Crocidura lea 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 lea

description: <i>Crocidura lea</i> Miller and Hollister, 1921 <i>Crocidura lea</i> Miller and Hollister, 1921: 102. Original description. <i>Crocidura</i> “pale lea” Esselstyn et al., 2019: 1715. Informal name. HOLOTYPE: USNM 217553, an adult male collected by H.C. Raven on 3 August 1916. Prepared as a skin and skull. External measurements from the holotype are 111 mm × 51 mm × 14 mm; ear length and weight were not recorded. TYPE LOCALITY: “Temboan, northeastern Celebes ” (Miller and Hollister, 1921: 102; fig. 25). Temboan is located at 0.979° N, 124.605° E, 650 m elevation in the Southeast Minahasa Regency, North Sulawesi Province, 6 km south of Kalait. See the gazetteer (appendix) for details of our interpretation of Raven’s field notes. GEOGRAPHIC DISTRIBUTION: Probably endemic to the northern peninsula. We identified specimens of this species from the northwest area of endemism (Mt. Dako, Central Sulawesi Province and Mt. Buliohuto, Gorontalo Province) and the type locality is in the north-east area of endemism (Temboan, North Sulawesi Province). We did not find this species on Mt. Ambang (north-east area of endemism), where its ecological position is apparently held by another tiny, but not closely related shrew, <i>Crocidura tenebrosa</i> (fig. 25). We recorded <i>C. lea</i> at both high and low elevations on both Mts. Dako (512 and 1600 m) and Buliohuto (400– 600 and 1200 m; table 3). We did not collect any specimens of this species at our 410 m site on Mt. Dako. The type locality (Temboan) is at roughly 650 m. 120°E 122°E 124°E 126°E 1.5°N C. lea 0° C. baletei C. tenebrosa C. levicula C. mediocris <i>C. parva</i> 1.5°S Recent sample sites 3°S Miller and Hollister (1921) type localities 100 km 4.5°S 0–1000 m 1000–2000 m 6°S&gt; 2000 m EMENDED DIAGNOSIS: <i>Crocidura lea</i> is a very small shrew (tables 2, 8) with a brown dorsal pelage, gray-brown ventral pelage, and pale feet. The pinnae are prominent and slightly paler than the surrounding fur, especially at the lobe. The mystacial vibrissae are generally unpigmented, but a few, typically the longer ones, are brown or black proximally. Dorsally, foot color transitions from gray-brown at the wrist and ankle to pale pinkish white on the digits (fig. 27B). The claws are small, unpigmented, and sometimes lack the surrounding tuft of white hairs present in many other species. Ventrally, the forefeet mostly lack pigment, but the hind feet are darker. In the hind feet, color transitions from brown posteriorly from the heel, and to a lesser extent laterally, to white at the digits. However, pigment is concentrated in the pads, particularly around the base of the thenar, hypothenar, and in some cases the 4th (lateralmost) interdigital pad (fig. 27B). The tail is shorter than head-and-body length, sometimes dorsoventrally bicolored, and thinly covered with very small, dark brown applied hairs. A few sparse bristles are present along the proximal three-fourths of the tail (fig. 27B). The skull is short, narrow, and somewhat rounded in overall shape (fig. 28B). Relative to body size, the skull is slightly longer than expected, and relative to skull length, the braincase (BB /CIL) is especially narrow, but the interorbital region is not unusual in its relative width (IOW /CIL; fig. 10). The maxillary process is weak (fig. 28B). The I3 A B 0.5 8.5 BREADTH 8. 2 0 Species <i>C C.. lea baletei</i> BRAINCASE7.5 COMP -0.5 <i>C C C C.... parva levicula mediocris tenebrosa</i> 16 17 18 CONDYLOINCISIVE LENGTH -2 -1 0 1 2 COMP. 1 (U2) is smaller than the C (U3). The parastyle of P4 is modest. COMPARISONS: <i>Crocidura lea</i> is much smaller than all Sulawesi <i>Crocidura</i> outside of the Small- Bodied Group (fig. 9). Among the Small-Bodied species, only C. <i>baletei</i> and <i>C. tenebrosa</i> are known from the northern peninsula. <i>Crocidura levicula</i>, <i>C. parva</i>, and <i>C. mediocris</i> are apparently restricted to the central core and eastern and southwestern peninsulas (fig. 25). We therefore focus our comparisons on the cooccurring, northern peninsula species. However, first we note that the allopatric <i>C. levicula</i> has a broader skull (figs. 10, 26; tables 8, 9), shorter tail, shorter hind feet, and more tail bristles than does <i>C. lea</i> (figs. 27, 29; table 2). Among the Small-Bodied shrews of the northern peninsula, <i>C. lea</i> has a slightly paler and more strongly dorsoventrally bicolored pelage than <i>C. baletei</i> or <i>C. tenebrosa</i>. Both of the latter species are relatively uniform in color, <i>C. baletei</i> being medium brown and <i>C. tenebrosa</i> dark brown. <i>Crocidura lea</i> also has paler skin wherever it is exposed than either <i>C. baletei</i> (somewhat darker) or <i>C. tenebrosa</i> (much darker). Dorsally, the feet of both <i>C. baletei</i> and <i>C. tenebrosa</i> are uniformly dark, whereas the foot color transitions posteriorly from brown at the heel to white or pink at the digits in <i>C. lea</i> (fig. 27). Similarly, the tails of both <i>C. baletei</i> and <i>C. tenebrosa</i> are not obviously bicolored, and both are darker. Tail bristles are present on approximately three-fourths of the tail length in all three species, but they are much more abundant and longer on the tails of <i>C. baletei</i> and <i>C. tenebrosa</i> than in <i>C. lea</i> (fig. 27). Applied hairs on the tail are also much more visible in the two new species than in <i>C. lea</i>. Tail length in <i>C. lea</i> is greater, on average, than in <i>C. baletei</i>, and substantially so compared to <i>C. tenebrosa</i> (fig. 9; table 2). The skulls of <i>C. lea</i> are comparable in length, but significantly narrower at the braincase, interorbital region, maxillary process, and to a lesser extent at the rostrum than those of <i>C. baletei</i> and <i>C. tenebrosa</i> (figs. 10, 30; table 8). <i>Crocidura lea</i> occupies distinct morphometric space from the other northern peninsula endemics in bivariate plots of skull length and width and the first two principal components from an analysis of 12 cra- TABLE 8 a The sample mean ± one standard deviation, the observed range in parentheses, and the sample size. nial measurements (fig. 26). The C (U3) is larger than I3 (U2) in <i>C. lea</i>, but they are usually subequal in occlusal surface area in <i>C. baletei</i> and <i>C. tenebrosa</i>. The parastyle of P4 is more prominent in <i>C. tenebrosa</i> than in <i>C. lea</i> or <i>C. baletei</i>. COMMENTS: An 1111 bp cytochrome <i>b</i> sequence from the holotype (USNM 217553) provides strong evidence that the tiny shrews we collected on Mts. Buliohuto and Dako (samples from Mt. Dako were referred to as “ <i>Crocidura</i> pale lea” by Esselstyn et al., 2019) represent this species. Interestingly, we did not find <i>C. lea</i> on Mt. Ambang, our sample site nearest the type locality. Rather, the tiny shrews we collected from Mt. Ambang represent an undescribed spe- cies (<i>C. tenebrosa</i>) that does not appear to be a close relative of <i>C. lea</i>. We suspect that <i>C. lea</i> is widespread at relatively low elevations on the northern peninsula, but is displaced by smallerrange, darker-colored endemics in some higherelevation areas (fig. 13; table 3). Our only surveys on Mt. Ambang were centered on 1500 m elevation (fig. 3). For BPP species delimitation results, see the <i>C. tenebrosa</i> account below. Ruedi (1995) reported <i>Crocidura lea</i> from Mt. Rorekatimbo, but we refer those specimens to <i>C. normalis</i> (see below) based on a cytochrome <i>b</i> sequence from Ruedi et al. (1998). None of our phylogenetic estimates found strong support for the closest relative(s) of <i>Croc-</i> <i>idura lea</i>, but some consistencies are nevertheless apparent. Our mitochondrial analyses placed <i>C. lea</i> as the sister to <i>C. rhoditis</i> and <i>C. pseudorhoditis</i> (fig. 4) or <i>C. elongata</i> (fig. 5), while our UCE species tree placed <i>C. lea</i> as sister to the clade containing <i>C. elongata</i>, <i>C. rhoditis</i>, and <i>C. pseudorhoditis</i> (fig. 7). Other nuclear-based inferences lacked support (fig. 8; supplementary data S 6). TABLE 9 <b>Results of Principal Components Analysis of Craniodental Measurements of the Small-Bodied Group of <i>Crocidura</i></b> a Table entries for variables are component loadings. SPECIMENS EXAMINED: Mt. Buliohuto (LSUMZ 38254–38262, 38271–38274; NMV C37692, C37715, C37716, C37720, C37721, C37735, C37744, C37829), Mt. Dako (LSUMZ 36947– 36950, 36952, 36954–36956, 36958, 36960, 36964, 36968–36972), Temboan (USNM 217553). TABLE 9 <b>Results of Principal Components Analysis of Craniodental Measurements of the Small-Bodied Group of <i>Crocidura</i></b> a Table entries for variables are component loadings. SPECIMENS EXAMINED: Mt. Buliohuto (LSUMZ 38254–38262, 38271–38274; NMV C37692, C37715, C37716, C37720, C37721, C37735, C37744, C37829), Mt. Dako (LSUMZ 36947– 36950, 36952, 36954–36956, 36958, 36960, 36964, 36968–36972), Temboan (USNM 217553).
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 52-57, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835
description: {"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.", "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.", "Ruedi, M., M. Auberson, and V. Savolainen. 1998. Biogeography of Sulawesian shrews: testing for their origin with a parametric bootstrap on molecular data. Molecular Phylogenetics and Evolution 9: 567 - 571."]}