Fourteen New, Endemic Species Of Shrew (Genus Crocidura) From Sulawesi Reveal A Spectacular Island Radiation Author Esselstyn, Jacob A. Author Achmadi, Anang S. Author Handika, Heru Author Swanson, Mark T. Author Giarla, Thomas C. Author Rowe, Kevin C. text Bulletin of the American Museum of Natural History 2021 2021-12-15 2021 454 1 109 https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-454/issue-1/0003-0090.454.1.1/Fourteen-New-Endemic-Species-of-Shrew-Genus-Crocidura-from-Sulawesi/10.1206/0003-0090.454.1.1.full journal article 2886 10.1206/0003-0090.454.1.1 3a7cdaa4-c2f2-4370-9301-c2ab339b2e0d 0003-0090 5788835 7982B923-4CDC-44ED-A598-8651009DC7CC Crocidura pseudorhoditis , new species LSID : urn:lsid:zoobank. org:act: 980DF37A-D8F0-4DBF-877C- A11C04F55FC1 Crocidura rhoditis Ruedi, 1995: 255 . Misidentification. 11.5 Mt. Ambang Crocidura rhoditis Esselstyn et al., 2019: 1716 . Misidentification. BREADTH 10.5 11 Sulawesi-wide HOLOTYPE : MZB 43002 (= LSUMZ 39310), an adult female, collected on 22 February 2016 by H. Handika. The specimen was prepared as a cleaned skull and skeleton, study skin, stomach, intestine, and frozen tissues. External measurements from the type are 148 mm × 70 mm × 16 mm × 11 mm = 11 g . The voucher specimen and a tissue sample will be permanently curated at MZB, with another tissue sample retained at LSUMZ. BRAINCASE 9.5 10 FIG. 23. Box plots summarizing head-and-body length, condyloincisive length, and braincase breadth measurements for members of the Rhoditis Group. Data from Crocidura rhoditis and C. pseudorhoditis are divided into two boxes. Dark gray boxes show all available measurements from across the species range while light gray boxes show only samples taken on Mt. Ambang, where the two species occur in syntopy. Note that the differences between these two species are greater on Mt. Ambang than they are for the islandwide sample. 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. All measurements in mm. 9 N = 5 N = 27 N = 35 N = 76 N = 10 N = 20 ralis aust C. pallida C. rhoditis C. pseudorhoditis C . SPECIES TYPE LOCALITY: Indonesia ; Sulawesi Utara ; Bolaang Mondgondow , Passi Timur ; Insil; Mt. Ambang , near Lake Aliyah ; 0.76385° N , 124.41188° E , 1481 m elevation. ETYMOLOGY: We combine “pseudo” with “rhoditis” because this species looks very similar to C. rhoditis . GEOGRAPHIC DISTRIBUTION: We recorded this species from the northwestern portion of the west-central area of endemism (Mts. Torompupu and Rorekatimbo, Central Sulawesi Province ), and the north-west and north-east areas of endemism of the northern peninsula ( fig. 20 ; Mt. Dako, Central Sulawesi Province ; Mt. Buliohuto, Gorontalo Province ; and Mt. Ambang, North Sulawesi Province ). The absence of records from the north-central area of endemism is almost certainly due to the lack of general mammal surveys and specimens from this region. Both specimens we report from Mt. Rorekatimbo were trapped by Ruedi (1995) ; we did not collect any Crocidura pseudorhoditis on Mt. Rorekatimbo, despite working at similar elevations. We found this species at a range of elevations ( fig. 13 ), the low on Mt. Buliohuto ( 500 m ) and the high on Mt. Rorekatimbo ( 2200 m ; table 3 ). DIAGNOSIS: Crocidura pseudorhoditis is a moderately large shrew ( tables 2 , 7 ) with a somewhat stocky build, medium gray, moderately bicolored pelage, a tail that is shorter than headand-body length, and dorsal foot surfaces that range from light gray-brown to pinkish white. The forefeet are paler than the hind feet and the claws are unpigmented and accompanied by a small tuft of white hairs on the hands, but on the hind feet, some of these hairs are dark. The tail is barely dorsoventrally bicolored proximally, but this transitions to a uniformly colored tail for the distal half of tail length. The tail has only sparse bristles over the proximal half of its length ( fig. 21C ). The lips are gray-brown, but the nose is distinctly paler. The palmar surface ranges from gray-brown to white, being palest on the digits. The color of the plantar surface also transitions from darker to paler colors toward the digits ( fig. 21C ), but all aspects are darker than on the forefeet. The skull is long ( table 7 ) and broader at the braincase and interorbital region than expected given its length ( figs. 10 , 22B ). The rostrum is long relative to the postpalatal portion of skull length, reflected in the RL/CIL ( fig. 10 ). The suture between the squamosal and parietal bones is often open and in the shape of a sickle blade just below the opening of the sinus canal ( fig. 22B ). The maxillary bridge is wide. COMPARISONS: Crocidura pseudorhoditis is smaller than C. rhoditis , C. elongata , and C. quasielongata , but larger than C. caudipilosa , Rhoditis Group members C. australis and C. pallida , all members of the Small-Bodied Group, and all members of the Ordinary Group except C. nigripes , which is similarly sized ( fig. 19 ; tables 2 , 7 ). The body of C. pseudorhoditis is more robust than those of C. microelongata , C. australis , and C. pallida , but less stocky than in C. rhoditis and C. caudicrassa ( fig. 17 ). Crocidura pseudorhoditis is slightly darker (both pelage and feet) than C. rhoditis and tail bristles, though not abundant, are still more prevalent than in C. rhoditis ( fig. 21 ). The thenar and hypothenar pads on the hind foot are both more rounded than the oblong pads of C. rhoditis ( fig. 21 ). Crocidura pseudorhoditis is similar in pelage color to C. australis and C. pallida . Braincase breadth relative to skull length is greater than in many of the species with somewhat similar body sizes, including C. rhoditis , and all Elongata Subgroup and Thick-Tailed Group species. However, C. pallida is comparable in this regard, and C. australis has an even greater relative skull breadth no matter where it is measured ( fig. 10 ). The braincase is a bit more angular in C. pseudorhoditis than in C. rhoditis , with the widest part of the cranium in C. pseudorhoditis forming an obtuse point ( fig. 22 ). However, the angularity of the braincase does not reach the degree seen in C. nigripes . The ratio of rostral length to skull length is large in C. pseudorhoditis and exceeded only by that of C. rhoditis ; the other Rhoditis Group members have comparatively short relative rostral lengths (RL/CIL) and outside the Rhoditis Group, C. nigripes and the two Thick- Tailed species (defined below) are comparable ( fig. 10 ). Crocidura pseudorhoditis is larger in head-and-body length, foot length, and all cranial dimensions than C. australis and C. pallida ( fig. 19 ; tables 2 , 7 ). The slight difference in size between C. pseudorhoditis and C. rhoditis appears to be enhanced where the two species cooccur (Mt. Ambang; fig. 20 ), especially condyloincisive length ( fig. 23 ). A principal components analysis of 12 cranial measurements that only included these two species readily separated them on the first axis, which represents size ( fig. 19 ; table 6 ). COMMENTS: While Crocidura pseudorhoditis is on average smaller than C. rhoditis , the two species are easily confused phenotypically and they occur in sympatry in the north-east area of endemism ( fig. 20 ; table 3 ). The large mitochondrial distances between C. rhoditis and C. pseudorhoditis , along with the slight but consistent differences in size, which appear to be enhanced where they are syntopic ( fig. 23 ), strongly suggest these are independently evolving populations. Our BPP analyses tested species limits with an alignment that contains 15 C. rhoditis and 58 C. pseudorhoditis and is 96% complete. The analyses supported these species’ distinction with 1.0 posterior probability with all prior combinations in all runs. One of the few cases of clarity from our phylogenetic analyses is the consistent and well-supported sister relationship between Crocidura rhoditis and C. pseudorhoditis ( figs. 4 , 5 , 7 , 8 ; supplementary data S6). Given their phenotypic similarity, this makes sense from a morphological perspective. In our UCE species tree, we found these two species sister to C. elongata ( fig. 7 ). SPECIMENS EXAMINED: Mt. Ambang ( LSUMZ 39030 , 39031 , 39034–39036 , 39039–39043 , 39045 , 39048 , 39049 , 39051 , 39052 , 39056 , 39059 , 39060 , 39063 , 39066 , 39067 , 39278 , 39280–39283 , 39293 , 39297 , 39298 , 39300– 39309 , 39311–39314 , 39316 , 39317 , 39322 ; MZB 43002 ; NMV C37985 , C37992 , C37996 , C38017 ), Mt. Buliohuto ( LSUMZ 38275–38278 , 38284 ; NMV C37793 ), Mt. Dako ( LSUMZ 36973 , 36978 , 36979 , 36986 , 36990 , 36991 , 36993 , 36995 , 36998 , 37000 , 37002 , 37005 , 37007 , 37008 , 37011–37016 , 37033 , 37034 ; MZB 38564–38566 ; NMV C37252 , C37254 , C37264 , C37268 , C37292 , C37297 , 37308 , 37315 , C37328 , 37329 , C37363 , C37367 ), Mt. Torompupu (NMV C40308), Mt. Rorekatimbo ( RMNH 38409 (= IZEA 4406 ), IZEA 4407 ) .