Fourteen New, Endemic Species Of Shrew (Genus Crocidura) From Sulawesi Reveal A Spectacular Island Radiation Author Esselstyn, Jacob A. Author Achmadi, Anang S. Author Handika, Heru Author Swanson, Mark T. Author Giarla, Thomas C. Author Rowe, Kevin C. text Bulletin of the American Museum of Natural History 2021 2021-12-15 2021 454 1 109 https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-454/issue-1/0003-0090.454.1.1/Fourteen-New-Endemic-Species-of-Shrew-Genus-Crocidura-from-Sulawesi/10.1206/0003-0090.454.1.1.full journal article 2886 10.1206/0003-0090.454.1.1 3a7cdaa4-c2f2-4370-9301-c2ab339b2e0d 0003-0090 5788835 7982B923-4CDC-44ED-A598-8651009DC7CC Crocidura musseri Ruedi, 1995 Crocidura musseri Ruedi, 1995: 254 . Original description. HOLOTYPE : MZB 16791 (= IZEA 4397 ), an adult female caught 18 August 1991 . Measurements from the holotype are TC (head-andbody length): 66 mm × Q (tail length): 57 mm × PP (hind-foot length): 13.5 mm = Pds (weight): 6.5 g . TYPE LOCALITY: Mt. Rorekatimbo, Central Sulawesi , at 1° 16’ S , 120° 15’ E , 2230 m elevation. GEOGRAPHIC DISTRIBUTION: Apparently restricted to high elevation areas (> 1800 m ) in the northern portions of the west-central area of endemism east of the Palu-Koro Fault in Central Sulawesi Province ( figs. 13 , 39 ; table 3 ). Ruedi (1995) reported specimens of this species from only the type locality on Mt. Rorekatimbo. We collected specimens that match Ruedi’s description and are mitochondrially identical (based on cytochrome b sequences from Ruedi et al., 1998 ) from the same mountain, at three nearby sites ( 1.3093° S , 120.3092 °E , 2020 m ; 1.2926° S , 120.3064° E , 2180 m ; 1.2884° S , 120.3104° E , 2250 m ). Musser collected additional specimens from 1800–2300 m elevation on the neighboring Mt. Nokilalaki (ca. 15 km W of Mt. Rorekatimbo). EMENDED DIAGNOSIS: A medium-sized shrew ( tables 2 , 14 ) with a thick, dark brown pelage that is only slightly paler on the venter. Foot, ear, tail, and rhinarium color closely match the pelage. Tail TABLE 14 Descriptive Statistics a for Craniodental Measurements (mm) for Species of the Ordinary Group of Sulawesi Crocidura

C. musseri	C. nigripes	C. normalis	C. ordinaria	C. solita
Condyloincisive length	20.13 ± 0.446 (19.17–20.78) 17	22.47 ± 0.733 (20.91–24.32) 80	18.73 ± 0.276 (18.29–19.66) 27	20.38 ± 0.491 (19.47–21.59) 15	20.13 ± 0.584 (19.25–21.55) 43
Braincase breadth	9.74 ± 0.233 (9.42–10.26) 18	9.94 ± 0.315 (9.32–10.78) 77	8.61 ± 0.168 (8.45–9.06) 27	9.79 ± 0.178 (9.48–10.06) 15	9.51 ± 0.263 (8.8–10.12) 43
Interorbital width	4.91 ± 0.164 (4.61–5.2) 18	4.71 ± 0.196 (4.3–5.08) 85	4.3 ± 0.114 (4.11–4.58) 28	4.86 ± 0.137 (4.65–5.12) 16	4.68 ± 0.167 (4.36–5.06) 43
Rostral length	8.16 ± 0.205 (7.76–8.39) 17	9.31 ± 0.337 (8.45–10.09) 86	7.36 ± 0.163 (7.09–7.75) 28	8.25 ± 0.285 (7.77–8.95) 16	8.15 ± 0.363 (7.2–9.17) 43
Postpalatal width	3.84 ± 0.12 (3.59–4.03) 18	4.02 ± 0.156 (3.7–4.38) 85	3.34 ± 0.106 (3.15–3.68) 28	3.96 ± 0.18 (3.7–4.36) 16	3.88 ± 0.147 (3.57–4.22) 43
Rostral width	2.65 ± 0.147 (2.46–3.03) 17	3.02 ± 0.17 (2.62–3.58) 86	2.47 ± 0.172 (2.08–2.71) 28	2.96 ± 0.139 (2.73–3.18) 16	2.73 ± 0.198 (2.31–3.2) 43
Postpalatal length	8.89 ± 0.222 (8.4–9.23) 18	9.79 ± 0.366 (8.9–10.54) 80	8.41 ± 0.231 (8.08–9.03) 27	9.1 ± 0.297 (8.7–9.87) 15	8.89 ± 0.304 (8.41–9.51) 44
Condyle to gle- noid fossa	7.93 ± 0.186 (7.52–8.24) 18	8.37 ± 0.281 (7.78–8.9) 80	7.72 ± 0.134 (7.47–8.01) 27	8.09 ± 0.194 (7.77–8.43) 15	7.93 ± 0.224 (7.35–8.49) 43
Upper toothrow length	8.84 ± 0.229 (8.35–9.19) 17	10.2 ± 0.359 (9.24–11.02) 86	8.07 ± 0.211 (7.69–8.7) 28	9.04 ± 0.254 (8.63–9.68) 16	8.91 ± 0.327 (8.07–9.89) 43
P4 to M3 length	5.08 ± 0.182 (4.7–5.34) 18	5.82 ± 0.238 (5.17–6.24) 86	4.49 ± 0.163 (4.23–5.02) 28	5.24 ± 0.167 (4.99–5.62) 16	5.11 ± 0.195 (4.72–5.67) 43
M2 to M2 labial width	5.86 ± 0.238 (5.44–6.16) 17	6.84 ± 0.253 (6.37–7.52) 86	5.12 ± 0.152 (4.85–5.63) 28	6.08 ± 0.286 (5.69–6.77) 16	5.9 ± 0.217 (5.53–6.61) 43
Palatal width	2.52 ± 0.085 (2.36–2.64) 18	2.74 ± 0.134 (2.45–3.19) 85	2.25 ± 0.096 (2.01–2.43) 28	2.49 ± 0.118 (2.31–2.8) 16	2.46 ± 0.105 (2.24–2.71) 43

a The sample mean ± one standard deviation, the observed range in parentheses, and the sample size. length is shorter than head-and-body length. The tail has inconspicuous applied hairs that are visible with the naked eye only on the distal third of tail length and sparse bristle hairs spread from the tail base to at least its midpoint ( fig. 36A ). The braincase is notably inflated, with great width, giving an overall wedge-shaped appearance when viewed from the dorsal aspect ( fig. 37A ). The squamosal-parietal suture is often open, leaving a tear-drop-shaped opening anteriorly, directly below the anterior opening of the sinus canal (sensu McDowell, 1958 ). COMPARISONS: Crocidura musseri is smaller than members of the Elongata Subgroup and all Rhoditis Group members except C. pallida . It is larger than all members of the Small-Bodied Group. Within the Ordinary Group, C. musseri is smaller than C. nigripes , slightly larger than the sympatric C. normalis , and comparable to C. ordinaria and C. solita . Relative to head-and-body length, C. musseri has a shorter tail than all members of the Long-Tailed and Rhoditis groups, C. nigripes , and the sympatric C. solita . The pelage of C. musseri is thicker than that of any other shrew species we recognize from Sulawesi , except C. caudicrassa . Overall, the color of C. musseri is substantially darker than specimens from the Long-Tailed and Rhoditis groups, C. lea , and C. mediocris . Within the Ordinary Group, C. musseri is slightly darker than C. solita and C. normalis . Relative to skull length, skull width at both the braincase and interorbital region is greater than most other spe- cies, including all members of the Ordinary Group ( figs. 10 , 37 ). The unusual width of the skull ( table 14 ) gives C. musseri a more wedge-shaped appearance than any other species when viewed from the dorsal aspect ( fig. 37A ). Crocidura nigripes has a more angular braincase, a more robust dentition, and a much narrower relative interorbital region (IOW/CIL). Relative rostral length (RL/CIL) in C. musseri is less than in C. nigripes , but greater than in C. normalis ( fig. 10 ). Crocidura musseri is distinguished from all sympatric shrews by a combination of its short tail length, thick and dark pelage, and wide, wedge-shaped skull with small teeth ( figs. 9 , 36 , 40 ). FIG. 36. Images showing the ventral surface of the right hind foot and dorsal surface of the tail base (approximately 1 cm from rump) and tail tip from two darkly pigmented members of the Ordinary Group: A, Crocidura musseri , FMNH 213267; and B, C. nigripes , LSUMZ 37018. Scale bars apply to all images in panel A and to foot (upper) and tail (lower) in panel B. COMMENTS: Our description of Crocidura musseri is based primarily on the series of specimens we collected in 2011 very near the type locality. This series matches the holotype (MZB 16791) and the description of Ruedi (1995) . Cytochrome b sequences from our series are identical to one from a paratype (IZEA 4403; Ruedi et al., 1998 ). None of our phylogenetic analyses supported a close relationship of C. musseri to any other species. Our mitogenome result placed C. musseri as sister to all other members of Sulawesi’s endemic radiation with some support ( fig. 5 ), while our UCE species tree placed it as sister to C. microelongata and C. pallida , but without statistical support ( fig. 7 ). SPECIMENS EXAMINED : Mt. Nokilalaki ( AMNH 223495 , 225516 , 225517 , 225521 , 225525–225527 , 225530 , 225531 ), Mt. Rorekatimbo ( FMNH 213249–213251 , 213253–213256 , 213258–213269 , 213442 , 213443 ; MZB 16791 ).