Crocidura pallida Esselstyn &amp; Achmadi &amp; Handika &amp; Swanson &amp; Giarla &amp; Rowe 2021, new species
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 pallida

description: <i>Crocidura pallida</i>, new species LSID: urn:lsid:zoobank. org:act: 1249AC44-0D01-4060-B5FD- 3000D9C4D29D HOLOTYPE: MZB 43004 (= FMNH 210607), an adult female collected by J.A. Esselstyn on 18 October 2010. The specimen consists of a dried skin, cleaned skull (fig. 24B) and skeleton, and tissue sample. It carried one embryo measuring 18 mm in crown-rump length. External measurements from the holotype are: 140 mm × 62 mm × 15 mm × 8 mm = 11 g. The voucher specimen and a tissue sample will be permanently curated at MZB, with another tissue sample retained at FMNH. TYPE LOCALITY: Indonesia, Sulawesi Selatan, Luwu Utara, Sukamaju, Mt. Balease; 2.4995° S, 120.4874° E, 900 m elevation. ETYMOLOGY: We use the Latin <i>pallida</i> to highlight the pale color of the feet of this species. GEOGRAPHIC DISTRIBUTION: Recorded from the west-central (Mt. Gandang Dewata, West Sulawesi Province; Mts.Torompupu and Balease, Central Sulawesi Province; Mt. Latimojong and Rindingallo, South Sulawesi Province), east-central (Mts. Katopasa and Tompotika, Central Sulawesi Province), and south-east areas of endemism (Mt. Mekongga, Southeast Sulawesi Province; fig. 20). Across these areas, we found this species over a broad elevational range, from approximately 100 to over 2500 m (fig. 13; table 3). DIAGNOSIS: <i>Crocidura pallida</i> is a moderately sized shrew (figs. 9, 23; tables 2, 7) with very pale feet (fig. 21B) and a somewhat pale ventral side of the tail. The tail is shorter than the head-andbody length (fig. 9; table 2). The dorsal pelage is gray to gray-brown while the ventral pelage is pale gray. The pinna and the dorsal side of the tail match the dorsal pelage, but the dorsal side of the feet are distinctly paler. The dorsal surface of the hand is nearly white, but some pigment is present near the wrist. The hind feet show a similar pattern but are modestly darker. Ventrally, the feet are also pale, especially on the digits, which are usually white. The darkest parts of the hind foot are usually the thenar and hypothenar pads (fig. 21B). The pads of the forefeet are rarely pigmented. Tail bristles are sparse to nearly absent (fig. 21B), extending along no more than the proximal third of the tail length. The tail varies from uniformly colored to moderately bicolored with a paler ventral side. The skull is typical of <i>Crocidura</i> of this size. The braincase is generally rounded, but a subtle lateral point is evident at the mastoid region when viewed from a dorsal aspect (fig. 24B). The braincase is somewhat inflated dorsoventrally, and it is wide relative to skull length (fig. 24B), as are the interorbital region and rostrum (fig. 10; table 7). The wide interorbital region gives the maxillary process a relatively weak appearance. In some individuals, the nasal passage is particularly inflated and laterally bulging, further obscuring the maxillary process. The rostrum is somewhat long, relative to skull length (fig. 10). The maxillary bridge is usually narrow, with an anteriorly placed lacrimal foramen. The posterior portion of the hard palate is narrow, sandwiched between broad molars. COMPARISONS: <i>Crocidura pallida</i> is smaller than <i>C. rhoditis</i>, <i>C. pseudorhoditis</i>, <i>C. nigripes</i>, and members of the Elongata Subgroup and Thick-Tailed Group. It is considerably larger than all members of the Small-Bodied Group and somewhat larger than all members the Ordinary Group except <i>C. nigripes</i>. In absolute terms and relative to head-and-body length, the tail length is comparable to members of the Ordinary, Thick-Tailed, and Rhoditis groups, but substantially shorter than in all members of the Long- Tailed Group and considerably longer than in any member of the Small-Bodied Group (fig. 9). Outside of the Rhoditis Group, most species have much darker feet than <i>C. pallida</i>. These include <i>C. caudipilosa</i> and <i>C. microelongata</i> of the Long-Tailed Group, <i>C. normalis</i>, <i>C. musseri</i>, and <i>C. nigripes</i> of the Ordinary Group, both Thick-Tailed Group members, and all members of the Small-Bodied Group except <i>C. lea.</i> Within the Rhoditis Group, <i>C. pallida</i> is smaller in absolute measurements (fig. 9) and more delicately built than <i>C. rhoditis</i> and <i>C. pseudorhoditis</i> (fig. 17); it is paler with a relatively narrower braincase and interorbital region than the otherwise similarly proportioned <i>C. australis</i>. Rostral length makes up a smaller proportion of skull length (RL /CIL) in <i>C. pallida</i> than in either <i>C. rhoditis</i> or <i>C. pseudorhoditis</i>, but this trait is comparable in <i>C. australis</i> (fig. 10). COMMENTS: Substantial mitochondrial genetic divergence is evident between populations from Mt. Katopasa, Mt. Tompotika, and the remaining sites (up to 0.089 Jukes-Cantor distance; fig. 4; supplementary data S3). However, these populations form a cohesive set of morphological specimens, form a clade in our phylogenetic analyses of nuclear genes (figs. 7, 8; supplementary data S6), and all of the mitochondrial variation is partitioned geographically (i.e., no sympatry between divergent mitochondrial clades). We therefore did not divide them further. We identified a single specimen of <i>Crocidura pallida</i> from Mt. Latimojong (MVZ 237618), a locality where <i>C. solita</i> is abundant. Although <i>C. pallida</i> is slightly larger, these two can be difficult to distinguish. As such, it is possible that this specimen is a slightly large individual of <i>C. solita</i> with mtDNA introgressed from <i>C. pallida</i>. Unfortunately, we did not obtain nuclear loci (exons or UCEs) from this specimen and there- fore cannot test for introgression. Contamination of this cytochrome <i>b</i> sequence is unlikely because it is unique in our alignment. We favor the hypothesis that MVZ 237618 is <i>C. pallida</i> because its cytochrome <i>b</i> sequence differs slightly from <i>C. pallida</i> sequences from nearby localities (i.e., Mts. Gandang Dewata and Balease) and phenotypically, it is nearer the averages for <i>C. pallida</i> than <i>C</i>. <i>solita</i>. Phylogenetic estimates were not consistent regarding the relationships of <i>Crocidura pallida</i>. Our mitochondrial gene trees put it as sister to a clade of Small-Bodied species, <i>C. caudipilosa</i>, and <i>C. normalis</i> (figs. 4, 5). However, our nuclearbased inferences placed <i>C. pallida</i> as part of the large basal polytomy (figs. 7, 8; supplementary data S6). SPECIMENS EXAMINED: Mt. Balease (FMNH 210580–210592, 210608, 210609, MZB 43004), Mt. Gandang Dewata (FMNH 218687–218702, 218989; MZB 34872, 34886, 34888, 34889), Mt. Katopasa (LSUMZ 39527, 39529–39538; MVZ 238115–238118; NMV C40187, C40192, C40199, C40206, C40214, C40217, C40307, Z56723, Z62366, Z61754, Z62414), Mt. Latimojong (MVZ 237618), Mt. Mekongga (MWFB 8059, 8115, 8125, 8139, 8150, 8161, 8162, 8195, 8196, 8438, 8439), Rindingallo, Tana Toraja (MSB 93256), Mt. Tompotika (FMNH 213366–213369), Mt. Torompupu (NMV C40307).
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 49-52, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835