Crocidura religiosa (I. Geoffroy-Saint Hilaire, 1826 )

Woodman, Neal, Koch, Claudia & Hutterer, Rainer, 2017, Rediscovery of the type series of the Sacred Shrew, Sorex religiosus I. Geoffroy Saint-Hilaire, 1826, with additional notes on mummified shrews of ancient Egypt (Mammalia: Soricidae), Zootaxa 4341 (1), pp. 1-24 : 12-14

Crocidura religiosa (I. Geoffroy-Saint Hilaire, 1826)

Sorex religiosus I. Geoffroy Saint-Hilaire, 1826 ; 1827a, b, 1831; Fischer, 1829; Lichtenstein, 1829; Ehrenberg, 1835; Partington, 1837; Wagner, 1841; Knight, 1843; Smedley et al., 1845.

Crocidura religiosa: Fitzinger, 1868 ; de Winton, 1902; Cabrera, 1925; Flower, 1932; Setzer, 1957; Bodenheimer, 1960; Hoogstraal, 1962; Heim de Balsac & Meester, 1971; Gureev, 1971, 1979; Hutterer et al., 1982; Hutterer, 1993, 2005; Wolsan & Hutterer, 1998; Grimmberger et al., 2009; Aulagnier et al., 2009, Happold, 2013.

Crocidura nana (not of Dobson, 1890): Heim de Balsac & Verschuren, 1968; Heim de Balsac & Mein, 1971; Osborn & Helmy, 1980; Le Berre, 1990; Hoath, 2003; Ikram, 2005a.

Lectotype. ÄM 690, complete embalmed mummy ( Figs. 3 , 4 , 5 , 7 , 8 , S 2 ), acquired by J. Passalacqua in 1823; collector and preparator unknown.

Type locality. Specimen recovered from: Egypt, Qena Governorate, near Thebes, western bank of the Nile River, necropolis Dra ′ Abu el-Naga ′ (25°44’07”N, 32°37’14”E), tomb of Queen Mentuhotep. The exact source of the preserved specimen is unknown, but presumed to be from the vicinity of Thebes.

Diagnosis. Small and greyish-brown Crocidura with paler underparts and limbs. Fur short and silky. Tail thick at base and covered with long bristle-like hairs over most of its length. Head and body length 54 mm, tail 26-40 mm, hindfoot short (8–10 mm s.u., 9–11 mm c.u.). Skull short (GLS 14.4-16.1 mm, Table 1) and slender; braincase flat and dorsal profile straight. Upper toothrow (I1-M3) 5.9-6.8 mm, height of coronoid process (COR) 3.0-3.7 mm. Dentition not specialized. First upper incisor robust, but of medium size ( Figs. 3 , 5 , S 2 ). Upper unicuspid teeth with well-developed cinguli. Upper premolar (P4) with a short parastyle. M1 and M2 with well-separated protocone and hypocone. Upper third molar small. Cutting surface of lower incisor smooth.

Description of the lectotype. Specimen ÄM 690 is fully preserved and embalmed ( Fig. 4A ). The cover of resin and other preservatives obscures the colour of the fur and longer bristles. Head and body length is 46 mm, tail length 28 mm (61% of head and body length), and hind foot length 8 mm. An X-ray image of the body ( Fig. 4B ) shows that the skeleton is complete. The age of the specimen based on suture fusion and tooth wear is young adult. The skull ( Figs. 3 , 5 , S 2 ) is short (GLS 15.3 mm) and slender with a flat dorsal profile. The rostrum is long (PL 6.0 mm) and narrow (MB 4.8 mm), and the interorbital region is comparatively long and narrow (IO 3.2 mm). The braincase is long and wide (BB 6.8 mm). Lambdoid crests are developed. The mandible is short and robust (COR 3.6 mm), with a slender angular process. Length of the upper toothrow (I1-M3) is 6.2 mm. The first upper incisor is robust with a long anterior hook and a pronounced secondary cusp ( Figs. 3 , 5 ). There are three pairs of unicuspid teeth, the first one largest, the followed posteriorly by two that are subequal in size. Cinguli on the unicuspids are well pronounced. The upper premolar (P4) has a short parastyle. M1 and M2 are almost equal in size, and the protocone and hypocone on these teeth are clearly separated. The cutting surface of the lower incisor is smooth.

We provide an interactive 3D scan of the skull of the lectotype online in the Supplementary material ( Fig. S2 ).

Paralectotypes. We consider all of the 17 surviving specimens of small shrews as part of the original type series of S. religiosus I. Geoffroy Saint-Hilaire, 1826 a . However, two specimens belong to a different species and must be excluded. Of the 15 remaining specimens identified as C. religiosa based on external measurements ( Table 3 ), we were able to examine only four in detail using micro-CT-scans and high resolution X-rays ( Table 4). We therefore propose to regard only specimens ÄM 688, 7051, 7056 as paralectotypes until such time that the remaining specimens also can be examined in detail.

Distribution. Today, C. religiosa is confined to the Upper Nile valley and delta ( Fig. 1 ). Its current population status is unknown ( Hutterer et al., 2008; Happold, 2013). The most recently reported collection dates from 1988 ( Handwerk, 1990). A possible Pleistocene record from Bir Tarfawi indicates the species also may have occurred in southern Egypt at that time, but the identity of the fossil fragments needs to be confirmed ( Kowalski et al., 1989; Butler, 1998).

Vernacular name. We propose to use "Sacred Shrew" as the English common name for this small species. The name coined for it by I. Geoffroy Saint-Hilaire (1826: 294–295) is “musaraigne sacrée,” first translated into English by Partington (1837) and subsequently used by Smedley et al. (1845), Reichenbach (1852: "Heilige Spitzmaus"), Fitzinger (1868), Bodenheimer (1960), and Woodman (2015). “Egyptian Pigmy Shrew” has been used by Le Berre, 1990, Wolsan & Hutterer (1998), Wilson & Cole (2000), Hutterer (2005), Aulagnier et al. (2008), and Happold (2013), while Osborn & Helmy (1980), Osborn & Osbornová (1998), and Hoath (2003) used “Dwarf Shrew.”

Illustrations. No photograph of a live C. religiosa exists. The best approximation of the appearance of the living shrew is a colour plate by the Victorian artist Pierre Jacques Smit (1863–1960; Schoonraad, 1964), which shows the silky greyish-brown pelage, pale (whitish) limbs, and the thick and hairy tail ( Fig. 6 ). Other drawings in Le Berre (1990) and Aulagnier et al. (2009) show the dorsal fur as too pale and are less likely to represent the natural colors.

Figures

FIGURE 1. Map of northeastern Egypt, showing the distribution of Crocidura religiosa along the Nile River valley. Symbols: solid triangle, original type locality at Thebes; open triangles, other ancient necropoli (Lortet & Gaillard, 1903; Boessneck, 1988; Kessler, 2007); solid square, Corbet’s (1978) neotype locality at Giza; solid dots, modern specimen localities (de Winton, 1902; Bonhote, 1909, 1912; Flower, 1932; Setzer, 1957; Osborn & Helmy, 1980; Handwerk, 1990, 1998). A presumed Pleistocene record from Bir Tarfawi, southern Egypt (Kowalski et al., 1989) is not included.

FIGURE 2. Mummified remains of Sorex religiosus (figure 1, top), the middle-size shrew (figure 2, middle: Crocidura floweri?), and Crocidura olivieri (figure 3, bottom) from Thebes (I. Geoffroy Saint-Hilaire, 1827 b: Plate 4, Figure 1). The shrews illustrated in figs. 1 & 2 are now lost; the specimen of C. olivieri is identifiable as ÄM 7044. Image from the Biodiversity Heritage Library (www. biodiversitylibrary. org). Digitized by Smithsonian Libraries, Washington. Original page is ca. 258 x 190 mm.

FIGURE 3. Dorsal views of the skulls of mummified (micro-CT-scans) and extant (camera lucida drawings) specimens of shrews. C. religiosa: A, Passalacqua specimen ÄM 690 (GLS 15.3 mm), C, Giza, Egypt, BMNH 10.6. 18.5 (GLS 15.6 mm). C. pasha: B, Passalacqua specimen ÄM 697 (GLS 14.1 mm), D, Khartoum, Sudan, BMNH 9.9. 2.31 (GLS 14.1 mm).

FIGURE 4. Lectotype ÄM 690 of Sorex religiosus I. Geoffroy Saint-Hilaire, 1826. A, External view of mummy; B, X-ray of entire specimen. Scale in mm. Photograph courtesy of S. Steiss, Berlin; X-ray image courtesy of C. Schmidt, Berlin.

FIGURE 5. X-ray (A) and micro-CT-scans (B-E) of the skull of the lectotype AM 690 of Sorex religiosus I. Geoffroy Saint- Hilaire, 1826 (GLS 15.3 mm): A, B, lateral views; C, sagittal cross section: D, dorsal view; E, ventral view. See the interactive 3 D scan in Fig. S 2.

FIGURE 6. Crocidura religiosa, as seen by the Victorian artist P. J. Smit. Detail from plate 23 in Anderson & de Winton (1902).

FIGURE 7. Micro-CT-scans of a cross section through the head and the lateral view of the skull of an embalmed shrew (ÄM 690) from the Passalacqua Collection. The skull is surrounded by dry tissue and mineral dust. Width of bony braincase is 6.8 mm. Please see interactive 3 D scan in Fig. S 2.

FIGURE 8. Taphonomic features of the Passalacqua shrew mummies. A, distorted thoracic vertebrae of ÄM 690; B, distorted thoracic vertebrae of ÄM 7050; C, broken braincase of ÄM 7055; D, broken braincase of ÄM 7051. X-rays (A-C) and Micro-CTscan (D).