Crocidura levicula 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 levicula

description: Crocidura levicula Miller and Hollister, 1921   
 Crocidura levicula Miller and Hollister, 1921: 103. Original description.   
N = 7 N = 21 N = 6  
HOLOTYPE: USNM 219450, an adult female collected by H.C. Raven on 13 February 1918. Specimen prepared as a skin and skull. External measurements from the type are 108 mm × 44 mm × 11 mm; ear length and weight were not recorded. TYPE LOCALITY: “Pinedapa, Middle Celebes” (fig. 25). Pinedapa is located near the north coast of Central Sulawesi Province, in Poso Regency, just west of the town of Mapane. Musser (2014) placed the locality at 1.4167° S, 120.5833° E, 31 m elevation.  
2.3  
WIDTH 2.2 2.1  
PALATAL 1.9 2.0  
1.8  
N = 6 N = 22 N = 6  
WIDTH 4.2 INTERORBITAL 3.8 4.0  
GEOGRAPHIC DISTRIBUTION: The type locality is near sea level in the northwestern part of Sulawesi’s west-central area of endemism. We found this species to occur in the northern part of the west-central area of endemism (Mts. Rorekatimbo and Torompupu, Central Sulawesi Province), in two lowland areas near the boundary between the west-central and south-east areas of endemism (Wasponda and Tolala, Central Sulawesi Province), and in the east-central area of endemism (Mts. Katopasa and Tompotika, Central Sulawesi Province; fig. 25). These records span a broad elevational range, from approximately 400 m on Mt. Tompotika to 2000 m on Mt. Rorekatimbo (fig. 13; table 3).  
N = 7 N = 21 N = 6  
 C. baletei C. lea C. tenebrosa   
SPECIES  
EMENDED DIAGNOSIS: A very small (tables 2, 8), medium- to dark-brown shrew, with almost uniformly colored pelage, feet, pinna, and tail. The ventral pelage is only slightly paler than the dorsal pelage. The digits are similarly pigmented as the other foot parts. The tail is unusually short (table 2), with an abundance of bristles (some of which are pigmented) extending along the proximal two-thirds of tail length (fig. 29A). The hind foot is short, with darkly pigmented pads. The claws are pale, perhaps lightly pigmented, and surrounded by a small tuft of brown hairs (fig. 29A). The skull is small but broad, more so at the braincase than at the interorbital region (figs. 10, 31A). Relative to body size and skull breadth, the skull is quite short, particularly the rostrum (fig. 10). The C (U3) is equal to or larger in occlusal area than the I3 (U2).  
 COMPARISONS: Crocidura levicula is one of the smallest shrew species on Sulawesi, only being comparable to other members of the Small-Bodied Group (fig. 9). It has a shorter relative tail length than any other shrew species on Sulawesi (fig. 9). Its hind-foot length is shorter than in all Sulawesi shrews except one other member of the Small-Bodied Group, C. parva (fig. 9; table 2). Crocidura levicula is also darker in color than both C. lea and C. baletei, but not C. tenebrosa. Absolute tail length is slightly less in C. levicula than in C. parva. Tail bristles are more abundant on specimens of C. levicula than in C. lea, C. tenebrosa, and C. parva. The skull of C. levicula is small, delicate, and broad at the braincase and to a lesser degree at the interorbital region (fig. 10). Its skull width relative to length is greater than in C. lea, C. parva, and C. mediocris, and to a lesser degree, greater compared to C. tenebrosa (fig. 26). Crocidura levicula has a lesser skull length and breadth than C. baletei (fig. 26), but these two species have similar relative skull widths (fig. 10). Samples of Crocidura levicula are nearly nonoverlapping with other members of the Small-Bodied Group in morphometric space in bivariate plots of skull length versus width and along the first two axes from a PCA of 12 cranial measurements (fig. 26). In C. levicula, the C (U3) is equal to or greater in occlusal surface area than is I3 (U2) (fig. 31), which is similar to the conformation in C. lea.   
 COMMENTS: Miller and Hollister (1921) named this species based solely on the holotype, and unfortunately, we failed to obtain DNA from this specimen. We followed the Tsai et al. (2020) phenol-chloroform extraction protocol and used two separate extractions. The addition of resalting the ethanol supernatants did not yield a quantity of DNA that could be detected with a Qubit 2.0 fluorometer using the dsDNA High Sensitivity Assay Kit (Thermo Fisher Scientific, Waltham, MA). As such, our identification of Crocidura levicula is based on the morphological similarity of the holotype to specimens we collected in areas nearest the type locality.   
 One specimen (NMV Z63390) from Mt. Torompupu appears to have an introgressed mitochondrion from syntopic Crocidura ordinaria, a member of the Ordinary Group detailed below. Our identification of this specimen is based on morphology alone; we did not sequence any nuclear DNA from this specimen and therefore cannot test our introgression hypothesis.   
 In our phylogenetic estimates, UCE data confidently placed Crocidura levicula as sister to C. caudipilosa (figs. 7, 8). However, our nuclearexon and mitochondrial inferences placed this species as a member of a clade of mostly Small- Bodied species that also includes C. caudipilosa (fig. 5; supplementary data S6).   
 Crocidura levicula was delimited by all BPP analyses. See the C. tenebrosa account above for details.   
SPECIMENS EXAMINED: Mt. Katopasa (LSUMZ 39512–39516; NMV C40180, C40181, C40200, C40203, C40207, Z55557), Pinedapa (USNM 219450), Mt. Rorekatimbo (FMNH 213192, 213193, 213271), Tolala (FMNH 210576, 210577), Mt. Tompotika (FMNH 213343 – 213365), Mt. Torompupu (LSUMZ 39446–39449; NMV C40292, C40294, C40306, C40311, Z63365), Wasponda (FMNH 210578, 210579).
description: 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 62-63, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835