Crocidura nigripes Miller and Hollister 1921
Creators: name: Esselstyn, Jacob A.
name: Achmadi, Anang S.
name: Handika, Heru
name: Swanson, Mark T.
name: Giarla, Thomas C.
name: Rowe, Kevin C.
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Soricomorpha
Soricidae
Crocidura
Crocidura nigripes

description: <i>Crocidura nigripes</i> Miller and Hollister, 1921 <i>Crocidura nigripes</i> Miller and Hollister, 1921: 101. Original description. <i>Crocidura nigripes lipara</i> Miller and Hollister, 1921: 101. Original subspecies description. HOLOTYPE: USNM 217545, an adult male collected by H.C. Raven on 4 August 1916 and prepared as a skin and skull. External measurements from the holotype are 131 mm × 51 mm × 14 mm; ear length and weight were not recorded. TYPE LOCALITY: “Temboan (southwest from Tondano Lake), northeastern Celebes ” (Miller and Hollister, 1921: 101; fig. 1). We estimate Temboan is in the Southeast Minahasa Regency of North Sulawesi Province, 6 km south of Kalait, at 0.979° N, 124.605° E, 650 m. See the gazetteer (appendix) for a full justification of our interpretation of Raven’s notes. GEOGRAPHIC DISTRIBUTION: <i>Crocidura nigripes</i> is generally regarded as a Sulawesi endemic that occurs broadly across lowland areas of the island (fig. 1), but the species was tenta- tively reported from Obi Island by Fabre et al. (2018). This latter record is based solely on the similarity of cytochrome <i>b</i> sequences between <i>C. nigripes</i> and a single specimen from Obi; we have not examined the Obi specimen. On Sulawesi, we recorded this species from the north-east (Mt. Ambang, Temboan, and Lembeh, North Sulawesi Province), north-central (Toraut, North Sulawesi Province), north-west (Mt. Buliohuto, Gorontalo Province and Mt. Dako, Central Sulawesi Province), west-central (Mts. Balease, Rorekatimbo, and Torompupu, Toare, Tolai, and Sungai Miu, Central Sulawesi Province; Mt. Gandang Dewata (including our lowland sample area at Salu Tiwo), West Sulawesi Province), and east-central areas of endemism (Mt. Katopasa and Peleng Island, Central Sulawesi Province). Notably, we did not record the species from the south-west or southeast areas of endemism. Similarly, Musser (1987) reported the species only from the west-central area of endemism and northern peninsula. The full elevational extent of <i>C. nigripes</i> records across these areas extends from near sea level to just over 2000 m (fig. 13; table 3). EMENDED DIAGNOSIS: A medium-sized shrew (tables 2, 14) with a medium build and uniformly dark pelage, feet, and tail (light chocolate to black; fig. 36). The tail is shorter than head-andbody length. The hind foot is somewhat short relative to head-and-body length (fig. 17), and usually dark brown to black. The skull is broad, and quite angular, with a prominent maxillary process, broad palate, and dentition with a large occlusal area relative to palatal area (fig. 37B). The interorbital region is unusually narrow, which enhances the prominence of the braincase breadth and maxillary process (fig. 10). Rostral length makes up a large proportion of skull length (fig. 10). COMPARISONS: <i>Crocidura nigripes</i> is generally very dark in color with a short tail and moderate body size, but there is substantial variation in size TABLE 12 a The sample mean ± one standard deviation, the observed range in parentheses, and the sample size. and color. The feet are often nearly black, which will, in most cases, distinguish this species from all Sulawesi shrews except the darkest members of the Small-Bodied Group, which are much smaller. Typically, <i>C. nigripes</i> is smaller than <i>C. elongata</i>, <i>C. quasielongata</i>, <i>C. caudicrassa</i>, and <i>C. rhoditis</i>, comparable in size to <i>C. microelongata</i>, <i>C. pseudorhoditis</i>, <i>C. brevicauda</i>, and <i>C. australis</i>, and larger than the remaining species known from Sulawesi. It is the largest species in the Ordinary Group. These size differences can be observed using measures of skull length, head-and-body length, mass, or hind foot length (figs. 9, 17). Tail length is shorter than head-and-body length, which will distinguish <i>C. nigripes</i> from all similarly sized shrews (as judged by HBL, CIL, etc.) except the two Thick-Tailed species, which have greater relative hind-foot lengths (HF/HBL) and stockier bodies (figs. 9, 17). The skull of <i>C. nigripes</i> has a more angular braincase, more prominent maxillary process (enhanced by the narrow interorbital region), and more robust dentition (larger occlusal surface area) than all other Sulawesi shrews. The relative interorbital width (IOW /CIL) is substantially less than in all other species except <i>C. elongata</i> and <i>C. quasielongata</i> (fig. 10), two species with much longer skulls, tails, hind feet, and bodies. COMMENTS: Phylogenetically, <i>Crocidura nigripes</i> is more closely related to shrew species endemic to Sundaland than to any others that live on Sulawesi (Ruedi et al., 1998; Esselstyn et al., 2009, 2019; Hinckley et al. 2021). Our phylogenetic analyses conducted on far more species than previous studies reinforce this conclusion. We placed <i>C. nigripes</i> as sister to <i>C. palawanensis</i>, a species endemic to the Palawan Island group in the southwestern Philippines (figs. 7, 8). Hinckley et al. (2021) estimated <i>C. nigripes</i> as a close relative of <i>C. foetida</i>, a Bornean endemic that was not included in our analyses. Miller and Hollister (1921) described two subspecies, <i>Crocidura nigripes nigripes</i> from Temboan and <i>C. n. lipara</i> from Gimpoe, in Central Sulawesi Province. Ruedi (1995) treated northern peninsula populations as <i>C. n. nigripes</i> and specimens from other parts of the island as <i>C. n. lipara</i>. He noted size differences with this geographic division, but also highlighted the limited sample of available specimens. We treat <i>C</i>. <i>n</i>. <i>lipara</i> as a synonym of <i>C</i>. <i>nigripes</i> because we have not identified any geographically partitioned morphological (fig. 38; tables 15, 16) or genetic diversity with our improved sampling. Jukes-Cantor cytochrome <i>b</i> distances between specimens sampled from across Sulawesi are &lt;0.01 (fig. 4; supplementary data S 3). As such, we see no reason to maintain the subspecific distinction between <i>C. n. nigripes</i> and <i>C</i>. <i>n</i>. <i>lipara</i>. SPECIMENS EXAMINED: Mt. Ambang (LSUMZ 39044, 39279, 39320), Mt. Balease (MZB 38491; FMNH 210593, 210595, 210611), Mt. Buliohuto (LSUMZ 38279–38281, 38283, 38286, 38287, 38292–38295, 38297; NMV C37778), Mt. Dako (LSUMZ 37017–37019, 37022, 37025–37029, 37032, 37035–37049; NMV C37257, C37265, C37307, C37318, C37368, C37369), Mt. Gandang Dewata (FMNH 218708), Gimpoe (= Gimpu) (USNM 219444, 219446), Lake Lindu (USNM 218668, 501223), Mt. Katopasa (LSUMZ 40925; NMV C40227, C40229, C40230), Koelawi (= Kulawi) (FMNH 47361), Kuala Navusu (AMNH 226105, 226106, 226108, 226109, 226110, 226112), Peleng Island (AMNH 109215), Pinedapa (USNM 219435), Mt. Rorekatimbo (FMNH 213190, 213191, 213252), Salu Tiwo (FMNH 218703, 218704, MZB 38488, 38489), Toraut (RMNH 38389 [= IZEA 4413], 38390 [= IZEA 4415]), Lembeh (FMNH 31844), Sungai Miu (AMNH 223989), Sungai Tolewonu, Tolai (AMNH 226537–226539), Temboan (FMNH 43859; USNM 217536, 217541– 217545), Toare (= Tuare or Toware) (FMNH 47362, USNM 219434, 219437–219440, 219443, 219447), Mt. Torompupu (LSUMZ 39469–39474).
description: Published as part of <i>Esselstyn, Jacob A., Achmadi, Anang S., Handika, Heru, Swanson, Mark T., Giarla, Thomas C. &amp; Rowe, Kevin C., 2021, Fourteen New, Endemic Species Of Shrew (Genus Crocidura) From Sulawesi Reveal A Spectacular Island Radiation, pp. 1-109 in Bulletin of the American Museum of Natural History 2021 (454)</i> on pages 71-75, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835
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