (27); the Halmahera group (6); Papua New Guinea keasti were much paler dorsally than the males. (13); Ambon (10); Seram (16); Bum (2); Sulawesi In the study region, N. cephalotes is reported from (9); Selaru (13) and Yamdena (32). The localities of the following islands: Ambon (fixed as the type these specimens are shown in Figure 1. Apart from locality by Andersen 1912); Seram; Bum, Larat the Halmahera group specimens (Australian Island (Tanimbar group); Timor and Sulawesi Museum, Sydney), all specimens are currently (Andersen 1912; Hill in Corbet and Hill 1992). The lodged in the Western Australian Museum. records from Larat Island and Timor Island appear Seventeen measurements of skull, dentary and to devolve on single specimens collected late last dental characters and six of external body century (Andersen 1912, Goodwin 1979). characters (all in mm) were recorded from adult Between 1987 and 1993 expeditions by staff from specimens. the Western Australian Museum and Museum The measurements recorded were (all Zoologicum Bogoriense carried out a survey of the measurement involving teeth were to alveoli): GSL, 126 D.}. Kitchener, W.e. Packer, A. Suyanto • o ~_J,~ 140 •~. ~ + + ·t·- ... ~~'~'O' ~ J - ~ G·t· .. \ t, ~. , D' •• Jo~l ~ .' ....~ .~,~ ~•X• o ..... " .. & . '.~. '\. P .. • ~ A ~!J~/7 A c:. .. ~.,f/ ·\:)6 + + Figure 1 Locality of Nyctimene in this study; N. albiventer albiventer, .; N. a. subsp. indet., G; N. a. papuanus, A.; N. keasti keasti, 0; N. k. tozeri subsp. nov.,~; N. cephalotes cephalotes, .; N. c. aplini subsp. nov., X. greatest skull length; CBL, condylobasal length; Adults were diagnosed as those specimens with PIF, minimum length from posterior margin of the following sutures fused: basioccipital ­ incisive foramen to palate posterior margin; RL, basisphenoid, basisphenoid - presphenoid and rostrum length, from anteriomost internal margin palatine - maxillary (these sutures are illustrated in of orbit to nares; RH, rostrum height, from upper Heaney and Peterson 1984: Figure 4). canine alveoli to level of dorsal surface of nasals; The effect of sex and island on all characters was BB, braincase breadth above zygoma; ZW, examined by multiple regressions for the islands: zygomatic width; MlMl and ClCl, width across Yamdena, Selaru, Kai Kecil, Kai Besar, Aru, MIMl and ClCl respectively, from the labial side; Ambon, Seram, Sulawesi and Papua New Guinea. ClMl and C M upper and lower canine to last l 2 Examination of the residuals from regression molar length; lOB, minimum interorbital breadth; analyses gave no indication of heteroscedasticity. POB, minimum postorbital breadth; MFB, Stepwise canonical variate (discriminant mesopterygoid fossa breadth, at the widest point function) analyses (OFA) were run for skull, of the palatal flange; p4p4, palatal width between dentary and dental characters and external the lingual aspect of p4p4; ML, mandible length, characters using all 23 characters for males and from condyle to anteriomost point of dentary; CH, females combined, after first testing for sexual dentary coronoid height; TV, tail to anus length; dimorphism. A reduced set of five of these EAR, basal notch to apex length; FA, forearm characters was used in all presented OFA because length; and MC3-5, metacarpal 3,4 and 5 length. in all instances they provided similar OF plots to The skull, dentary and dental characters were the full set of 23 characters. This reduced set of measured to an accuracy of 0.01 mm, while the characters was selected in all these analyses external body characters were measured to 0.1 because the sample size of the smallest a priori mm. Terminology used in the description of skull, group selected was always less than the total dentary, dental and external body characters number of characters. This reduced set of follows Hill and Smith (1984). Pelage descriptions characters was selected because they provided follow the colour terminology of Smithe (1975). values that minimise Wilk's Lambda. Systematic review of Nyctimene ceplla/otes and N. a/biventer 127 STATISTICS: RESULTS AND DISCUSSION Univariate statistics Mean, standard deviation, mInimUm and maximum values and sample size for each island are presented in Table 1, this paper, and Kitchener \ et al. (1993: Table 1) for all characters examined. \ \ • Multiple regressions ! • o Multiple regressions were run for skull, dentary, ! o I dental and external characters on sex and nine f o. island populations. The islands of Buru, Banda f Neira, Halmahera, Ternate and Moratai were c: excluded because of their small sample size and/ Q u or absence of either males or females (Table 2). ­c: ::> ... I o Sex Metacarpal 3-5 lengths and postorbital breadth / o -0 / were significantly influenced by sex (F = 6.11, P 00600S°"- 196 / = 0.015; F1,96 = 4.67, P = 0.033; F1,96 ='4.653, P rt0Gl. (] "0, o o ... 0 \ . \ 0.033; and F1 96 =4.639, P =0.034, respectively. o \ o' . \ o Island . \• 'V' Only postorbital breadth was not highly «.01) / significantly related to island. Clearly there was -- ./ considerable difference in morphology between these island populations. - 8 +-~,--~,....--r~-r~-,--~-,-~,--~,....--r~- - 5 -4 Interactions Function 2 There were no significant interactions. Figure 2 Canonical variate analysis on the following island populations: Aru, [;] ; Kai Besar, 0; Kai Multivariate analyses Kecil, 0; the Halmahera group (Halmahera, Because the multiple regression analysis Ternate and Morotai), -; Papua New Guinea, involved testing a large number of interactions, ...; Selaru 0; and Yamdena, II (Tanimbar some of these tests will be significant at group); Seram, +; Ambon, e; and Sulawesi, 0.05>P>0.01 by chance alone. For this reason, none X; Banda Beira +, and Buru *, were ungrouped. The DFA plots of functions 1 and of the characters measured were considered to be 2 were based on a selection of five characters markedly influenced by sex. As a result both males (skull, dentary, and external body), with and females are combined for all 23 characters males and females combined. examined in the following DFA. the Ambon group (Ambon-Seram-Buru-Sulawesi); All populations the Kai group (Kai and Tanimbar islands); and the The DFA was first run using all 23 skull, dental, Halmahera group (Halmahera, Aru and Papua dentary and external body characters and the New Guinea). When a DFA was again run using islands Aru, Kai Besar, Kai Kecil, the Halmahera the reduced set of five characters and these three group (Halmahera, Temate and Morotai) Papua island groups, the analysis extracted two New Guinea, Selaru and Yamdena (Tanimbar significant functions (Figure 3). Function 1, which Islands), Seram, Ambon and Sulawesi. Banda explained 78.9% of the variance, separated the Neira and Buru were ungrouped because there Ambon and Kai groups from the Halmahera were only two individuals from each of these group. The character loading heavily on the islands. The above analysis was then repeated standardised canonical variate coefficients (>0.6) using only five characters (forearm length, FA; ear on this function was C j M 2 length (Table 3). 1 length EAR; C M length, C M ; width across M1M 1 2 1 2 Function 2, which separated the Ambon group from labial side, MIM1; and rostraI length, RL).
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