Crocidura normalis, Esselstyn & Achmadi & Handika & Swanson & Giarla & Rowe, 2021

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 : 78-82

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

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 ).

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 ).

Figures

FIG. 41. Images showing dorsal, ventral, and lateral views of the skull and lateral and occlusal views of the dentary of three species belonging to the Ordinary Group: A, Crocidura normalis, FMNH 213457; B, C. ordinaria, FMNH 218726; and C, C. solita, FMNH 213044.

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. 39. Map of Sulawesi showing localities sampled for shrews. Colored areas enclose localities with known records of members of the Ordinary Group. To maintain clarity of presentation, the widespread Crocidura nigripes is not included.

FIG. 17. Box plots showing the length of the hind foot (HF) relative to the head-and-body length (HBL) and the ratio of mass to HBL. 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 along the x-axis according to the species groups used in the text (Thick = Thick-Tailed Group). Lengths are in mm and mass is in grams.

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. 40. Images showing the ventral surface of the hind foot and dorsal surfaces of the tail base (approximately 1 cm from rump) and tail tip from three members of the Ordinary Group: A, Crocidura normalis, FMNH 213174 (left hind foot); B, C. ordinaria, FMNH 218726 (right hind foot); and C, C. solita, FMNH 213015 (left hind foot). Upper scale bar for foot, lower bar corresponds to both tail images.

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. 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.

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.