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Crocidura elongata Miller and Hollister, 1921 Crocidura elongata Miller and Hollister, 1921: 101. Original description. . Crocidura ���dark elongata��� Esselstyn et al., 2019: 1715. Informal name. HOLOTYPE: USNM 217534, an adult male obtained by H.C. Raven on 1 August 1916. The specimen comprises a skin and skull. External measurements recorded from the type are 214 mm �� 120 mm �� 22 mm; no ear length or weight was recorded. TYPE LOCALITY: ���Temboan (southwest from Tondano Lake), northeastern Celebes��� (Miller and Hollister, 1921: 101; fig. 1). We estimate that the type locality is at 0.979�� N, 124.605�� E, 650 m elevation, which differs from other interpretations (e.g., Musser, 2014). See the gazetteer for a full explanation (appendix). GEOGRAPHIC DISTRIBUTION: Apparently restricted to the northern peninsula of Sulawesi, where clear records are known from the northeast (the type locality, Temboan and Mt. Ambang, North Sulawesi Province) and north-west (Mt. Dako, Central Sulawesi Province, and Mt. Buliohuto, Gorontalo Province) areas of endemism (fig. 16). The absence of records from the north-central area of endemism are almost certainly due to the general lack of mammal collecting from this region. Miller and Hollister���s (1921) paratypes from ���Pinedapa, eastern Middle Celebes ��� and the two specimens from Mt. Rorekatimbo reported by Ruedi (1995) almost certainly represent Crocidura microelongata (see below). Crocidura elongata is known from a moderately broad range of elevations, with the low elevation records between 500 and 600 m from Mts. Buliohuto and Dako and high-elevation records reaching 1600 m on Mt. Dako (table 3). The type locality at Temboan is around 650 m elevation. TABLE 2 Descriptive Statistics a for External Measurements (mm) and Mass (g) for Species of Sulawesi Crocidura a The sample mean �� one standard deviation, the observed range in parentheses, and the sample size. Long-Tailed Ordinary Rhoditis Small-Bodied Thick ��� ������ ��� ��� ��� ������ ��� 54 43 34 ��� 40 17 78 27 15 43 5 27 75 20 7 21 20 26 27 6 5 11 = = = = = = = = = N = = = = N = = = = = N = N = = N N N N N N N N N N N N N N N N N ��� ��� ��� ��� ��� ��� ��� ������ ��� ��� ��� ��� ��� ��� = 54 = 44 = 33 = 40 = 17 = 81 = 27 = 15 = 43 = 5 = 27 = 75 = 22 = 7 = 21 = 20 = 26 = 27 = 6 = 5 = 11 N N N N N N N N N N N N N N N N N N N N N ��� ��� ��� ��� ��� ��� ��� ��� ��� ��� ��� ������ ��� ��� ��� = 54 = 43 = 32 = 40 = 18 = 78 = 27 = 15 = 43 = 5 = 27 = 76 = 20 = 7 = 21 = 20 = 26 = 27 = 6 = 5 = 11 N N N N N N N N N N N N N N N N N N N N N ��� ��� ��� ��� ��� ��� ������ ��� 54 44 34 40 17 81 27 15 ��� 43 5 27 75 22 7 21 20 26 27 6 5 11 = = = = = = = = = N = = = = N = = = = = N = N = = N N N N N N N N N N N N N N N N N TABLE 3 Elevational Ranges (m) and Species Richness of Crocidura on Mountains of Sulawesi a a Mountains are grouped by peninsula or area of endemism. Some mountain names are abbreviated: Amb = Ambang, Bulio = Buliohuto, Gand = Gandang Dewata, Lati = Latimojong, Bal = Balease, Rore = Rorekatimbo, Torom = Torompupu, Bawa = Bawakaraeng, Kato = Katopasa, Tomp = Tompotika, and Mek = Mekongga). When museum databases recorded an elevational range for an individual specimen, we used the median of that range as the elevation for that specimen. Specimens from Salu Tiwo are grouped under Mt. Gandang Dewata. EMENDED DIAGNOSIS: A long-bodied, somewhat heavily built, moderately bicolored shrew with a very long tail and unusually long, pale hind feet (figs. 9, 14C, 17; tables 2, 5). The dorsal pelage is gray-brown overall; individual dorsal hairs are gray at the base and brown at the tip. The ventral pelage is paler, with individual hairs gray-based like those on the dorsum, except that the hair tips are silvery. The silvery appearance of the venter is most pronounced on the throat and chest area. In some specimens, however, this area has a reddish tint, due to variation in the color of the tips of some hairs. The mystacial vibrissae are dark proximally but white distally. The ears are prominent and pale. Dorsally, the feet are pale, transitioning from light brown near the wrist and ankle to nearly white on the digits. Ventrally, the feet show the same transition, but pigment is concentrated around the base of the pads (fig. 14C). The claws are translucent. The hind feet are long, even relative to head-and-body length (fig. 17). The long tail is subtly bicolored, with a brown dorsum and pale brown venter. Tiny applied hairs are present along the entire length of the tail, but they are barely visible to the naked eye (fig. 14C). However, these hairs are slightly longer and white near the tip of the tail, creating a very small white distal tuft. The long bristle hairs that are common at the base of the tail of many Crocidura are nearly absent in this species. The skull is long and slender, with a notably narrow braincase, interorbital region, and palate (figs. 10, 18A). Although the interorbital region is narrow relative to skull length, it is less constricted than the braincase. The long skull of C. elongata is attributable primarily to elongation of the postpalatal region; the rostral length is short relative to skull length (fig. 10). The lambdoidal ridge is prominent for a shrew of this size. The interorbital region is relatively straight (i.e., not strongly tapered) when viewed from the dorsal aspect. The dentition is prominent relative to the palatal breadth (fig. 18A). NMKR BobAaTe NM = BoAfkCApb VKR V COMPARISONS: Crocidura elongata is readily distinguished from most shrew species on Sulawesi by its long body and even longer tail, relatively pale color, and long and narrow hind feet and skull (tables 2, 5). Its head-and-body length is considerably longer than all species except C. rhoditis and C. quasielongata. Absolute hind-foot and ear lengths are on average greater in C. elongata than in any other shrew on Sulawesi (fig. 9; table 2). However, the other members of the Elongata Subgroup, first described below, can be difficult to distinguish (fig. 11; table 4). Compared to C. elongata, C. microelongata is smaller bodied, with a darker pelage and shorter average tail length. The thenar pad on the hind foot of C. elongata (fig. 14C) is considerably longer than in either C. microelongata (fig. 14D) or C. quasielongata (fig. 14B). The skull of C. microelongata is also shorter but jO = AT T = tfaTe SKR = Coltk S elongata C. microelongataC. quasielongata C. pmbCfbp TABLE 4 Results of Principal Components Analysis of Craniodental Measurements of the Elongata Subgroup of Crocidura a Table entries for variables are component loadings. nearly as wide at the braincase and interorbital region, hence its relative width (BB /CIL and IOW /CIL) is greater than that of C. elongata (figs. 10, 12). Compared to C. elongata, C. quasielongata has a similar head-and-body length, but, on average, a shorter tail and paler pelage (fig. 12, table 2). The skull of C. elongata is very similar in length to that of C. quasielongata (figs. 12, 18). However, in C. elongata the rostrum makes up a smaller proportion and the postpalatal region makes up a greater proportion of skull length than in C. quasielongata (figs. 10, 12). In addition, relative braincase breadth (BB / CIL and BB / IOW) is, on average, slightly less in C. elongata than in C. quasielongata. A principal components analysis of 12 cranial measurements shows that these two species have broad, though not complete, overlap in multivariate morphometric space (fig. 11). COMMENTS: A nearly complete cytochrome b sequence (1109 bp) from a topotypical paratype (USNM 217535) is nearly identical to those from specimens we collected on Mt. Ambang (fig. 4), which is approximately 32 km to the southwest. Our inference based on mitogenomes also place USNM 217535 as a close relative to specimens from Mt. Ambang (fig. 5). These animals��� mitochondrial sequences are also closely related to a series from Mt. Buliohuto and to specimens from Mt. Dako referred to as ���dark elongata��� by Esselstyn et al. (2019). Maximum Jukes-Cantor distances between these sample localities is 0.05 (fig. 4; supplementary data S 5). Samples of this species from Temboan and Mts. Ambang, Buliohuto, and Dako formed a well-supported clade in our analyses of UCEs (figs. 7, 8) and concatenated nuclear exons (supplementary data S 6). The phylogeographic study of Eldridge et al. (2018), which examined the correspondence of genetic diversity in Elongata Subgroup shrews to the area of endemism paradigm expectations, conflated the three species of this subgroup with Crocidura elongata. More recently, specimens of C. elongata from Mt. Dako were referred to as ���dark elongata��� by Esselstyn et al. (2019) because the authors were unable to determine if either of two long-tailed species on Mt. Dako represented true C. elongata. Mt. Dako is the only area where we found C. elongata occurring in syntopy with another member of the Elongata Subgroup (C. quasielongata). Despite the confusing history of specimens in this subgroup, none of our analyses suggested a sister relationship between any two of these species. In our UCE species-tree inference, C. elongata was moderately supported as the sister to C. rhoditis and C. pseudorhoditis (fig. 7), but in our mitogenome estimate it was placed as sister to C. lea, although without statistical support (fig. 5). Ruedi (1995) suggested a scansorial lifestyle for this species based on its long, naked-appearing tail and long hind feet. While this is certainly possible, direct evidence for a scansorial lifestyle is lacking, and these traits could be linked alternatively to a saltatorial locomotory style (Brosset, 1988). Some very limited evidence indicates that Crocidura caudipilosa, which has a long, but less extreme tail combined with a more typical hind-foot length, is a skilled climber (Esselstyn et al., 2019). For coalescent species delimitation results, see the Crocidura quasielongata account below. SPECIMENS EXAMINED: Mt. Ambang (LSUMZ 39008���39013, 39015���39018, 39057, 39058, 39061, 39248���39251, 39257���39264, 39318; NMV C38009, C38032), Mt. Buliohuto (LSUMZ 38238, 38240, 38243���38247, 38251���38253; NMV C37742, C37752, C37760), Mt. Dako (LSUMZ 36905���36907, 36909, 36916, 36919, 36921, 36923, 36924, 36932; NMV C37248, C37249, C37303), Temboan (USNM 217534, 217535)., Published as part of Esselstyn, Jacob A., Achmadi, Anang S., Handika, Heru, Swanson, Mark T., Giarla, Thomas C. & 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) on pages 23-33, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835, {"references":["Esselstyn, J. A., A. S. Achmadi, H. Handika, T. C. Giarla, and K. C. Rowe. 2019. A new climbing shrew from Sulawesi highlights the tangled taxonomy of an endemic radiation. Journal of Mammalogy 100: 1713 - 1725.","Musser, G. G. 2014. A systematic review of Sulawesi Bunomys (Muridae, Murinae) with the description of two new species. Bulletin of the American Museum of Natural History 392: 1 - 313.","Ruedi, M. 1995. Taxonomic revision of shrews of the genus Crocidura from the Sunda Shelf and Sulawesi with description of two new species (Mammalia: Soricidae). Zoological Journal of the Linnean Society 115: 211 - 265.","Eldridge, R. A., A. S. Achmadi, T. C. Giarla, K. C. Rowe, and J. A. Esselstyn. 2018. Geographic isolation and elevational gradients promote diversification in an endemic shrew on Sulawesi. Molecular Phylogenetics and Evolution 118: 306 - 317.","Brosset, A. 1988. Le peuplement de mammiferes insectivores des forets du nord-est du Gabon. Revue d'Ecologie, La Terre et la Vie 43: 23 - 46."]}

Crocidura normalis, new species LSID: urn:lsid:zoobank. org:act: A639DEFB-7B56-4B2F-9916- 333E95A8B499 HOLOTYPE: MZB 43010 (= FMNH 213437), an adult male, collected by J.A. Esselstyn on 27 March 2011. The specimen was prepared as a study skin, cleaned skull (fig. 41 A) and skeleton, and frozen tissues. External measurements from the holotype are: 124 mm × 55 mm × 13 mm × 8 mm = 4.9 g. The voucher specimen and a tissue sample will be permanently curated at MZB, with an additional tissue sample retained at FMNH. TYPE LOCALITY: Indonesia, Sulawesi Tengah, Poso, Huasa, Sedoa, Lore Lindu National Park, Mt. Rorekatimbo; 1.2884° S, 120.3104° E, 2250 m. ETYMOLOGY: Normalis is Latin for “normal,” used in recognition that this is yet another species of shrew with no striking phenotypic traits worthy of attaching a descriptive name. GEOGRAPHIC DISTRIBUTION: We recorded this species at relatively high-elevation sites in Sulawesi’s west-central (Mt. Gandang Dewata, West Sulawesi Province; Mts. Latimojong and Rorekatimbo, Central Sulawesi Province), eastcentral (Mt. Katopasa, Central Sulawesi Province) and south-east areas of endemism (Mt. Mekongga, Southeast Sulawesi Province; figs. 13, 39). We found this species at sites ranging from 1400 to 2600 m (fig. 13, table 3). DIAGNOSIS: Crocidura normalis is a mediumsized shrew (tables 2, 14), with a light build (fig. 17) and dark brown pelage, feet, lips, pinna, and tail. Dorsoventrally, the color is relatively uniform. The mystacial vibrissae are short relative to body length and darkly pigmented proximally for about half of their length. The tail can be as long as head-and-body length but is usually shorter (fig. 9). It is relatively densely covered with bristles, many of which are darkly pigmented from the base of the tail along approxi- mately two-thirds of its length (fig. 40A). The small, applied hairs that typically cover the tail of Crocidura are less distinctly noticeable than in many other species. The feet are brown to black, usually uniformly so, including the digits and the thenar and hypothenar are subequal in area (fig. 40A). The claws are translucent. The skull is relatively gracile, being narrow across its entire length for a Crocidura of this size (fig. 10; table 14). Rostral length makes up a comparatively small proportion of skull length (fig. 10). The maxillary process is not prominent, the palate is narrow, and the occlusal surface area of the dentition is small relative to skull size (fig. 41A). The maxillary bridge is narrow. COMPARISONS: Crocidura normalis is smaller than all members of the Long-Tailed, Thick- Tailed, and Rhoditis groups, and larger than all members of the Small-Bodied Group. As the smallest member of the Ordinary Group, C. normalis is only slightly larger than C. mediocris (the TABLE 15. Results of Principal Components Analysis of Craniodental Measurements of Crocidura nigripes a Table entries for variables are component loadings. largest member of the Small-Bodied Group). It differs from C. mediocris in having a darker, denser pelage (individual hairs approximately 5 mm at middorsum) and longer tail both absolutely and relative to HBL (fig. 9). Within the Ordinary Group, C. musseri has a wider braincase relative to skull length, more robust body (fig. 17), longer rostrum relative to skull length (fig. 10), and paler feet than C. normalis. Crocidura nigripes has a short tail, like that of C. normalis, but is substantially larger, usually has darker feet, and the skull of C. nigripes shows a more prominent dentition, longer rostrum, and narrower interorbital region relative to skull length (fig. 10). Crocidura ordinaria and C. solita are slightly larger than C. normalis, have longer tails, both absolutely and relative to HBL, have hypothenar pads larger than thenar pads, and have greater relative rostral lengths (RL/CIL), braincase breadths (BB /CIL), and interorbital widths (IOW /CIL) (fig. 10). COMMENTS: In addition to the coalescent species delimitation analyses described above in the Crocidura parva account, we used BPP to test species limits between C. normalis and C. mediocris. The dataset consisted of 20 specimens of C. mediocris and 36 individuals of C. normalis. The sequence alignment is 93% complete. All replicates supported these species with 1.0 posterior probability. Unfortunately, we do not have nuclear exon data from the two specimens of C. normalis from Mt. Mekongga, which formed a clade independent of C. normalis from other localities in the cytochrome b and mitogenome gene trees (figs. 4, 5). In our UCE inferences, the Mekongga specimens of C. normalis are represented and the species is monophyletic in both our species tree and concatenated estimates (figs. 7, 8). If specimens from additional localities from the south-east and west-central areas of endemism are collected, reexamining these populations would be worthwhile. In both UCE estimates, C. normalis is a member of a clade comprising several Small-Bodied Group members and C. caudipilosa (figs. 7, 8). TABLE 16 Results of Principal Components Analysis of External Measurements of Crocidura nigripes a Table entries for variables are component loadings. Ruedi (1995) referred three specimens (IZEA 4393, 4394, 4042) to Crocidura lea and later Ruedi et al. (1998) sequenced a fragment of cytochrome b in one of these individuals (IZEA 4394). That sequence is identical to our C. normalis cytochrome b sequences from Mt. Rorekatimbo. Although we have not examined these three specimens, we presume they all represent C. normalis. SPECIMENS EXAMINED: Mt. Gandang Dewata (MZB 34809, 34810, 34812, 34821; FMNH 218604–218613, 218652–218658, 218973– 218975), Mt. Katopasa (NMV Z61813), Mt. Latimojong (FMNH 213006–213008, 213010, 213011, 213013, 213014, 213030, 213032; MVZ 237595, 237622–237624), Mt. Mekongga (MWFB 8151, 8154), Mt. Rorekatimbo (FMNH 213174– 213189, 213257, 213438; MZB 43010).