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Species Limits and Distribution of the Malagasy Carnivoran Genus Eupleres (Family Eupleridae)

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Species Limits and Distribution of the Malagasy Carnivoran Genus Eupleres (Family Eupleridae) Article inAuthor's press - uncorrected Copy proof Mammalia 74 (2010): 177–185 ᮊ 2010 by Walter de Gruyter • Berlin • New York. DOI 10.1515/MAMM.2010.018 Species limits and distribution of the Malagasy carnivoran genus Eupleres (Family Eupleridae) Steven M. Goodman1,2,* and Kristofer M. Helgen3 Eupleridae are becoming reasonably well established (Yoder 1 Field Museum of Natural History, 1400 South Lake Shore et al. 2003), the distribution and species limits of several Drive, Chicago, IL 60605, USA genera remain poorly studied. Herein we address these 2 Vahatra, BP 3972, Antananarivo 101, Madagascar, aspects for the enigmatic genus Eupleres (the falanouc), e-mail: [email protected]; which possesses a highly reduced dentition and other [email protected] remarkable anatomical features (Gray 1870, Carlsson 1902, 3 National Museum of Natural History, Smithsonian Pocock 1915, Scapino 1981, Popowics 2003, Gaubert et al. Institution, P.O. Box 37012, MRC 108, Washington, 2005) and for which little information is available regarding DC 20013-7012, USA natural history and geographical distribution (Albignac 1972, 1973a, 1974). *Corresponding author In his extensive review of the Carnivora of Madagascar, Albignac (1973a) considered Eupleres to be monospecific and represented by two subspecies: E. g. goudotii Doye`re, Abstract 1835, from the eastern wet forests and E. g. major Lavauden, 1929, from the transitional dry deciduous-humid forests of A review was conducted of members of the endemic Mala- the northwest. Albignac (1973a) presented several external gasy carnivoran genus Eupleres (family Eupleridae) based morphological characters to separate these two subspecies. on published and unpublished records and museum speci- In subsequent taxonomic reviews of the Carnivora of mens. Classically, one species has been recognized in this Madagascar, this monospecific arrangement for Eupleres has poorly known genus – E. goudotii, divided into two geo- been followed (Wozencraft 1993, 2005), although there has graphical forms with non-overlapping distributions: E. g. been some indication that these geographical forms could goudotii distributed in the mesic forests of the east and E. g. warrant specific recognition (Lavauden 1929, Albignac major found in the dry areas of the northwest. Drawing on 1973b). The purpose of this paper is to review morphological external and craniodental comparisons, we demonstrate that Author's Copy characters and measurements distinguishing these two forms, Author's Copy these two forms are highly distinctive morphologically and their geographical distribution, and to assess their specific can be readily distinguished from each another. Furthermore, status. there is some evidence that they both can occur on the slopes of Montagne d’Ambre in the far north of the island. On this basis, we recognize these taxa as distinct species, E. goudotii Methods and materials and E. major. Patterns of morphological variation Keywords: distribution; Eupleres; Madagascar; morphology; taxonomy. To assess patterns of morphological variation in the different recognized forms of Eupleres we examined and measured specimens in the following institutions: American Museum Introduction of Natural History (AMNH); Muse´um national d’Histoire naturelle (MNHN); Museum of Comparative Zoology, Har- Until recently it was assumed that the native Carnivora of vard University (MCZ); Natural History Museum, London Madagascar represented several deeply divergent lineages wBMNH; formerly The British Museum (Natural History)x; within the order, and hence that they arrived on the island Yale Peabody Museum (YPM), Yale University; Museum fu¨r via multiple colonization events (Gregory and Hellman 1939, Naturkunde, Humboldt Universita¨t, Berlin (ZMB). Some of Simpson 1945). However, recent molecular comparisons these specimens are incomplete, missing teeth or damaged have shown that all native Malagasy Carnivora genera in other ways. All craniodental measurements included here- (Cryptoprocta, Fossa, Eupleres, Galidia, Galidictis, Sala- in were taken by the second author, at an accuracy of 0.1 mm noia, and Mungotictis) form a monophyletic group repre- for cranial variables and 0.01 mm for dental variables. senting a single colonization event (Yoder et al. 2003) and The following measurements were taken for all adult (and all are now placed in an endemic family, Eupleridae (Wozen- nearly mature subadult) specimens – skulls: greatest length craft 2005). This family represents one of the more extra- of skull (GSL), condylobasal length (CBL), zygomatic ordinary adaptive radiations among the world’s Carnivora breadth (ZB), breadth of braincase at zygomatic roots (BB), (Goodman 2009). Although higher level systematics of the interorbital width (IOW), length of anterior palatal foramina 2010/201004 Author's Copy Article inAuthor's press - uncorrected Copy proof 178 S.M. Goodman and K.M. Helgen: Species limits of Eupleres (Eupleridae) (APFL), maximum length of nasal bones (NL), palatal length inflated structures, the fused interdigital pads form a large (PL), auditory bulla length (AUDL), auditory bulla width proportion of the foot-pad surface, and the metatarsal pads (AUDW), and mandible length (ML); teeth – length of upper (thenar and hypothenar) are more developed. Finally, a con- first molar (LM1); width of upper first molar (WM1); length siderable portion of the plantar surface posterior to the of upper second molar (LM2); width of upper second molar metatarsal pads is naked in major and largely covered with (WM2); external alveolar distance between upper first fur in goudotii. molars (M1–M1); external alveolar distance between upper canines (C–C); length of maxillary toothrow (C–M2); width Cranial measurements and morphology There are of lower first molar (wm1); length of second lower molar several cranial and mandibular measurements that distinguish (lm2); length of third lower premolar (lp3). goudotii and major (Table 2, Figure 1). In general, major has a distinctly more robust skull, both in cranial length (e.g., greatest length of skull) and breadth (e.g., zygomatic Results breadth), as well as a longer and more robust mandible. In fact, there is a large discrepancy, with no overlap, in condy- Morphology lobasal length between adults of goudotii (F87 mm) and major (G90 mm), offering a convenient single metric for External morphology External measurements tabulated discriminating adult skulls. Furthermore, major has a dis- both by Albignac (1973a) from non-cited sources and from tinctly longer palate, but shorter anterior palatal foramina museum study skins at MCZ (Table 1) demonstrate the con- than those of goudotii, an important cranioproportional dis- sistently larger body size of major relative to goudotii. tinction. One of the most noticeable diagnostic differences Another striking distinction between major and goudotii is between these two animals is the much more expansive audi- in pelage coloration; major is distinctly darker brown than tory bullae in major, as reflected both in average values for goudotii, which is paler and more fawn brown in overall length and width dimensions (tabulated in Table 2), but most coloration (both species are paler ventrally than dorsally). notably expressed by inflation of the bullae in the vertical Albignac (1973a) noted sexual dimorphism in both of these (dorsoventral) dimension (difficult to measure consistently forms, with males being slightly larger than females, but and not tabulated here). based on current limited sample sizes we are unable to test for these differences in a meaningful way. Dental measurements Many of the specimens available Another consistent difference between these two forms is for this study are missing teeth, so that there are numerous in the position and size of the foot-pads and fur cover on the dental variables for which few measurements are available, distal portions of the fore and hind feet (see Albignac 1973a, particularly for major. In general, the upper molar teeth in Figures 9–12). On the palmar surface of the forelimb of major are more robust in length (e.g., length first upper nominate goudotii the digital pads are only moderately Author's Copy molar, length second upper molar) and width (e.g., width first Author's Copy developed, and the interdigital and metacarpal pads are large- upper molar, width second upper molar) than in goudotii. ly fused and form a large central surface, whereas in major The length of the maxillary toothrow in major is notably the digital pads are markedly larger and inflated, and the longer than in goudotii. Furthermore, the distance across the hypothenar and thenar pads are notably separated from the rostrum (e.g., distance outer C–C) and palate (distance outer inflated interdigital pads (terminology follows Brown and M1–M1) are greater in major compared with goudotii. The Yalden 1973). Furthermore, in major the palmar surface pos- lower molar measurements in goudotii show little overlap in terior to the metacarpal pads is largely naked and furred in length and width to those measurements of major (e.g., sec- goudotii. On the plantar surface of goudotii, the digital pads ond to last lower width, second to last lower width, length are reduced, and the largely fused interdigital pads form a of fourth to last tooth in lower jaw) (Table 3). Hence, animals central rounded unit with a notable separation from the dig- assigned to nominate goudotii and major have conspicuously ital pads, whereas in major the digital pads are conspicuously different
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