Crocidura caudipilosa Esselstyn et al., 2019

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, Bulletin of the American Museum of Natural History 2021 (454), pp. 1-109 : 21-23

Crocidura caudipilosa Esselstyn et al., 2019

Crocidura caudipilosa Esselstyn et al., 2019:

1718. Original description.

HOLOTYPE: MZB 41456 (= LSUMZ 36945 ), an adult male collected 8 March 2013 by J.L. Patton. The skin, skull, skeleton, and a tissue are held at MZB, while an additional frozen tissue is curated at LSUMZ. External measurements and weight recorded from the holotype are 182 mm × 98 mm × 19 mm × 9 mm = 12 g.

TYPE LOCALITY: Indonesia, Sulawesi Tengah, Toli Toli, Malangga Selatan, Mt. Dako ; 1.10642° N, 120.9106° E, 512 m elevation.

GEOGRAPHIC DISTRIBUTION: Widespread on Sulawesi across most areas of endemism, with records from the west-central (Mts. Rorekatimbo and Torompupu, Central Sulawesi Province; Mt. Gandang Dewata, West Sulawesi Province; Mt.

Latimojong, South Sulawesi Province), north- west (Mt. Dako, Central Sulawesi Province; Mt. Buliohuto, Gorontalo Province), north-east (Mt. Ambang, North Sulawesi Province), south-west (Mt. Bawakaraeng, South Sulawesi Province), east-central (Mt. Katopasa, Central Sulawesi Province), and south-east (Mt. Mekongga, Southeast Sulawesi Province) areas of endemism. The lack of records from the north-central area of endemism is almost certainly due to the lack of collecting effort in this region. Occurs over a wide elevational range from approximately 500 m on Mt. Buliohuto to at least 2600 m on Mt. Gandang Dewata ( fig. 13 ; table 3 ).

ABBREVIATED DIAGNOSIS: Esselstyn et al. (2019) made detailed comparisons to other Sulawesi species found on Mt. Dako, of the north-west area of endemism. Briefly, Crocidura caudipilosa is a slender grayish to brownish shrew with a hairy tail ( fig. 14A ) that is moderately longer than the head and body ( table 2 ). The feet are brown in dorsal view, with paler digits. The skull is gracile, with rounded features and a weakly developed dentition ( fig. 15 ). Summary statistics for measurements of material we refer to this species are provided in tables 2 and 5 .

COMPARISONS: This species is readily distinguished from all other Crocidura on Sulawesi by the density (combined number and length) of applied hairs on the tail (short hairs that lie close to the tail, not the longer bristles that typically project away from the tail on many species of Crocidura ), but it is also distinguishable by various combinations of its body size, tail length relative to head-and-body length ( fig. 9 ), gray to brown pelage, slight bicoloration (darker dorsum, paler venter), slender skull, and relatively small teeth. Its tail is shorter relative to headand-body length than in any of the Elongata Subgroup members, but longer (absolutely and relatively) than in any other congeneric species

on Sulawesi ( fig. 9 ). Body size is considerably smaller than in C. elongata and C. quasielongata , but only slightly smaller than in C. microelongata ( fig. 9 ). The skull and hind feet are not particu- larly elongate, as they are in members of the Elongata Subgroup. The ratio of braincase breadth to interorbital width is greater, on aver- age, than in any other shrew species on Sulawesi, and considerably so compared to any of the Elongata Subgroup members ( fig. 10 ). COMMENTS: Individuals of Crocidura caudipi- losa have been caught (NMV Z56802) and observed (NMV Z62413) climbing trees (Essel- styn et al., 2019). The extent of time this species spends in trees is unknown, but the possibility that some shrews on Sulawesi exploit arboreal resources is a potentially promising explanation for how so many species coexist on the island. Musser (1982: 81) reported collecting an unde- scribed species of Crocidura “in moss growing 8 feet above ground around a tree trunk.” How- ever, he never described this species, and it is unclear whether he was referencing a member of the Long-Tailed Group or some other species Crocidura caudipilosa is unusually widespread across Sulawesi, but Jukes-Cantor distances calcu- lated from mtDNA sequences were <0.024 ( fig. 4 ; supplementary data S3), suggesting recent, wide- spread movement of at least maternally inherited markers. Nuclear loci sampled from across the island show the same pattern, with variation detected at only five of 549 nucleotides among 56 sequences of apolipoprotein b, for instance. This species is sister to Crocidura levicula in our UCE- ( figs. 7 , 8 ) and nuclear exon-based phylogenies, although statistical support from the latter estimate was absent (supplementary data S6). Our mitochondrial estimates placed C. caudipilosa in an unresolved clade of several small, darkly colored species of shrew, among which C. levicula is included ( figs. 4 , 5 ). This clade, which was well supported by our UCE analyses, contained no other member species of the Long-Tailed Group ( figs. 7 , 8 ).

SPECIMENS EXAMINED: Mt. Ambang ( LSUMZ 39243–39247 ), Mt. Bawakaraeng ( MVZ 237625– 237627 ; NMV Z57152, Z57013, Z57043, Z57044), Mt. Buliohuto ( LSUMZ 38288–38291 , 38296 , 38588 , 38656 ; NMV C37746, C37815), Mt. Dako ( LSUMZ 36940 , 36942 , 36945 ; NMV C37267, C37304, C37305, C37330, C37351), Mt Gandang Dewata ( FMNH 218759–218762 , 218768 , 218983– 218986 ; MZB 34746, 34747 , 34951 , 34952 ), Mt. Katopasa ( LSUMZ 39355–39357 ; MZB 39840, 39871 , 39922 ; NMV C40186, C40219, C40220, Z61815, Z62413, Z62415), Mt. Latimojong ( FMNH 213020–213028 , 213427 ; MVZ 237612– 237615 ; MZB 41649; NMV C38591), Mt. Mekongga ( MWFB 8158 , 13512 ), Mt. Rorekatimbo ( FMNH 213246–213248 , 213441 ), Mt. Torom- pupu ( LSUMZ 39358 , 39359 ).

Figures

FIG. 13. Elevational records of all species of Crocidura known from Sulawesi. Each point represents a specimen. For specimens associated with a minimum and maximum elevation, we used the center of the given elevational range. Sample sizes are given above the x-axis. Species are grouped according to the species groups used in the text (Thick = Thick-Tailed Group).

FIG. 14. Images showing the ventral surface of the left hind foot and dorsal surfaces of the tail base (approximately 1 cm from rump) and tail tip from the four members of the Long-Tailed Group: A, Crocidura caudipilosa, LSUMZ 36940; B, C. quasielongata, FMNH 218551; C, C. elongata, LSUMZ 39009; and D, C. microelongata, FMNH 212998. Scale bars represent 5 mm and apply to their nearest images within each panel. The thenar (T) and hypothenar (H) pads are labeled on panel B.

FIG. 15. Images showing dorsal, ventral, and lateral views of the skull and occlusal and lateral views of the dentary of Crocidura caudipilosa (FMNH 218986).

FIG. 9. Box plots showing variation in external measurements from all species of Sulawesi shrew. Plots show the median, 1st and 3rd quartiles, the maximum value within 1.5 × interquartile range (distance between 1st and 3rd quartiles; IQR), the minimum value within 1.5 × IQR, and outliers (black circles). Sample sizes are shown along the x-axis. Species are grouped according to the species groups used in the text (Thick = Thick- Tailed Group). All measurements in mm. HBL = head-and-body length.

FIG. 10. Box plots of relative skull measures showing braincase breadth (BB), interorbital width (IOW), and rostral length (RL) divided by condyloincisive length (CIL) and BB divided by IOW for all species of Sulawesi shrew. Plots show the median, 1st and 3rd quartiles, the maximum value within 1.5 × interquartile range (distance between 1st and 3rd quartiles; IQR), the minimum value within 1.5 × IQR, and outliers (black circles). Sample sizes are shown along the x-axis. Species are ordered according to the species groups used in the text (Thick = Thick-Tailed Group).

FIG. 4. Maximum-likelihood estimate of the gene tree of Sulawesi Crocidura derived from an alignment of 851 individuals and 1111 characters from the mitochondrial gene cytochrome b. Bootstrap support is shown along branches. Clades corresponding to species are collapsed for ease of presentation. Tips are labeled with the species name, the number of tips (T), and number of localities (L), as labeled in figure 1, and the maximum intraspecific (MI) Jukes-Cantor distance calculated from a reduced alignment. Two species are paraphyletic and their respective, within-clade MI values are shown separately. For species described by Miller and Hollister (1921), the holotype or paratypes are included for C. elongata, C. lea, and C. rhoditis. Branch lengths between C. nigripes and other taxa are shortened for presentation. See supplementary data S2 for the full tree.

FIG. 7. Estimated species tree from analysis of 3940 ultraconserved element loci in ASTRAL. Samples from Sulawesi are labeled with the species name, locality, and catalog number. Asterisks indicate type specimens from Miller and Hollister (1921). Local posterior probabilities <0.95 are shown. Tip branch lengths are arbitrary.

FIG. 8. Estimated phylogenetic relationships from a maximum likelihood analysis of 983 concatenated ultraconserved elements. Samples from Sulawesi are labeled with the species name, locality, and catalog number. Asterisks indicate type specimens from Miller and Hollister (1921). Ultrafast bootstrap values <95 are shown.

FIG. 5. Maximum likelihood estimate of the mitochondrial gene tree derived from an analysis of 14,007 characters (representing protein-coding and rRNA genes) from 83 samples. Bootstrap support values <95 are shown at nodes. Tips are labeled with the species, locality, and voucher number.