Crocidura elongata 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 elongata

description: Crocidura elongata Miller and Hollister, 1921   
 Crocidura elongata Miller and Hollister, 1921: 101. Original description.   
.  
 Crocidura “dark elongata” Esselstyn et al., 2019: 1715. Informal name.   
 HOLOTYPE: USNM 217534, an adult male obtained by H.C. Raven on 1 August 1916. The specimen comprises a skin and skull. External measurements recorded from the type are 214 mm × 120 mm × 22 mm; no ear length or weight was recorded. TYPE LOCALITY: “Temboan (southwest from Tondano Lake), northeastern Celebes” (Miller and Hollister, 1921: 101; fig. 1). We estimate that the type locality is at 0.979° N, 124.605° E, 650 m elevation, which differs from other interpretations (e.g., Musser, 2014). See the gazetteer for a full explanation (appendix).   
 GEOGRAPHIC DISTRIBUTION: Apparently restricted to the northern peninsula of Sulawesi, where clear records are known from the northeast (the type locality, Temboan and Mt. Ambang, North Sulawesi Province) and north-west (Mt. Dako, Central Sulawesi Province, and Mt. Buliohuto, Gorontalo Province) areas of endemism (fig. 16). The absence of records from the north-central area of endemism are almost certainly due to the general lack of mammal collecting from this region. Miller and Hollister’s (1921) paratypes from “Pinedapa, eastern Middle Celebes ” and the two specimens from Mt. Rorekatimbo reported by Ruedi (1995) almost certainly represent Crocidura microelongata (see below). Crocidura elongata is known from a moderately broad range of elevations, with the low elevation records between 500 and 600 m from Mts. Buliohuto and Dako and high-elevation records reaching 1600 m on Mt. Dako (table 3). The type locality at Temboan is around 650 m elevation.   TABLE 2 Descriptive Statistics a  for External Measurements (mm) and Mass (g) for Species of Sulawesi Crocidura   
 a The sample mean ± one standard deviation, the observed range in parentheses, and the sample size.   
 Long-Tailed Ordinary Rhoditis Small-Bodied Thick ● ●● ● ● ● ●● ● 54 43 34 ● 40 17 78 27 15 43 5 27 75 20 7 21 20 26 27 6 5 11 = = = = = = = = = N = = = = N = = = = = N = N = = N N N N N N N N N N N N N N N N N ● ● ● ● ● ● ● ●● ● ● ● ● ● ● = 54 = 44 = 33 = 40 = 17 = 81 = 27 = 15 = 43 = 5 = 27 = 75 = 22 = 7 = 21 = 20 = 26 = 27 = 6 = 5 = 11 N N N N N N N N N N N N N N N N N N N N N ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ● = 54 = 43 = 32 = 40 = 18 = 78 = 27 = 15 = 43 = 5 = 27 = 76 = 20 = 7 = 21 = 20 = 26 = 27 = 6 = 5 = 11 N N N N N N N N N N N N N N N N N N N N N ● ● ● ● ● ● ●● ● 54 44 34 40 17 81 27 15 ● 43 5 27 75 22 7 21 20 26 27 6 5 11 = = = = = = = = = N = = = = N = = = = = N = N = = N N N N N N N N N N N N N N N N N   TABLE 3  Elevational Ranges (m) and Species Richness of Crocidura on Mountains of Sulawesi  a  
 a Mountains are grouped by peninsula or area of endemism. Some mountain names are abbreviated: Amb = Ambang, Bulio = Buliohuto, Gand = Gandang Dewata, Lati = Latimojong, Bal = Balease, Rore = Rorekatimbo, Torom = Torompupu, Bawa = Bawakaraeng, Kato = Katopasa, Tomp = Tompotika, and Mek = Mekongga). When museum databases recorded an elevational range for an individual specimen, we used the median of that range as the elevation for that specimen. Specimens from Salu Tiwo are grouped under Mt. Gandang Dewata.   
 EMENDED DIAGNOSIS: A long-bodied, somewhat heavily built, moderately bicolored shrew with a very long tail and unusually long, pale hind feet (figs. 9, 14C, 17; tables 2, 5). The dorsal pelage is gray-brown overall; individual dorsal hairs are gray at the base and brown at the tip. The ventral pelage is paler, with individual hairs gray-based like those on the dorsum, except that the hair tips are silvery. The silvery appearance of the venter is most pronounced on the throat and chest area. In some specimens, however, this area has a reddish tint, due to variation in the color of the tips of some hairs. The mystacial vibrissae are dark proximally but white distally. The ears are prominent and pale. Dorsally, the feet are pale, transitioning from light brown near the wrist and ankle to nearly white on the digits. Ventrally, the feet show the same transition, but pigment is concentrated around the base of the pads (fig. 14C). The claws are translucent. The hind feet are long, even relative to head-and-body length (fig. 17). The long tail is subtly bicolored, with a brown dorsum and pale brown venter. Tiny applied hairs are present along the entire length of the tail, but they are barely visible to the naked eye (fig. 14C). However, these hairs are slightly longer and white near the tip of the tail, creating a very small white distal tuft. The long bristle hairs that are common at the base of the tail of many Crocidura are nearly absent in this species. The skull is long and slender, with a notably narrow braincase, interorbital region, and palate (figs. 10, 18A). Although the interorbital region is narrow relative to skull length, it is less constricted than the braincase. The long skull of C. elongata is attributable primarily to elongation of the postpalatal region; the rostral length is short relative to skull length (fig. 10). The lambdoidal ridge is prominent for a shrew of this size. The interorbital region is relatively straight (i.e., not strongly tapered) when viewed from the dorsal aspect. The dentition is prominent relative to the palatal breadth (fig. 18A).   
NMKR  
BobAaTe NM  
=  
BoAfkCApb VKR V  
 COMPARISONS: Crocidura elongata is readily distinguished from most shrew species on Sulawesi by its long body and even longer tail, relatively pale color, and long and narrow hind feet and skull (tables 2, 5). Its head-and-body length is considerably longer than all species except C. rhoditis and C. quasielongata. Absolute hind-foot and ear lengths are on average greater in C. elongata than in any other shrew on Sulawesi (fig. 9; table 2). However, the other members of the Elongata Subgroup, first described below, can be difficult to distinguish (fig. 11; table 4). Compared to C. elongata, C. microelongata is smaller bodied, with a darker pelage and shorter average tail length. The thenar pad on the hind foot of C. elongata (fig. 14C) is considerably longer than in either C. microelongata (fig. 14D) or C. quasielongata (fig. 14B). The skull of C. microelongata is also shorter but   
jO  
=  
AT T  
=  
tfaTe SKR  
=  
Coltk S  
 elongata C. microelongataC. quasielongata C. pmbCfbp   TABLE 4  Results of Principal Components Analysis of Craniodental Measurements of   the Elongata Subgroup of Crocidura   
 a Table entries for variables are component loadings.   
 nearly as wide at the braincase and interorbital region, hence its relative width (BB/CIL and IOW/CIL) is greater than that of C. elongata (figs. 10, 12). Compared to C. elongata, C. quasielongata has a similar head-and-body length, but, on average, a shorter tail and paler pelage (fig. 12, table 2). The skull of C. elongata is very similar in length to that of C. quasielongata (figs. 12, 18). However, in C. elongata the rostrum makes up a smaller proportion and the postpalatal region makes up a greater proportion of skull length than in C. quasielongata (figs. 10, 12). In addition, relative braincase breadth (BB/ CIL and BB/IOW) is, on average, slightly less in C. elongata than in C. quasielongata. A principal components analysis of 12 cranial measurements shows that these two species have broad, though not complete, overlap in multivariate morphometric space (fig. 11).   
 COMMENTS: A nearly complete cytochrome b sequence (1109 bp) from a topotypical paratype (USNM 217535) is nearly identical to those from specimens we collected on Mt. Ambang (fig. 4), which is approximately 32 km to the southwest. Our inference based on mitogenomes also place USNM 217535 as a close relative to specimens from Mt. Ambang (fig. 5). These animals’ mitochondrial sequences are also closely related to a series from Mt. Buliohuto and to specimens from Mt. Dako referred to as “dark elongata” by Esselstyn et al. (2019). Maximum Jukes-Cantor distances between these sample localities is 0.05 (fig. 4; supplementary data S5). Samples of this species from Temboan and Mts. Ambang, Buliohuto, and Dako formed a well-supported clade in our analyses of UCEs (figs. 7, 8) and concatenated nuclear exons (supplementary data S6).   
 The phylogeographic study of Eldridge et al. (2018), which examined the correspondence of genetic diversity in Elongata Subgroup shrews to the area of endemism paradigm expectations, conflated the three species of this subgroup with Crocidura elongata. More recently, specimens of C. elongata from Mt. Dako were referred to as “dark elongata” by Esselstyn et al. (2019) because the authors were unable to determine if either of two long-tailed species on Mt. Dako represented true C. elongata. Mt. Dako is the only area where we found C. elongata occurring in syntopy with another member of the Elongata Subgroup (C. quasielongata). Despite the confusing history of specimens in this subgroup, none of our analyses suggested a sister relationship between any two of these species. In our UCE species-tree inference, C. elongata was moderately supported as the sister to C. rhoditis and C. pseudorhoditis (fig. 7), but in our mitogenome estimate it was placed as sister to C. lea, although without statistical support (fig. 5).   
 Ruedi (1995) suggested a scansorial lifestyle for this species based on its long, naked-appearing tail and long hind feet. While this is certainly possible, direct evidence for a scansorial lifestyle is lacking, and these traits could be linked alternatively to a saltatorial locomotory style (Brosset, 1988). Some very limited evidence indicates that Crocidura caudipilosa, which has a long, but less extreme tail combined with a more typical hind-foot length, is a skilled climber (Esselstyn et al., 2019).   
 For coalescent species delimitation results, see the Crocidura quasielongata account below.   
SPECIMENS EXAMINED: Mt. Ambang (LSUMZ 39008–39013, 39015–39018, 39057, 39058, 39061, 39248–39251, 39257–39264, 39318; NMV C38009, C38032), Mt. Buliohuto (LSUMZ 38238, 38240, 38243–38247, 38251–38253; NMV C37742, C37752, C37760), Mt. Dako (LSUMZ 36905–36907, 36909, 36916, 36919, 36921, 36923, 36924, 36932; NMV C37248, C37249, C37303), Temboan (USNM 217534, 217535).
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 23-33, DOI: 10.1206/0003-0090.454.1.1, http://zenodo.org/record/5788835