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Russell A. Mittermeier, Don E. Wilson (2018): Tenrecidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 134-172, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6808230

11. Indian Hedgehog Paraechinus micropus French: Hérisson indien / German: Indischer Igel / Spanish: Erizo de India Other common names: Indian Desert Hedgehog, Pale Hedgehog Taxonomy. Erinaceus micropus Blyth, 1846, “Bhawulpore.” Interpreted by D. R. Frost and colleagues in 1991 as “Bahawalpur, Pakistan, 29°23’N, 71°39’L.” This species is monotypic. Distribution. Arid zones of SE Pakistan (extreme SE Balochistan, Punjab, and Sindh) and NW India (W Rajasthan, Gujarat, and extreme NW Maharashtra). Descriptive notes. Head-body 130-230 mm, tail 10-40 mm, ear 26-32 mm, hindfoot 24-26 mm; weight 300-600 g. The Indian Hedgehog is small and rounded, with pointed nose and short legs. It has dark legs and relatively small feet and claws. Color is highly variable, with some individuals tending toward melanistic and others toward albinistic. True albinos have been reported. The Indian Hedgehog has well-defined, dark,facial mask thatis variable in shape and color. Dorsal pelage has spines, with rough surfaces and strong grooves along their length; they average 19-23 mm long and have fairly extensive creamy-white tips. Spines can be banded with dark brown or black and white or yellow, but usually only one of these colors predominates; some individuals have red-golden tint. There is wide and conspicuous center parting of spines on crown of head and distinct whitish band across forehead. Ears protrude beyond adjacent spines, and snoutis pointed. Upper parts of face and cheeks are grayish white. Underparts can be blotched dark brown and white, varying from entirely brown to entirely white. Legs are dark. Limbs, abdomen, and area around eyes and muzzle are covered with reddish brown to reddish gray hairs. Hallux is somewhat reduced but well developed. Tail is short and stumpy and sparsely covered with hairs. Karyotype is 2n = 48. Habitat. Primarily deserts and other arid areas in dry sandy areas, tropical thorn forests, semiarid scrublands, and agricultural lands. The Indian Hedgehog also occupies slightly more mesic habitats, especially regions influenced by monsoons. Food and Feeding. The Indian Hedgehog usually feeds at night, and its typical gait is a trot at ¢.30 cm/sec. When pursuing prey or when frightened, it may travel up to 64 cm/sec. Its diet consists mainly of insects (especially dung beetles) but also solifugid spiders, scorpions, frogs, toads,lizards, small snakes, eggs of ground-nesting birds, and rodents. It will eat ripe fallen fruits ofjujube dates (Zizyphus, Rhamnaceae). It holds down prey with forepaws and begins consuming its victim posterior-end first. Apparently, it stores excess food in its burrow for future use. Captive Indian Hedgehogs have survived 4-6 weeks without food and water. Breeding. The Indian Hedgehog is polygynous, and probably only one litter is produced each year. Males are reproductively capable well in advance of the breeding season, which extends throughout the monsoon period in April-September. Litters are born in July-September when insects are most abundant. Litters usually have 1-2 young (range 1-6). Neonates are 47-54 mm long. Captive Indian Hedgehogs sometimes eat their young. Activity patterns. The Indian Hedgehog usually is nocturnal and rather sedentary. Dayresting sites can be under a pile of brushy debris or a bush, but typically they are in burrows. The Indian Hedgehog does not hibernate. Although insects generally are plentiful even in winter, it undergoes temporary episodes of torpidity during drought or when food is scarce. When torpid,it remains curled up in its burrow or other shelter. It digs its own burrows and caches food. Movements, Home range and Social organization. Indian Hedgehogs excavate their own burrows that have one opening and are ¢.46 cm long. They also occupy burrows abandoned by other small mammals including those of the geographically sympatric Indian Long-eared Hedgehog (Hemiechinus collaris); they often occupy the same burrow and territory throughout the year. Indian Hedgehogs are usually solitary and nonaggressive. Males and females meet only to breed. Except for nursing mothers, only one Indian Hedgehog lives in a burrow at a time, although three shared one burrow in captivity. They engage in self-anointing behavior. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Indian Hedgehog is widespread and common, but local populations are increasingly fragmented and isolated due to desertification and agricultural development. Bibliography. Beer (2003), Bidwai & Bawa (1981), Corbet (1988), Eisentraut (1952), Frost et al. (1991), Gupta & Sharma (1961), Hutterer (2005a), Krishna & Prakash (1956), Mahabal et al. (2015), Nowak (1999), Prakash (1959, 1960), Reeve (1994), Roberts (1997), Shah & Aravinda Babu (1977), Stone (1995b), Walton & Walton (1973)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 324, DOI: 10.5281/zenodo.6639332

Family ERINACEIDAE (HEDGEHOGS AND GYMNURES) • Mediumssized insectivores/omnivores, with stiff spines on head and back and relatively short rostrum (hedgehogs) or soft or coarse fur and shrew-like snout (gymnures). • 10-75 cm. • Palearctic, Afrotropical, and Indo-Malayan Regions. • Deserts, steppes, urban grasslands, forests, farmlands, alpine meadows, parks, and gardens (hedgehogs) or humid, tropical forests (gymnures). • 10 genera, 26 species, 4b taxa. • 2 species Endangered, 1 species Vulnerable; none Extinct since 1600., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 288, DOI: 10.5281/zenodo.6639332

25. Mindanao Gymnure Podogymnura truei French: Gymnure de Mindanao / German: Mindanao-Rattenigel / Spanish: Gimnuro de Mindanao Other common names: Long-footed Woodshrew, Mindanao Moonrat, Mindanao Wood-shrew, Philippine Wood Shrew Taxonomy. Podogymnura true: Mearns, 1905, “Mount Apo at 6,000 feet [= 1829 m] altitude, southern Mindanao, Philippine Islands.” Interpreted by D. R. Frost and colleagues in 1991 as “6°59’N, 125°16’E.” This species is monotypic. Distribution. Mindanao I, S Philippines. Descriptive notes. Head-body 205-220 mm, tail 40-70 mm, ear 19-22 mm, hindfoot 29-38 mm; weight 52-82 g (adult males), 55-78 g (adult non-pregnant females), and 60-84 g (pregnant females). Female Mindanao Gymnures are slightly larger than males in some external measurements, but sexes are similar in cranial dimensions. Pelage is long, soft, and full, and it has stiffer guard hairs. Upper parts are predominantly ort mixed with coarse reddish-brown hairs. Underparts are hoary and slightly mixed with brown hairs. Tail is robust, moderately haired and buffy to purplish-flesh colored. Nose and toes are flesh colored. Claws are tan, with a reddish-brown spot at middle. Snout,lips, ears, and feet are purplish-flesh colored. Ears are large and semi-naked. Long whiskers protrude out of long snout, which give them good tactile sensation. Dental formulais 13/3, C1/1,P 3/3, M 3/3 (x2) = 40. Chromosomal complement has 2n = 40 and FN = 76. Habitat. Damp, montane, and mossy forests at elevations of 1300-2900 m. The Mindanao Gymnure is common in montane forests at elevations of ¢.1600-2000 m and abundant in mossy forests at ¢.2000-2900 m. It has been documented near a hole at the base of a large tree, among tangled roots of trees, among roots in thick moss, near logs along creeks in dense forests with undergrowth offerns, under grass on the edge of a lake, and near boulders in a fern-covered valley. It also occursin areas with standing water. Food and Feeding. Diet of the Mindanao Gymnure includes beetles and other insects, worms, other invertebrates, and carrion. It seems especially fond of earthworms. Breeding. The Mindanao Gymnure probably is polygynous. Nine pregnant females were reported in March, April, and June. During the same periods, 18 adult females had large mammae but were not pregnant, and four others were nulliparous. None of 15 females captured in May and November was pregnant. Average crown-rump length of embryos is 17 mm (range 4-40 mm). Litters have 1-2 young. Juveniles and subadults of both sexes were recorded in March—-May. Activity patterns. Mindanao Gymnures are nocturnal or crepuscular, and they remain in their burrows during the day. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. At low elevations, the Mindanao Gymnure occurs in restricted patches of forested habitats that are being destroyed by logging and slash-and-burn agriculture; populations in high-elevation forests that have limited commercial value are stable. Native mountain tribes use the meat of the Mindanao Gymnure and other species of mammals as sources of protein. Bibliography. Beer (2003), Bhatnagar & El-Azawi (1978), Frost et al. (1991), Heaney & Morgan (1982), Heaney, Balete et al. (1998), Heaney, Tabaranza et al. (2006), Hoogstraal (1951), Hibner et al. (1991), Hutterer (2005a), Li Yuchun et al. (2008), Mearns (1905), Nowak (1999), Rabor (1986), Reumer & Meylan (1986), Rickart (2003), Stone (1996b), Zima & Kral (1984)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 329-330, DOI: 10.5281/zenodo.6639332

26. Grandidier’s Shrew Tenrec Muicrogale grandidieri French: Microgale de Grandidier / German: GrandidierKleintenrek / Spanish: Tenrec musarana de Grandidier Taxonomy. Microgale grandidieri Olson et al., 2009, “Madagascar: Province de Toli- ara, Sept Lacs, 23°31°29-27S,44°09'33-2"E, 140 m. The specific name of the site is Antafiky.” Microgale grandidier: forms a clade with M. brevicaudata. There is considerable variation among populations of M. grandidieri from south-western and west-central Madagascar. Monotypic. Distribution. W & SW Madagascar, from the Namoroka Massif S to Onilahy River. Descriptive notes. Head-body 60-80 mm, tail 33-43 mm, ear 12-14 mm, hindfoot 11-13 mm; weight 8-10 g. Tail of Grandidier’s Shrew Tenrec is ¢.50% of head-body length. Hindfootis relatively short. Dorsal pelage is fine and relatively soft, a mix of dark brown and lighter brown giving an agouti appearance; ventral pelage is finer in texture and uniform medium gray, separated from dorsum by well-demarcated line. Ears are short and covered with fine dark brown or black hairs. Habitat. Lowland dry and dry spiny forests, selectively logged and disturbed gallery forests, secondary riparian forests, and dense dry deciduous forests on limestone karst (“tsingy”) at elevations of 50-430 m. Food and Feeding. No information. Breeding. No information. Activity patterns. Grandidier’s Shrew Tenrec is seasonally active. Despite surveys extending for more than a year,it was trapped only during warm season along northern bank of Onilahy River, not in cold dry season, suggesting that it becomes torpid for extended periods when food availability is reduced. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Grandidier’s Shrew Tenrec is known from only nine localities, and its population trends are unknown. It occurs in several protected areas and is apparently tolerant of some habitat disturbance. Major threats include forest clearing for agriculture, timber extraction, and charcoal production. Bibliography. Goodman et al. (2013), Olson et al. (2009), Soarimalala & Goodman (2011), Stephenson et al. (2016u)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 171, DOI: 10.5281/zenodo.6808230

Russell A. Mittermeier, Don E. Wilson (2018): Erinaceidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 288-330, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6639332

17. Daurian Hedgehog Mesechinus dawuricus French: Hérisson de Daourie / German: Daurischer Igel / Spanish: Erizo del Gobi Taxonomy. Erinaceus dauuricus Sundevall, 1842, “Dauuria.” Interpreted by D. R. Frost and colleagues in 1991 as “[former] U.S.S.R., Transbaikalia, Dauryia, 49°57’N, 116°55°E,” Russia. This species is monotypic. Distribution. SE Siberia & SW Russian Far East (Transbaikalia E to upper Amur Basin), C & E Mongolia, and NC, NE & C {China. Descriptive notes. Head-body 241-267 mm, tail 26-34 mm, ear 18-21 mm, hindfoot 41-44 mm; weight 1.1-1 kg. Ears of the Daurian Hedgehog are moderately long, with slightly coarse fur, and they extend a little beyond adjacent spines. Dorsal pelage has spines that are 19-23 mm long. Spines are dark brown at bases, dull whitish for nearly one-half their lengths, with brownish black bands and 3-5mm white tips. Ventral fur is rather coarse, underparts are whitish, and feet and tail are chestnut-brown. Dental formulais13/2,C1/1,P3/2,M 3/3 (x2) = 36. Chromosomal complement has 2n = 44-48 and FN = 87-95. Habitat. Semiarid habitats including grasslands, with shrubs and herbaceous plants, and dry mountain and wooded steppes. In Mongolia, the Daurian Hedgehog occupies rocky outcrops and low-density shrub habitats and avoids areas with high densities of shrubs. Habitats occupied change between early and late summer, shifting to greater use oflow-density shrub areas and decreased use of forb-dominated short grassy areas. Food and Feeding. The Daurian Hedgehog feeds on insects, especially beetles, cockroaches, and grasshoppers, many other invertebrates, and small vertebrates including rodents, snakes, frogs, and eggs of ground-nesting birds. Breeding. The Daurian Hedgehog mates in mid-May after emergence from hibernation. Gestation lasts 35-40 days; 3-7 young are born in June-July. Activity patterns. The Daurian Hedgehog is terrestrial and digs its own burrows or uses those of other small mammals. Burrows, crevices in rocks, and caves are used as shelters and places to hibernate in winter. Daytime shelters have one, rarely two, entrances and usually are in rocky outcrops or at bases of shrubs. Movements, Home range and Social organization. In Mongolia, home ranges were 113-2172 ha and were larger in early summer than late summer. Nightly movements averaged 3-4 km (range 0-3-12-2 km). The Daurian Hedgehog selects level terrain and areas with shrubs, specifically wild apricots (Amygdalus pedunculata, Rosaceae). It probably forages around wild apricots because they support higher densities of insects, and level areas might be used to reduce energetic costs of foraging. Actual foraging paths of the Daurian Hedgehog have high diversities of shrubs, abundance of wild apricots, and low diversities of grasses, other herbaceous species, and Artemisia (Asteraceae). Status and Conservation. Classified as Least Concern on The IUCN Red List. Loss of habitat through increased mining activities and grazing by increasing numbers oflivestock probably degrade habitats of the Daurian Hedgehog. Occasional vehicular mortality constitutes a low-level threat. Poisoning of plague carriers, including hedgehogs, is not conducted on a large scale, butit still occurs locally. Bibliography. Allen (1938), Batsaikhan et al. (2010), Beer (2003), Corbet (1988), Frost et al. (1991), Hutterer (2005a), Kong Fei, Guo Jianmin & Wu Jiayan (2016), Kong Fei, Wu Jiayan & Guo Jianmin (2016a, 2016b), Kong Fei, Wu Jiayan, Guo Jianmin & Wu Xiaomin (2016a, 2016b), Korablev et al. (1996), Mallon (1985), Murdoch et al. (2006), Nowak (1999), Ognev (1928), Reading et al. (2010), Reeve (1994), Smith & Yan Xie (2013), Stone (1995b), Sundevall (1842), Thomas (1908b), Zapletal et al. (2012, 2015)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 327, DOI: 10.5281/zenodo.6639332

11. Montane Shrew Tenrec Macrogale monticola French: Microgale des montagnes / German: Gebirgs-Kleintenrek / Spanish: Tenrec musarana de montana Other common names: Mountain Shrew Tenrec Taxonomy. Microgale monticola Goodman & Jenkins, 1998, “11 km WSW of Befingi- tra, Réserve Spéciale d’Anjanaharibe-Sud, 14°44’S, 49°26’E, 1550 m,” Madagascar. Microgale monticola forms a clade with M. tava. Monotypic. Distribution. Marojejy and Anjanharibe-Sud (Northern Highlands), and Makira Sud, N Madagascar. Descriptive notes. Head-body 72-90 mm, tail 98-117 mm, ear 12-16 mm, hindfoot 19-21 mm; weight 12-18 g. Tail of the Montane Shrew Tenrec is 120% of head—body length. Dorsal pelage is dark brown and slightly grizzled; venter is dark brown. Tail is dark brown above and paler below, with scales clearly visible beneath short scale hairs. Habitat. Humid forests at elevations of 1500-1950 m. The Montane Shrew Tenrec has been documented at lower elevations (500-720 m) in Makira Sud. Food and Feeding. No information. Breeding. Number of embryos was 1-2 in wild-caught Montane Shrew Tenrecs. One of these individuals was pregnant and lactating—the first observation of this occurring in Microgale. Activity patterns. Montane Shrew Tenrecs are terrestrial. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Vulnerable on The IUCN Red List. The Montane Shrew Tenrec is known from only four locations on two massifs in northern Madagascar. Populations are assumed to be stable. Major threats are loss of habitat from illegal logging (even in Marojejy National Park and Anjanharibe-Sud Reserve), unsustainable fuelwood harvesting, and gemstone mining. Bibliography. Everson et al. (2016), Goodman & Jenkins (1998, 2000), Goodman et al. (2013), Jenkins (2003), Olson & Goodman (2003), Soarimalala & Goodman (2011), Stephenson et al. (2016s)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 167, DOI: 10.5281/zenodo.6808230

23. Shrew Gymnure Neotetracus sinensis French: Gymnure de Chine / German: Spitzmausigel / Spanish: Gimnuro de China Other common names: Chinese Moonrat Taxonomy. Neotetracus sinensis Trouessart, 1909, “Ta-tsien-lou, province of Se-tchouen (China Occidental), at an altitude of 2545 metres.” Interpreted by D. R. Frost and colleagues in 1991 as “Kanding, Sichuan Sheng, 30°07’N, 102°02’E,” China. This species is monotypic. Distribution. Sichuan, Guizhou, and Yunnan in S China, and adjacent parts of NE Myanmar and N Vietnam. Descriptive notes. Head-body 108 mm (males) and 110 mm (females), tail 65 mm (males) and 62 mm (females), ear 18 mm (males and females), hindfoot 26 mm (males) and 25 mm (females); weight 36 g (males) and 38 g (females). Pelage of the Shrew Gymnureis soft, dense, and long. Upper parts are olive-brown, cinnamon-brown, reddish yellow, or mixed cream-colored and black, and sides of head and neck sometimes are tinged with reddish. Indistinct, black dorsal stripe can be present. Underparts are reddish, buffy gray, grayish white, or cream-colored over dark base. Snout is pointed. Tail is ¢.50% of length of head-body, somewhat slender, and sparsely covered with minute hairs. Feet are brown, five-toed, slender, and elongated, especially hindfeet; forefeet are tinted with reddish. Undersides of feet are naked or covered with short hair. Ears are brown, rounded, large, and covered with short hairs. Microstructures of hairs and vibrissae of the Shrew Gymnure are similar to those on the Long-eared Gymnures (Hylomys megalotis) and the Shorttailed Gymnure (Hylomys suillus); there are spear-shaped lead hairs and zig-zag guard hairs, but there are no downy hairs. Largest guard hairs of the Shrew Gymnure are long and form upper outer layer of pelage. Zig-zag hairs and wavy hairs are present among smaller guard hairs. Dental formula is 13/3, C 1/1, P 3/3, M 3/3 (x2) = 40. Karyotype is 2n = 32. Habitat. Subtropical evergreen forests, cool damp rainforests in montane areas, and close to and away from streams at elevations of 1500-2700 m. Food and Feeding. Diet of the Shrew Gymnure contains invertebrates, rhizomes, berries, and other vegetation. Of those examined, stomach of one individual contained earthworms and the other vegetable matter. Breeding. Breeding of Shrew Gymnures appears to occur year-round. Pregnant females have been observed in April and August. Two litters probably are born each year; litters have 5-7 young. Activity patterns. Shrew Gymnures are nocturnal and strictly terrestrial. They live in runways and underground burrows along moss and fern-covered banks and under logs, rocks, and other cover. It is not known if they make these runways because other small mammals also use them. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Shrew Gymnure is common in parts of its wide distribution, but it occupies relatively restricted habitats that are subject to increasing encroachment by humans. Bibliography. Abramov, Shchinov & Rozhnov (2008), Allen (1938), Anthony (1941), Beer (2003), Chernova, Kulikov & Abramov (2015), Chernova, Kulikov, Shchinov & Rozhnov (2012), Frost et al. (1991), Heaney & Morgan (1982), Hutterer (2005a), Jenkins & Robinson (2002), Lu Lu et al. (2013), Mu Huagiang et al. (2011), Nowak (1999), Smith & Yan Xie (2008, 2013), Stone (1995b), Trouessart (1909), Wu Yi et al. (2011), Ye Jianping et al. (2006)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 329, DOI: 10.5281/zenodo.6639332

3. Southern White-breasted Hedgehog Erinaceus concolor French: Hérisson de Turquie / German: Sidlicher WeiRbrustigel / Spanish: Erizo oriental de vientre blanco Other common names: \White-bellied Hedgehog, White-chested Hedgehog Taxonomy. Erinaceus concolor Martin, 1838, “apud Trebizond.” Interpreted by D. R. Frost and colleagues in 1991 as “Trabzon, Turkey, 40°00°N, 39°43’E.” This species is monotypic. Distribution. Turkey (Anatolia) E to S Caucasus (Georgia, Armenia, and Azerbaijan) and S to NW Iran, N Iraq, Syria, Lebanon, Israel, and Jordan and on Chios, Samos, and Rhodes Is (Greece). Descriptive notes. Head-body 200-260 mm, tail 20-30 mm, ear 20-30 mm, hindfoot 41 mm; weight 550-800 g (some individuals weigh up to 1-5 kg). Dorsal pelage of the Southern White-breasted Hedgehog has spines up to 37 mm long. As in other species of Erinaceus,it has faintly visible central parting of spines on crown of head, smooth spines lacking papillae, and very well-developed hallux on hindfeet. When it walks, spines on forehead lean forward, and others lean backward. Ears are relatively short. Face, legs, and underparts are covered with non-spiny hairs. Overall color is dark brown, with distinctive patch of white fur across chest. Albinos are extremely rare. It is similar in size and appearance to the West European Hedgehog (E. europaeus), but it can be distinguished by distinctive white chest that contrasts with dark abdomen. Dental formula is 13/2, C1/1,P 3/2, M 3/3 (x2) = 36. Chromosomal complement has 2n = 48 and FN = 92-94. Habitat. Deciduous forests, scrublands, grasslands, and alpine regions below tree line but typically not extensive coniferous forests because of lack of ground cover. The Southern White-breasted Hedgehog is less common in treeless moors, uplands, and marshlands because it usually gathers broad, dry leaves to construct nests in burrows and other shelters that are used for hibernation. It prospers in wooded parks, gardens, golf courses, and farmlands; however, populations in farmlands are small or lacking where hedgerows have been removed and where use of pesticides has reduced populations of invertebrate prey. During dry years, it might move onto irrigated fields. In Turkey, it inhabits apple, pear, and mulberry orchards; agricultural fields of beets, spinach, corn, cabbage, and other vegetables; areas with thick grasses around barns and bushes; and urban areas. Food and Feeding. The Southern White-breasted Hedgehog mainly eats insects and other invertebrates but also frogs, snakes,lizards, young birds, mice, some plant matter, fungi, and carrion. It can supplementits diet by scavenging food discarded by humans. Breeding. Breeding season varies geographically: March to late May in southern populations and April-May and into July in northern areas. Gestation lasts 31-36 days. Young are born in nests constructed under foliage or other objects. Litters have 5-6 young (range 3-8). Neonates have an average crown-rump length of 55 mm (range 45-60 mm) and weigh 12-16 g; white-tipped spines soon emerge on their back. Eyes open in 14-18 days, and teeth appear at 21-24 days. Maximum life expectancy is c.6-8 years in the wild and ten years in captivity. Activity patterns. The Southern White-breasted Hedgehog primarily is active at night and rests by day under piles of brush or in leaf nests in rocky crevices and burrows. Itis a good swimmer and climber. Its spines, besides serving as protection from predators, act as a cushion whenitfalls or deliberately drops from a height. Usual gait is slow, ambling walk, but it can run rapidly. Movements, Home range and Social organization. The Southern White-breasted Hedgehog travels ¢.300 m/night, and maximum walking speed is ¢.17 m/minute. It makes large aboveground nests of leaves and grass beneath bushes or under grass. It uses simple burrows on hillsides and gradual slopes and can excavate elaborate burrows. In winter, nests are in gently sloping burrows, often between roots of bushes and trees. Burrows are up to 152 cm long and 76 cm deep. In an area where there were few trees, a nest burrow that was lined with leaves was in dense grass in an orchard. The Southern Whitebreasted Hedgehog is usually solitary, except during the breeding season when a female has young or when several individuals share the same burrow in winter. Average home ranges are 1-6 ha for males (range 0-8-2-3 ha) and adult females (range 0-1-2-4 ha). The Southern White-breasted Hedgehog performs self-anointing behavior, which might be a sexualsignal, provide protection from predators, or serve some other purpose. Typical defensive posture is to roll into a ball, covering vulnerable underside, face, and limbs with spines. It is quite docile in captivity. Status and Conservation. Classified as Least Concern on The IUCN Red List. Southern White-breasted Hedgehogs are killed by motor vehicles on roads and by humans for food in some regions. Bibliography. Arslan et al. (2008), Beer (2003), Corbet (1988), Frost et al. (1991), Harrison Matthews (1952), Hutterer (2005a), Karatas et al. (2007), Kazemi et al. (2016), Kral (1967), KryStufek & Vohralik (2001), Martin (1838), Masseti (2012), Nowak (1999), Ozen (2006), Qumesiyeh (1996), Reeve (1994), Schoenfeld & Yom-Tov (1985), Stone (1995b), Vasilenko (1988), Zherebtsova (1992)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 320, DOI: 10.5281/zenodo.6639332

4. Amur Hedgehog Erinaceus amurensis French: Hérisson de Mandchourie / German: AmurIgel / Spanish: Erizo de Manchuria Other common names: Chinese Hedgehog, Manchurian Hedgehog Taxonomy. Erinaceus amurensis Schrenck, 1859, “In der Nahe der Stadt Aigun, im mandshurischen Dorfe Gulssoja am Amur,” eastern Siberia, Russia. This species is monotypic. Distribution. Amur River and tributaries in Russian Far East (E from Zeya River) and S through Korean Peninsula and C & E China to E Sichuan and N Guangdong. Descriptive notes. Head-body 158-300 mm, tail 17-42 mm, ear 16-26 mm, hindfoot 34-54 mm; weight c.1-4 kg (males) and 800-950 g (females). Weights in China are reported as 0.6-1 kg. The Amur Hedgehog is thick-bodied, with well-developed hallux on hindfeet, short tail, and relatively short ears. Dorsal pelage has sharp spines up to 24 mm long; spines are smooth and lack papillae. It has faintly visible, central parting of spines on crown of head. Face, legs, and underparts are covered with coarse hairs. Color varies from pale yellow to dark brown, and unpigmented spines usually are present among pale-tipped, yellow, and brown-banded ones. Its spines, besides serving as protection against enemies, act as a cushion when it falls or drops deliberately from a height. Dental formulais13/2,C1/1,P 3/2, M 3/3 (x2) = 36. Habitat. Various habitats from farmlands to deciduous forests, including grasslands, scrublands, suburban parks, gardens, and alpine regions below tree line. The Amur Hedgehog avoids truly arid areas and dry meadow-steppes. Its northern distribution is largely confined to northernmost limit of deciduous forests, and it generally does not enter extensive coniferous forests because of lack of ground cover. Food and Feeding. Diet of the Amur Hedgehog primarily contains invertebrates such as beetles, insect larvae, slugs, worms, and spiders. It also eats fungi, fruits, small vertebrates including frogs, snakes, lizards, mice, young birds, eggs, and carrion, and it scavenges on food discarded by humans. In northern China, almost 95% of dietary items were fly larvae; other invertebrates were earwigs, mole crickets, and beetles. One stomach was full ofjujube dates (Zizyphus jujuba, Rhamnaceae). Breeding. Amur Hedgehogs have 1-2 litters/year, with 4-6 young/litter. Activity patterns. The Amur Hedgehog is primarily active at night and rests under piles of brush by day orin leaf nests in rocky crevices and burrows. Usual gaitis slow, rolling walk, but it can run rapidly. It is a good swimmer and climber. It hibernates in winter, entering torpor in October in China and emerging in spring. Movements, Home range and Social organization. Except during breeding season or when a female has young, the Amur Hedgehog is solitary. It probably makes use of nests that are aboveground or in burrows, depending upon circumstances. Typical defensive posture is to roll into a ball, protecting abdomen, face, and limbs with sharp spines. It has been kept as a pet because it adjusts readily to captivity and becomes quite docile. Status and Conservation. Classified as Least Concern on The IUCN Red Lust. Bibliography. Beer (2003), Corbet (1988), Eo Kyung-Yeon et al. (2015), Harrison Matthews (1952), Hutterer (2005a), Liu Chengchao (1937), Nowak (1999), Reeve (1994), Smith & Yan Xie (2008, 2013), Stone (1995b)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 320-321, DOI: 10.5281/zenodo.6639332

4. Lowland Streaked Tenrec Hemacenteles semispinosus French: Tenrec rayé / German: Eigentlicher Streifentenrek / Spanish: Tenrec listado de tierras bajas Taxonomy. Ericulus semispinosus G. Cuvier, 1798, “Madagascar.” Hemicentetes semispinosus is distinguished from H. nigripes by craniodental characteristics and external appearance. They are sympatric in some localities. Monotypic. Distribution. Northern and Central highlands and E Madagascar. Descriptive notes. Head-body 104-176 mm, ear 12-20 mm, hindfoot 19-30 mm; weight 76-108 g. The Lowland Streaked Tenrec has no external tail; dorsal pelage is spinous, with sparse underfur and long, sparse guard hairs on dorsal surface. Base color is black, with broad lightcolored stripe on midline of head; conspicuous light-colored fan of spines occurs on top of head and nape of neck; light-colored longitudinal dorsal and lateral stripes on body are variably distinctive; and a few broad light-colored spines are scattered among dark spines within dark areas of dorsum. Contrasting light areas of body show individual variation from reddish buff to orange and yellow. Stout spines of stridulating organ in posterior mid-dorsal region are conspicuous reddish buff. Ventral pelage is bright reddish buffy brown or yellow, with mixture of sparse soft bristly hairs and spines. Head and limbs are covered with sparse but bristly fur. Large naked rhinarium of the Lowland Streaked Tenrec is covered with oval scales, and ears are moderately prominent. Forefeet are broad; first and fifth digits just reach to orjust beyond base of proximal digit; second, third, and fourth digits are long, with long, shallowly curved, stout claws. Hindfeet are moderately broad, with long claws on middle three digits. Dental formulais13/3,C1/1,P 3/3, M 3/3 (x2) = 40. Habitat. Humid forests; open woodland; degraded, agricultural and urban areas from near sea level to elevations of ¢.2050 m. Food and Feeding. [.owland Streaked Tenrecs mainly eat species of Annelida and Orthoptera and some coleopteran larvae. Feeding occurs during the day and in evening and early morning. Breeding. Gestation is ¢.58 days. Near Périnet (Analazamaotra Special Reserve), litters had 5-8 young but might be larger in captivity. Wild-caught females were pregnant with seven embryos in May, three embryos in July, and seven embryos in October. In a single captive litter of seven young, eyes opened at 7-10 days old, and solid food was eaten at c.3 weeks old. Young become sexually mature at 35-40 days old, and breeding can occur year-round under favorable conditions. Activity patterns. The Lowland Streaked Tenrec is active day and night. It excavates shallow burrows where it spends rest periods. In the wild, it has been seen between 13:00 h and 14:00 h and between 18:00 h and 20:15 h. A captive female and her five young had a major activity peak at midday, another major peak between 18:00 h and 19:00 h, and an extended peak between 04:00 h and 06:00 h; they were inactive in the afternoon. The Lowland Streaked Tenrec is apparently active year-round butless so in May-October. Individuals found at Ivohibe Special Reserve in August-September were very sluggish and cold to the touch after cold nights butstill with food in their intestines, indicating daily torpidity and partial seasonal torpidity. The Lowland Streaked Tenrec 1s a facultative hibernator; under favorable conditions, it can remain active year-round, although it still shows a reduction in activity during austral winter. It is semi-fossorial. Movements, Home range and Social organization. The Lowland Streaked Tenrec can be solitary but usually occurs in small family units or extended family groups of up to three generations that forage together. A single individual or a small family unit of an adult male and female and their young occupy a relatively simple shallow burrow, but a large family group or colonies of 20 or more individuals inhabit a relatively complex burrow system. Spines ofstridulating organs rub together as individuals move, and individuals probably can hear sounds produced by stridulating organs up to 10 m, helping to maintain contact among members of a group. In captivity, Lowland Streaked Tenrecs also produce tongue and lip clicks in experimental testing and when exploring an unfamiliar area, which are believed to be involved in echolocation. Predators include Fosas (Cryptoproctaferox), Spotted Fanalokas (Fossafossana), Ring-tailed Vontsiras (Galidia elegans), and large snakes. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Lowland Streaked Tenrec has a wide distribution and can be locally abundant, although overall population trends are unknown. It apparently can adapt to anthropogenic habitats, and it is hunted for food in some areas. Bibliography. Asher & Hofreiter (2006), Butler (1941), Dobson (1882), Eisenberg & Gould (1970), Everson et al. (2016), Goodman, Jenkins & Rakotondravony (2000), Goodman, Soarimalala et al. (2013), Gould (1965), Mar shall & Eisenberg (1996), Olson & Goodman (2003), Poux, Madsen, Glos et al. (2008), Poux, Madsen, Marquard et al. (2005), Rand (1935), Salton & Sargis (2008a, 2008b, 2009), Schmid & Stephenson (2003), Soarimalala & Goodman (2003, 2011), Stephenson (2003b, 2007), Stephenson & Racey (1994), Stephenson, Racey & Rakotondraparany (1994), Stephenson, Soarimalala & Goodman (2016e), Wever & Herman (1968)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 164-165, DOI: 10.5281/zenodo.6808230

22. Moonrat Echinosorex gymnurus French: Grand Gymnure / German: Grol3er Rattenigel / Spanish: Gimnuro Other common names: Malayan Moonrat, Raffles’ Gymnure Taxonomy. Viverra gymnura Raffles, 1822, “Sumatra.” Restricted by G. B. Corbet in 1988 to “Bencoolen, Sumatra,” Indonesia. Widely used specific name gymnura and subspecific name alba have been changed for gender agreement. Two subspecies recognized. Subspecies and Distribution. E.g.gymnurusRaffles,1822—MalayPeninsulaandSumatra,includingTebingtinggi1. E. g. albus Giebel, 1863 — Borneo, including Labuan I. Descriptive notes. Head-body 255-460 mm, tail 165-300 mm, ear 45-50 mm, hindfoot 55-75 mm; weight 0.4-2 kg. Females are larger than males. The Moonrat is the largest extant erinaceid. Shaggy pelage consists of short and thick underfur, covered by dense layer of long and coarse hairs. Coloris variable and primarily black on head and shoulders, pale on face and neck, and whitish on distal part of tail, or pelage generally is white with sparse scattering of black hairs. Face usually is marked with black spots or stripes near eyes. White forms (not albino) also occur. Body is exceedingly narrow, perhaps an adaptation for seeking food in narrow crevices. Underside of long, and mobile nose is grooved from its tip to a point between upper incisors. Scantily haired tail reveals that scales are arranged in rows around tail, except near base where they are arranged diagonally. Terminal one-third oftail is compressed. There is geographical variation in size and color. In Thailand, individuals in northern areas are smaller than those near the Equator. Individuals from western Borneo have greater proportion of black hairs than those from the east; intermediate color has been noted for Brunei. Dental formulais13/3, C1/1,P4/4,M 3/3 (x2) = 44. Habitat. Tropical lowlands and foothills, including primary, secondary, logged, and mangrove forests, rubber plantations, cultivated areas, and especially areas near streams and mangrove swamps. Food and Feeding. The Moonrat forages extensively on the forest floor, where it primarily eats earthworms, beetles, cockroaches, insect larvae, millipedes, centipedes, spiders, and scorpions. It might be partially aquatic and often enters water to hunt for frogs, fish, crustaceans, mollusks, and insects. It also takes other invertebrates on land and at water’s edge. It eats fruits of cultivated oil palms. Breeding. The Moonrat presumably is polygynous and breeds throughout the year. Pregnancies have been recorded in May, June, September, and November. Gestation lasts ¢.35-40 days. Litters have two young (range 1-2). Sizes of neonates are head-body 80-81 mm, tail 25-26 mm, hindfoot 14 mm, and weight 14-15 g. Females have 1-2 litters/year, usually two. Maximum reported lifespan in captivity is 55 months. Activity patterns. The Moonratis terrestrial and usually nocturnal but can be active day or night. Day-resting sites are under roots of trees and in hollow logs, empty holes, burrows, rocky crevices, or nipa palms (Nypa fruticans, Arecaceae) in mangrove swamps. The Moonrat constructs a nest of leaves or other materials. It emerges from its nest about one hour after sunset and usually is active for 45-90 minutes before returning to its nest. The Moonrat walks and runs with its body high off the ground and its snout raised. Snout explores surroundings by probing, similar to shrews (Soricidae). Local weather conditions, especially heavy rainfall, can increase foraging activities, probably to take advantage of emerging earthworms. Movements, Home range and Social organization. Moonrats are generally solitary and highly intolerant and aggressive toward conspecifics. Captive individuals emit hiss-puffs and low roars during encounters. The Moonrat engages in scent-marking behavior using strong, characteristic odor that emanates from two small glands near its anus; odor has been described as resembling rotten onions, ammonia, and stale sweat. It uses this potent scent to mark home ranges and deter predators; captive individuals use the odor to mark openings of nest boxes. On Borneo, Moonrats often forage in pairs at night along small muddy streams and damp areas in lowland primary and logged forests. Strange sounds cause a Moonrat to react suddenly and seek cover. When frightened, it freezes into hunched, open-mouthed postures, uttering a loud hissing cough. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Moonrat has a wide distribution and is abundant in suitable habitat. It occurs in many protected areas, including Kubah National Park and Kuching Wetlands National Park on Borneo. Bibliography. Beer (2003), Corbet (1988), Curtis (2003), Davis (1962), Frost et al. (1991), Gould (1978), Hayssen et al. (1993), Heaney & Morgan (1982), Hutterer (2005a), Jenkins & Robinson (2002), Lekagul & McNeely (1988), Lim (1967a), Medway (1983), Nowak (1999), Payne & Francis (2007), Phillipps & Phillipps (2016), Ralls (1976), Reeve (1994), Stone (1995b), Whittow et al. (1977)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 328-329, DOI: 10.5281/zenodo.6639332

26. Dinagat Gymnure Podogymnura aureospinula French: Gymnure de Dinagat / German: Dinagat-Rattenigel / Spanish: Gimnuro de Dinagat Other common names: Dinagat Moonrat, Dinagat Wood Shrew, Golden-spined Moonrat, Spiny Moonrat Taxonomy. Podogymnura aureospinula Heaney & Morgan, 1982, “Plaridel, Albor Municipality, Dinagat Island, Surigao del Norte Province, Republic of the Philip- pines.” Interpreted by D. R. Frost and colleagues in 1991 as “10°12°N, 125°35’E.” This species is monotypic. Distribution. Dinagat and Bucas Grande Is, S Philippines. Descriptive notes. Head-body 190-211 mm, tail 59-73 mm, ear 22-26 mm, hindfoot 39-42 mm. No specific data are available for body weight. The Dinagat Gymnure has spiny pelage and is second in size only to the Moonrat in the subfamily Galericinae. Dorsal pelage is short, spinous, golden-brown overall, and composed of three types of hairs: slate-gray underfur ¢.5 mm long; stiff, bristly, or spiny hairs c.15 mm long with black bases, others are golden-yellow, and many have black tips; and black, spiny hairs, c¢.12 mm long. Golden spines are twice as abundant as black hairs in middle of back. Black spines are densest at mid-dorsum, decreasing in abundance laterally, and disappearing on sides. Black-tipped golden spines are especially common on mid-dorsum and disappear on sides. Only golden spines are present on sides. Overall color of dorsum is golden-brown, with black spines and black-tipped golden spines, adding black speckling. Ventral pelage lacks spines and grades evenly from dorsal color to brownish gray over most of venter. Ventral hairs are of two types: soft, gray underfur c.5 mm long and slightly coarser guard hairs ¢.9 mm long, gray at bases, and tipped with pale brown. Pelage of rostrum and around eyes is short, dense, and spiny. Upper and lowerlips are covered in short, moderately dense, white, or pale brown fur. Vibrissae are dark at bases, very pale for most of their lengths, and up to 55 mm long. Rhinarium is long, naked, and distinctly bilobed, with nostrils opening laterally. Ears are relatively large and appear naked, but they have sparse covering of extremely short, white hairs. Forefeet and hindfeet have moderate coverings of short, white, or pale brown hairs dorsally and are nearly naked ventrally. Hindlegs appear almost naked distal to knee. Dorsal base oftail is sparsely furred, nearly naked, and 15 mm in diameter. Tail has sparse covering of short hairs. Dental formula is I 3/3, Cl1l/1,P3/3,M 3/3 (x2) =40. Habitat. Primary and secondary forests. The Dinagat Gymnure is known from a logged area in rolling low hills where there were many patches of remnant dipterocarp forests. These forests contained at least twelve families of plants including Dipterocarpaceae, which dominated the habitat with Anisoptera, Dipterocarpus, Hopea, and Shorea. Undergrowth was mainly rattans (Calamoideae) and ferns. Food and Feeding. Diet of the Dinagat Gymnure contains invertebrates. Breeding. No information. Activity patterns. The Dinagat Gymnure is nocturnal. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Endangered on The IUCN Red List. The Dinagat Gymnure is restricted to patches of forested habitats that are being destroyed by logging and slash-and-burn agriculture. Bibliography. Beer (2003), Frost et al. (1991), Heaney & Morgan (1982), Heaney & Rabor (1982), Heaney, Balete et al. (1998), Heaney, Dolar et al. (2010), Hutterer (2005a), Jenkins & Robinson (2002), Nowak (1999), duPont & Rabor (1973), Reeve (1994), Stone (1995b)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 330, DOI: 10.5281/zenodo.6639332

7. Somali Hedgehog Atelerix sclateri French: Hérisson de Somalie / German: Somalischer Igel / Spanish: Erizo de Somalia Other common names: Somalian Hedgehog Taxonomy. Erinaceus sclateri J. Anderson, 1895, “Taf, in Central Somaliland [= So- malia].” This species is monotypic. Distribution. Somalia. Descriptive notes. Head-body 206-263 mm, tail 14-20 mm, ear 22-27 mm, hindfoot 25-29 mm. No specific data are available for body weight. The Somali Hedgehog has five digits on hindfeet, and hallux is small. Dorsal pelage has spines that are dark gray-brown to grizzled black, smooth, finely striated, without papillae, up to 18 mm long, and 1 mm in diameter. Ventral pelage has non-spiny hairs that are white or buffy-white, often with dark gray-brown to grizzled black patches on posterior part of abdomen. Spiny dorsum and hairy venter are well delineated on lowerflanks. Forehead is white from cheek to cheek, there is narrow central parting of spines on crown of head, and black face mask has slight posterior extension of black onto cheeks. Snoutis short, and ears are small, broadly rounded, and shorter than spines between ears. Claws are moderately long, limbs are short and dark, and tail is short, with barely visible, pale hairs. Habitat. Maritime plain near hills in arid scrublands. Food and Feeding. The Somali Hedgehog eats invertebrates, small vertebrates, fruits, and fungi. Breeding. No information. Activity patterns. The Somali Hedgehog is nocturnal and terrestrial. Movements, Home range and Social organization. The Somali Hedgehog is solitary and probably self-anoint like other hedgehogs. Status and Conservation. Classified as Least Concern on The IUCN Red List. Bibliography. Anderson (1895), Beer (2003), Corbet (1988), Happold (2013c), Hutterer (2005a), Nowak (1999), Poduschka (1990), Reeve (1994)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 322, DOI: 10.5281/zenodo.6639332

24. Hainan Gymnure Neohylomys hainanensis French: Gymnure de Hainan / German: Hainan-Rattenigel / Spanish: Gimnuro de Hainan Other common names: Hainan Moonrat Taxonomy. Neohylomys hainanensis T. H. Shaw & Wong Song, 1959, “Pai-sa Hsian, Hainan Island.” Interpreted by D. R. Frost and colleagues in 1991 as “Baisha Xian, which is an administrative unit at 19°13’N, 109°26’E,” China. This species is monotypic. Distribution. Hainan I, China. Descriptive notes. Head-body 120-147 mm, tail 36-43 mm, ear 16-22 mm, hindfoot 24-29 mm; weight 50-70 g. Head of the Hainan Gymnureis blackish gray, mixed with brown. Back is rusty brown, mixed with gray. Long, black stripe on middle of back is darker on head and anterior part of body and paler toward posterior part of body. Sides are washed with pale olive-yellow, and underparts are pale gray or yellowish white. Ears, feet, and tail are almost naked, with minute scattered short hairs. Whiskers are black and up to 35 mm long. Dental formulais13/3,C1/1,P4/3,M 3/3 (x2) = 42. Chromosomal complement has 2n = 32 and FN = 64. Habitat. Damp forests including tropical rainforests and subtropical evergreen forests. Food and Feeding. Diet of the Hainan Gymnure includes invertebrates and plant material. Breeding. No information. Activity patterns. The Hainan Gymnure is nocturnal and terrestrial. It spends most of its time in subterranean burrows. Its cylindrical body, shorttail, and less-developed claws are adaptations for a fossoriallifestyle. Movements, Home range and Social organization. Hainan Gymnures are solitary. Status and Conservation. Classified as Endangered on The IUCN Red List. Extent of occurrence of the Hainan Gymnureis less than 5000 km?®. It is only known from Hainan Island, where remaining patches of evergreen forests are shrinking due to harvesting of timber and expansion of agriculture. Bibliography. Beer (2003), Bhatnagar & EI-Azawi (1978), Corbet (1988), Frost et al. (1991), Hubner et al. (1991), Hutterer (2005a), Jenkins & Robinson (2002), Li Yuchun et al. (2008), Nowak (1999), Reumer & Meylan (1986), Shaw & Wong Song (1959), Smith & Yan Xie (2013), Stone (1995b), Zima & Kral (1984).

14. Indian Long-eared Hedgehog Hemiechinus collaris French: Hérisson a collier / German: Indischer Langohrigel / Spanish: Erizo de orejas largas de India Other common names: Collared Hedgehog, Hardwicke's Hedgehog, Long-eared Desert Hedgehog Taxonomy. Erinaceus collaris Gray, 1830, “Doab.” Restricted by R. C. Wroughton in 1910 to “area between the rivers Jumna and Ganges,” India. This species is monotypic. Distribution. NW, EC & SE Pakistan and NW & NC India. Descriptive notes. Head-body 140-180 mm, tail 23 mm, ear 32-38 mm; weight 257-306 g. No specific data are available for hindfoot measurements. The Indian Long-eared Hedgehog exhibits a tendency toward melanism. It does not have central parting of spines on forehead or face mask. Snout is long and pointed. Legs are long, forefeet have curved claws, and hallux on hindfootis well developed. Long ears probably are adaptations for thermoregulation in its desert environment. Ears usually remain folded back over spines on crown, even when an individualis unrolled. Dorsal pelage has spines that are 17-30 mm long in adults. Spines appear blackish brown, usually banded with dark brown and white, pale at bases, and spines are black on their distal one-half including tips; overall effect is blackish brown appearance. Ventral pelage hassoft fur of variable color from whitish to blackish. Some individuals have collar of white hairs from under chin to ears. Stumpytail and abdomen are sparsely covered with blackish brown hair, face usually is paler and has dark brown or grayish black hair, and ears and upper parts of crown have gray or whitish hairs. Dental formula is I 3/2, C1/1,P 3/2, M 3/3 (x2) = 36. Karyotype is 2n = 48. Habitat. Various habitats including lowland areas of the Indus River, semiarid grasslands, plains, scrublands, deserts, dry steppes, and especially numerous in irrigated agricultural lands. The Indian Long-eared Hedgehog tends to avoid hot, dry deserts. Food and Feeding. The Indian Long-eared Hedgehog is omnivorous and will grab and eat any prey that it can overcome. When foraging,it takes short, rapid steps and appears to depend primarily on scent to locate food. It sniffs with its snout in roots of grass or leaflitter in search of insect larvae. Venomous snakes are grabbed anywhere on the body, and with spines erect and head lowered,it keepsits face partially concealed so that the reptile strikes against spines and injures itself. When an Indian Longeared Hedgehog seizes a snake or limb of a large animal, it will roll itself into a ball and begin chewing on the held portion. Diet primarily contains insects, especially dung beetles and orthopterans (mole crickets, crickets, and grasshoppers), but other insects and arthropods; earthworms; small vertebrates including frogs, toads, lizards, snakes, eggs and nestlings of ground-dwelling birds, and small mammals; carrion; and fruit also are eaten. Cannibalism is common. Females will eat their offspring, and males will attack and eat any young they encounter. Breeding. In India, breeding occurs in June-September, which is between extreme heat in May-June and cooler weather in December—January. Young benefit from increased availability of food following monsoons in July-August. Similarly, in Pakistan, mostlitters are born August-September at the height of the monsoon when insect prey is most abundant. The male copulates by standing almost vertically on his hindlegs, while the female lies on her abdomen with hindlegs extended on the ground. Gestation probably lasts 30-40 days. Young are born in burrows. If natal burrow is disturbed, the mother will abandon it. Young are transported by the mother only at night. To carry them, she grasps loose skin on the neck or side with her mouth. Litters average 2-3 young (range is 1-6). A litter of four young was born over a five-hour period. Neonates have closed eyes and seem to locate their mother by smell. Nursing occurs with the mother lying on her side, and infants knead mammae with their forefeet and suck loudly. Neonates average 8:3 g, are 47 mm long, and are nearly hairless, except for scattered spines that are 2 mm long, whitish, and soft. Within c.5 hours of birth spines have grown to ¢.8 mm and begin to darken but are still soft. When c.5 days old, numerous pigmented spines are visible, and by the time young are two weeks old, their back is thickly covered with spines. Young do not leave nest burrows until! four weeks old, after which they accompany their mothers on foraging trips. It is not known at what age young become independent of their mothers but probably at 6-7 weeks. Activity patterns. In summer,the Indian Long-eared Hedgehog escapes intense desert heat by being strictly nocturnal in foraging activity. It emerges at dusk and forages for 5-6 hours. When food is plentiful, it returns to its burrow by about midnight and does not emerge again until the following evening. It is an active digger and excavatesits own burrows. Opening of burrow is entirely or partially concealed under a bush, and if the ground is hard,it can be short, extending no more than 30 cm. In softsandy soil, the burrow can be 150 cm deep. Burrow terminates in slightly wider chamber that is unlined. An individual that descends head first into and out ofits burrow can turn around inside the small chamber. Burrow usually slopes down to ¢.30-35 cm below the surface and will be used by the same individual for an entire year. Movements, Home range and Social organization. The Indian Long-eared Hedgehog is solitary and unsocial, and it never shares a burrow with conspecifics. It is relatively silent, and the only vocalization known is a snake-like hiss that is emitted when disturbed in its burrow or threatened by a predator. This hiss generally is accompanied by sudden upward jerking of the body that can be a painful experience for a predator. The Indian Long-eared Hedgehog exhibits self-anointing behavior that begins by chewing on vegetable matter, generally grass roots, which produces copious frothy saliva. It then contorts itself so that its muzzle is twisted around to its back and flanks, and the liquid vegetable paste or frothy saliva is plastered onto the skin without injury to the tongue. Self-anointing usually is stimulated by some strong smell. Reasons for this behavior are unknown. Anointing might be a way of grooming otherwise inaccessible skin or helping to remove dried skin because the paste eventually dries and flakes off, or it might be a sexual signal because it occurs only in adults during the breeding season. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Indian Long-eared Hedgehog does not come into conflict with humans because it generally lives on edges of cultivation or in deserts. Bibliography. Asawa & Mathur (1981), Beer (2003), Corbet (1988), Goyal & Mathur (1974), Habibi (2004), Herter (1965), Hutterer (2005a), Kanwar et al. (1980), Krishna & Prakash (1955, 1956), Maheshwari (1982, 1984), McCann (1937), Molur et al. (2005), Nowak (1999), Osborn & Helmy (1980), Pandey & Munshi (1987), Prakash (1955a, 1955b, 1959, 1960), Reeve (1994), Roberts (1997), Sharma, G.P. et al. (1975), Sharma, S.K. (1991, 1996), Stone (1995b), Walton & Walton (1973), Wroughton (1910)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 326, DOI: 10.5281/zenodo.6639332

6 . Four-toed Hedgehog Atelerix albiventris French: Hérisson a ventre blanc / German: \WeiRbauchigel / Spanish: Erizo africano de vientre blanco Other common names: African Pygmy Hedgehog, Central African Hedgehog, White-bellied Hedgehog Taxonomy. Erinaceus albwventris Wagner, 1841, type locality not given. Restricted by J. Anderson and W. E. de Winton in 1902 to “in all probability, from Senegambia Confederation [= Senegal + The Gambia],” West Africa. This species is monotypic. Distribution. Senegal E to Eritrea, Djibouti, and Somalia and S through East Africa to Zambezi River. Descriptive notes. Head-body 140-258 mm,tail 9-25 mm, ear 15-28 mm, hindfoot 26-34 mm; weight 250-680 g, but up to 900 g in captivity. The Four-toed Hedgehog is small, with hallux (first digit) absent or very small; dorsal pelage has 15-17mm spines; there is narrow central parting of spines on crown of head; ventral pelage has non-spiny hairs; limbs are short and hairy; and tail is short and thinly haired. Back is speckled gray-brown, black, and white; eyes are black; ears are small, dark, rounded, and shorter than adjacent spines; muzzle is brownish to black; and face, legs, and underside are covered with white hairs. Some individuals have black face mask. Dental formulais13/2,C1/1,P3/2,M 3/3 (x2) = 36. Karyotype is 2n = 48. Habitat. Grasslands, thickets, scrublands, shrublands, woodlands, agricultural lands, hills, mountains, and frequently suburban areas occupying gardens and buildings. The Four-toed Hedgehog lives in areas with dry soils and is absent from deserts, marshes, and dense forests. It seems to be drawn to relatively open, dry, or seasonal habitats, with sparse or patchy grasses, especially overgrazed areas with dense populations of ungulates where there is an abundance of trampled herbage and dung to support termites and other insects. Food and Feeding. The Four-toed Hedgehog is omnivorous. It will eat fungi and plant material such as fallen fruits, roots, and groundnuts, but it primarily preys upon invertebrates, especially termites, beetles, earthworms, millipedes, ants, grasshoppers, slugs, snails, and crabs. It also eats small vertebrates including snakes, lizards, frogs, and young and eggs of ground-nesting birds. It usually forages alone and hunts mainly by scent. It will dig for prey in soft, loose soils. Hearing and sight also are involved with searching for food. Immobile foods are toyed with before being ingested, and mobile prey is snapped up and chewed noisily. Lizards and mice are seized and shaken to death. Snakes are approached with caution and are bitten severely and repeatedly while the Four-toed Hedgehog protects its face with spines to avoid being struck;it will continue the attack until it has broken the spine or eviscerated the snake. It readily consumes carrion and will scavenge road kills at night. Breeding. Times of reproductive activity of the Fourtoed Hedgehog vary in different parts of its distribution, but it is sexually active throughout the year in central and northern Africa. Farther south in Zambia and Zimbabwe, it breeds seasonally. Courtship is ritualistic. When a female is in search of a mate, she calls with a whine. Males serenade females during courtship with bird-like call of repeated squeaks that vary greatly from almost a whistle to a coarse squawk. Males court females by persistently following them. Male walks around a female in estrus with his snout pointed toward her and puffing; this may last for several days. When the male approaches, the female reacts aggressively by raising her spines and snorting. She respondsto a serenading male by vigorously butting him as she hisses and snorts, and she rejects him by running away, by pushing him on his flank with spines on her head, and with other evasive moves. Such behavior can last for hours. Copulation occurs when the male mounts the female from behind after she has flattened her spines, lowered her abdomen to the ground, and pressed her hindquarters outward. The male has an especially long penis, which might be because spines on the rump of the female prevent the male from getting too close to her during copulation. Gestation lasts 35-37 days. Young are born during any month in central and northern Africa, but in southern areas, young are born during warm and wet months, mainly October—March. In Zambia and Zimbabwe, young are born in November, ¢.2 months after females emerge from estivation. Nests are in well-hidden cavities, lined with vegetative material. After parturition, the female licks the young, eats the afterbirth, and places young on her abdomen to nurse. Young vigorously search for a nipple; after nursing begins, swallowing noises and motion of the throat are apparent. Litters usually have 3-5 young (range 1-10). Young are altricial and nearly hairless, eyes and ears are closed, they weigh ¢.10 g (range 8-13 g), they are 20-26 mm long, and their spines are pliable for the first few daysafter birth. Neonatal mortality is high due to desertion and cannibalism by mothers. Young grow rapidly; they weigh 25 g by day 7, 50 g by day 13, 70-75 g by day 20, 110-120 g by day 28, 130-150 g by day 34, and 170-190 g by day 40. Young huddle together, especially during periods of inactivity. Legs are thin and fragile, and feet are formed with nails intact. Young exhibit boxing, hissing, self-anointing, and anti-predator behaviors before their eyes open. Eyes open at 14-15 days. Deciduous teeth erupt at c.3 weeks, permanent teeth at 7-9 weeks, and consumption ofsolid food begins at ¢.24 days. Young typically are weaned by 4-6 weeks of age when they weigh 170-195 g; then, they begin to accompany their mother and typically leave her by c.7 weeks of age. In captivity, sexual maturity is attained as early as 61 days, but in the wild,it is reached at c.1 year of age. Females can give birth several times each year. There is no record of a reproductively active, captive-born female more than two years of age. Only one wild-caught female that was in the lab for 33 months was able to reproduce longer; females in the wild probably are reproductively active for an even shorter time. Activity patterns. The Four-toed Hedgehog is predominantly nocturnal, but it can be both crepuscular and nocturnalto avoid daytime heat and desiccation. In captivity, two peaks of activity are evident: 21:00-24:00 h and ¢.03:00 h. It rests by day curled up in a ball or reclined full-length under rocks, logs, or matted grass or in leaflitter, termitarium, rocky crevice, or hole in the ground. Such refuges change daily unless they are being used to rear young or as a hibernation site. During warm rainy season when food is abundant, the Four-toed Hedgehog gains weight and enters torpor when weather is cool and dry. In southern Africa, hibernation takes place in June-September, with individuals remaining torpid for up to six weeks and emerging during warm intervals. In some parts ofits distribution,it probably does not hibernate. The Four-toed Hedgehog can climb and swim; it typically moves slowly butis capable of bursts of speed. Movements, Home range and Social organization. Four-toed Hedgehogs are common in suitable habitats. Individuals are observed more often at beginning of wet season. Home range is small and usually within 200-300 m of a burrow or day-resting site. Four-toed Hedgehogs are generally solitary, except during courtship or when a female has young. When two individuals meet, there is much growling, hissing, snorting, spitting, and butting of heads. Main defense mechanism used against predators is to contract a series of muscles, rolling into a compact ball shape, with head, limbs, and underparts protected by sharp spines. Feces have a strong odor and might be a means of advertising presence of an individual. Self-anointing behavior occurs when the Fourtoed Hedgehog comes into contact with an irritating substance. It takes the substance into its mouth and creates a mixture with saliva that is rubbed onto spines. Reasons for self-anointing are unknown, but it might be used to attract mates or byjuveniles to get attention from their mothers. Self-anointing is innate becauseit is performed before young open their eyes. Several vocalizations are emitted that enable communication between individuals. Five types of audible sounds have been recorded. Twitter is a quiet sound emitted through the closed mouth, often accompanied by sniffing and usually associated with unfamiliar situations; each twitter is of short duration (5-40 ms), and several are emitted in pulses lasting for several seconds. Hiss is a short noise of lower pitch than the twitter and emitted during stressful situations. Snort is similar to the hiss but louder and emitted when severely stressed or attacked, often repeated rapidly. Scream is a rare sound, emitted under extreme stress. Serenade is a series of low-pitched sounds, ranging from a whistle to a coarse squawk and emitted by males during courtship behavior. Status and Conservation. Classified as Least Concern on The IUCN Red List. In eastern Africa, many superstitions surround the Four-toed Hedgehog. Its skin or spines often are used as fertility charms. In Uganda and Kenya, a bumper harvestis believed to be obtained by placing skin of a hedgehog on seeds before sowing, and for maximum yields of cotton, its skin should be burned on the cotton field. Humans eatit as bushmeat. In Nigeria, holes where it lives are located, and they are captured by digging, pouring water into the hole, or filling the hole with smoke. When the individual comes out,itis killed for food. The Four-toed Hedgehog is vulnerable to collisions with motor vehicles, but its overall population faces no major conservation threats. Bibliography. Allen (1922), Anderson & de Winton (1902), Ansell & Dowsett (1988), Beer (2003), Booth (1960), Brecht et al. (1997), Brockie (1976), Brodie Ill et al. (1982), Brodie Jr (1977), Cansdale (1960), Catania (2005), Catania et al. (2000), Fairley et al. (1999), Graffam et al. (1998), Gregory (1975, 1976), Haltenorth & Diller (1988), Happold (1969, 1987 2013a), Herter (1965), Hoefer (1994), Hibner et al. (1991), Hutterer (2005a), Kingdon (1974a, 1997), Meritt (1981), Nowak (1999), Ofori et al. (2015), Okaeme & Osakwe (1988), Okaeme et al. (1988), Reeve (1994), Santana et al. (2010), Smith (1992, 1995), Symonds (1999), Watson (1951)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 321-322, DOI: 10.5281/zenodo.6639332

20. Dwarf Gymnure Hylomys parvus French: Gymnure nain / German: Zwergrattenigel / Spanish: Gimnuro enano Other common names: Dwarf Moonrat Taxonomy. Hylomys parvus H. C. Robinson & Kloss, 1916, “Korinchi Peak, 10,000’ [= 3048 m], West Sumatra,” Indonesia. This species is monotypic. Distribution. Restricted to highlands of Mt Kerinci, WC Sumatra. Descriptive notes. Head-body 100-127 mm, tail 15-32 mm, ear 17-19 mm, hindfoot 22-25 mm. No specific data are available for body weight. The Dwarf Gymnure has notched space between premaxillary tips, soft-textured pelage, and delicate skull and dentition. Dental formula is I 3/3,C1/1,P4/4,M 3/3 (x2) = 44. Habitat. Restricted to high-elevation moss forest covering peak of Mount Kerinci, Sumatra, at elevations of 2000-3000 m. Food and Feeding. No information. Breeding. No information. Activity patterns. There is no specific information for this species, but the Dwarf Gymnure 1s terrestrial. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Vulnerable on The IUCN Red List. Extent of occurrence of the Dwarf Gymnure is less than 20,000 km?, it is known from fewer than ten locations, and there is continuing decline in the extent and quality ofits habitat in the Sumatran Mountains. Bibliography. Hutterer (2005a), Jenkins & Robinson (2002), Ruedi & Fumagalli (1996), Ruedi et al. (1994), Stone (1995b).

2. Northern White-breasted Hedgehog Erimaceus roumanicus French: Hérisson des Balkans / German: Nordlicher WeilRbrustigel / Spanish: Erizo de los Balcanes Other common names: Eastern European Hedgehog Taxonomy. Erinaceus europaeus roumanicus Barrett-Hamilton, 1900, “Gageni, Roumania [= Romania].” Erinaceus roumanicus was first described as a subspecies of E. europaeus and then included as a subspecies of E. concolor, current genetic and morphologic data suggest that E. concolor and E. roumanicus are distinct species with parapatric distributions. Erinaceus roumanicus is sympatric with E. europaeus in parts of central Europe. Five subspecies recognized. Subspecies and Distribution. E.r.roumanicusBarrett-Hamilton,1900—C&EEurope(EofalinefromOderRivertoNAdriaticSea). E.r.bolkayiV.Martino,1930—Montenegroandadjacentcountries. E.r.drozdovskiiV.Martino&E.Martino,1933—Macedonia,EuropeanTurkey,andadjacentcountries. E.r.nesiotesBate,1906—CreteandIonianandAegeanIs,Greece. E. r. pallidus Ognev, 1928 — SW Russia S to N Caucasus and E to W Siberia. Distributions of most subspecies are not well known. Descriptive notes. Head—body 208-306 mm, tail 24-31 mm, ear 28-29 mm, hindfoot 40— 44 mm; weight 0.6-1.2 kg. As in other species of Erinaceus, the Northern White-breasted Hedgehog has faintly visible, central parting of spines on crown of head, smooth spines lacking papillae, and well-developed hallux on hindfeet. Color of underside is mix of dirty white and dirty brown. Size varies in a smooth cline across the distribution, with largest individuals in southern populations and smallest in northern areas. Dental formulais 13/2, C1/1,P 3/2, M 3/3 (x2) = 36. Chromosomal complement has 2n = 48 and FN = 94. Habitat. Farmland, parks, and gardens in rural and urban areas, scrubby habitats at edges of forests, and shrubby vegetation from sea level to elevations of at least 1400 m. Food and Feeding. There is no specific information available for this species, but diet and feeding behavior of the Northern White-breasted Hedgehog are probably similar to the West European Hedgehog (E. europaeus). Breeding. Female Northern White-breasted Hedgehogs can produce two litters peryear: one at the end of May and another at the end of August. Activity patterns. There is no specific information available for this species, but activity patterns of the Northern White-breasted Hedgehog probably are similar to those of the West European Hedgehog. Movements, Home range and Social organization. There is no specific information available, but movements, home range, and social organization probably are similar to those of the West European Hedgehog. Status and Conservation. Classified as Least Concern on The [UCN Red List. Many Northern White-breasted Hedgehogs are killed by collisions with cars, but this is unlikely to cause widespread population declines. It occurs in many protected areas throughoutits wide distribution. No specific conservation measures are necessary at present. Bibliography. Arslan et al. (2008), Barrett-Hamilton (1900), Bate (1906), Corbet (1988), Geisler & Gropp (1967), Gropp et al. (1969), Hutterer (2005a), Krystufek & Vohralik (2001), Krystufek, Tvrtkovié et al. (2009), Lapini (1999), Martino & Martino (1933), Masseti (2012), Ognev (1928), Pavlinov & Lissovsky (2012), Poduschka (1969), Reeve (1994), Suchentrunk et al. (1998)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 320, DOI: 10.5281/zenodo.6639332

12. Bare-bellied Hedgehog Paraechinus nudiventris French: Hérisson a ventre nu / German: Nacktbauchigel / Spanish: Erizo de vientre desnudo Other common names: Madras Hedgehog Taxonomy. Erinaceus nudiventris Horsfield, 1851, “Madras,” India. Interpreted by D. R. Frost and colleagues in 1991 as either “the city at 13°05’N, 80°17’E or the older name for the Tamil Nadu province,” India. This species is monotypic. Distribution. Andhra Pradesh, Tamil Nadu, and Kerala, S India. Descriptive notes. There are no specific measurements available. T. Horsfield in 1851 noted that size of the Bare-bellied Hedgehog is about one-half that of the Indian Longeared Hedgehog (Hemiechinus collaris). G. B. Corbet and J. E. Hill in 1992 noted that compared with the Indian Hedgehog, the Bare-bellied Hedgehog has shorter ears, brown collar, and very rudimentary third upper premolars (P?). Tail is short and concealed. Spines have a rough surface, they are strongly grooved along their lengths, and they are yellowish white at bases and tips, with blackish band in the middle. Head and ears are naked and sooty black throughout. Throat, neck to near ears, chest, and abdomen are covered with naked skin of a dirty chestnut-brown, changing to blackish on abdomen and dirty yellowish on sides. It has wide center parting of spines on crown of head and distinct white band across forehead. Ears protrude beyond adjacent spines. Hallux is somewhat reduced but consistently well developed. Habitat. Rocky and shrubby deserts and arid regions with Acacia (Fabaceae) shrubs and trees. Food and Feeding. Diet of the Bare-bellied Hedgehog consists primarily of insects but also small vertebrates, eggs of ground-nesting birds, and scorpions. Captive individuals sometimes eat their young. Breeding. The Bare-bellied Hedgehog probably produces only onelitter each year. Activity patterns. The Bare-bellied Hedgehog is terrestrial and excavates burrows under logs, leaves, and near rocks. Most of the day is spent sleeping, and it wakes up and begins searching for food at dusk. Movements, Home range and Social organization. Bare-bellied Hedgehogs are solitary and engage in self-anointing behavior. Captive individuals generally do not bother one another if fed well. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Barebellied Hedgehog is abundantin its restricted distribution and presumably has a large overall population. Increasing desertification is resulting in fragmentation of populations. Loss of habitats due to collection of fuel wood, logging, agriculture, and urbanization are the major conservation threats. Bibliography. Corbet (1988), Corbet & Hill (1992), Frost et al. (1991), Horsfield (1851), Hutterer (2005a), Marimuthu & Asokan (2014), Nowak (1999), Reeve (1994), Stone (1995b)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 324-325, DOI: 10.5281/zenodo.6639332

9. Desert Hedgehog Paraechinus aethiopicus French: Hérisson du désert / German: Athiopischer Igel / Spanish: Erizo del desierto Other common names: Ethiopian Hedgehog Taxonomy. FErinaceus aethiopicus Ehren- berg, 1833, “in desertis dongalanis habi- tat.” Interpreted by G. B. Corbet in 1988 as “Dongola Desert, Sudan” and restricted by D. R. Frost and colleagues in 1991 to “approximately at 19°22’N, 30°45’E,” northern Sudan. Paraechinus aethiopicus might be polytypic, but subspecific taxonomy requires reassessment. Monotypic. Distribution. Sahara Desert from Mauritania E to Egypt, Sudan, and Eritrea, WC Ethiopia (Awash), and deserts in Middle East (Syria, Iraq, Israel, Jordan, and Arabian Peninsula); insular populations on Djerba (Tunisia), Bahrain, and Tunb (Persian Gulf). Descriptive notes. Head—body 130-240 mm, tail 10-30 mm, ear 23-58 mm, hindfoot 27-39 mm; weight 285-700 g. Dorsal pelage of the Desert Hedgehog has dark spines that are banded, whitish at their bases, with two wide black bands separated by white band, and white tips. Surfaces of spines are rough, with numerous papillae and many longitudinal grooves. Ventral pelage is rather sparse, and hairs are mostly white but can be brown posteriorly or have brown patches on chest in some individuals. Ventral pelage varies geographically in color, which can be variable mix of white, brown, and black, but some individuals have entirely brown venter. Head has broad white forehead from cheek to cheek, wide central parting of spines on crown of head, and welldefined gray-black face mask on muzzle and around eyes that extends posteriorly on lower cheeks. Ears are large, dark, slightly pointed, and usually longer than adjacent spines. Limbs are rather long and dark brown. Feet are black, forefeet have five digits, hindfeet have five digits, and hallux can be slightly reduced. Tail is short, barely visible, and covered with small pale hairs. Dental formulaisI13/2,C1/1,P 2-3/2, M 3/3 (x2) = 34 or 36. Chromosomal complement has 2n = 48 and FN = 96. Habitat. Dry scrublands, dry steppes, stony plains, mountainous desert country, and more mesic habitats such as oases and vegetated riverine valleys in arid and semiarid regions in and around the Sahara and Arabian deserts. Food and Feeding. The Desert Hedgehog forages at night and primarily is insectivorous. Diet also includes scorpions, other invertebrates, and small vertebrates such as frogs, lizards, snakes, and eggs of ground-nesting birds. Breeding. Breeding of the Desert Hedgehog begins after hibernation has ended. Females can produce up to four litters each year, with peaks in February-March and June. It nests in clumps of vegetation or uses burrows under shrubs or among rocks. The Desert Hedgehog does not line its nest; a female gave birth to one young in a sand and peat hollow, making no use of hay, wood shavings, and leaves that were available. Gestation lasts 34-46 days. Litters have 2-7 young. Neonates weigh 6-9 g. Weight at ten days of age is 20 g, 20 days is 40 g, 30 days is 60 g, 40 days is 80 g, and 60 daysis 150 g. Rate of growth to six weeks is 1-7 g/day, and it is 3-5 g/day at 6-9 weeks old. Young begin rolling-up for the first time at 7-22 days old, eyes open at 21-22 days,thermoregulation is fully developed, solid foods are eaten at c.38-44 days, and weaning occurs at 40-58 days. Growth to weaning is longer than in other species of hedgehogs. Longevity might be 6-10 years. Activity patterns. The Desert Hedgehog is usually nocturnal, and it can excavate burrows that have many openings. In Egypt,it forages on the coastal plain at night. During day, it rests in crevices in cliffs, under clumps of vegetation, or in burrows. When nights are cold, and food is less abundant in winter,it enters torpor and is rarely seen. In Algeria, it is torpid in November—-March, but it usually emerges every few days to forage. During winter in Qatar, body temperature was higher than ambient temperature throughout the day, and the difference was more extreme during midday. Desert Hedgehogs bask in direct sunlight, which appears beneficial for thermoregulation in the desert where there is plenty of solar radiation. Movements, Home range and Social organization. The Desert Hedgehog is solitary and generally uncommon. Nevertheless,it can be more common in some localities because populations are scattered and isolated. It is relatively abundant in irrigated gardens and fields in parts of its distribution. Long limbs enable fast walking and trotting. One individual was followed at a steady 10 km/h for five minutes. The Desert Hedgehog can be rather sedentary, occupying a single burrow and the same home range all year—behaviors that could be construed as evidence of territoriality. It exhibits self-anointing behavior. When attacked,it rolls into a ball and makes a noise similar to the throbbing of a miniature engine. Periodically, it gives a sort of half-grunt, half-bark sound. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Desert Hedgehog is reported to be widespread species, with a presumed large population.It is believed to be reasonably tolerant of habitat modification and is not considered globally threatened. Nevertheless, increasing desertification in its distribution is resulting in fragmentation of populations. In Algeria, the Desert Hedgehog is hunted and eaten by humans. In Somaliland, humans are rather superstitious about hedgehogs, never harming them because they consider them stars that have fallen from the heavens. Bibliography. Abu Baker, Mohedano et al. (2016), Abu Baker, Reeve et al. (2016), Al-Saleh & Khan (1985), Amr (2000), Beer (2003), Bhatnagar & EI-Azawi (1978), Brodie (1977), Corbet (1988), Delany & Farook (1989), Drake-Brockman (1910), Eisentraut (1952), Frost et al. (1991), Happold (2013e), Harrison & Bates (1991), Hayssen et al. (1993), Heuglin (1861), Hufnagl (1972), Hutterer (2005a), Nader & Al-Safadi (1993), Nowak (1999), Osborn & Helmy (1980), Petter (1954), Qumsiyeh (1996), Reeve (1994), Sellami et al. (1989), Stone (1995b), Thomas (1919), Walton & Walton (1973), Yamaguchi et al. (2013)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 323-324, DOI: 10.5281/zenodo.6639332

1. West European Hedgehog Erinaceus europaeus French: Hérisson commun / German: Braunbrustigel / Spanish: Erizo europeo Other common names: Brown-breasted Hedgehog, Common Hedgehog, European Hedgehog, Northern Hedgehog Taxonomy. Erinaceus europaeus Linnaeus, 1758, “Europa.” Restricted by O. Thomas in 1911 to “Wamlingbo, S. Gothland Id.,” Sweden. Erinaceus europaeus is sympatric with FE. roumanicus in central Russia, Poland, Czech Republic, Slovakia, and the Balkan Peninsula. Monotypic. Distribution. W Europe, from British Is and Iberian Peninsula E to Fennoscandia and N & C European Russia, also on Corsica, Sardinia, Sicily, and many smaller islands. Introduced into New Zealand during second halfofthe 19" century and recently into Azores. Descriptive notes. Head-body 209-245 mm, tail 24-32 mm, ear 27-28 mm, hindfoot 43-50 mm; weight 526-556 g (some individuals weigh up to 1-2 kg). The West European Hedgehog has well-developed hallux on hindfeet, short tail, and relatively short ears. Pelage, except non-spiny hairs on face, legs, and underparts, has dense smooth spines, lacking papillae, banded yellow and brown, and with pale tips. As on other species of Erinaceus, the West European Hedgehog has narrow spine-free parting on crown ofhead. It can be distinguished from the Southern White-breasted Hedgehog (E. concolor) by absence of white breast patch. In southern Spain, the West European Hedgehog is pale, with many entirely white spines, and can be distinguished from the similar North African Hedgehog (Atelerix algirus) by narrow spine-free parting on crown of head. Albinos are relatively common, leucistic individuals are rare, and melanistic individuals are very rare. Dental formulais 13/2, C1/1,P 3/2,M 3/3 (x2) = 36. Chromosomal complement has 2n = 48 and FN = 92. Habitat. Forests with ground vegetation and adequate food and grasslands especially adjacent to deciduous woodlands, hedgerows, or scrublands. In the Alps, the West European Hedgehog occurs up to elevations of 2000 m but does not live above tree line. It generally does not enter extensive coniferous forests because oflack of ground cover. It is common in parks, gardens, and golf courses, and it seems to favor edge habitats between open and more densely vegetated areas. Rocky outcrops and piles of rocks can serve as shelters. Populations in cultivated areas are small or lacking where removal of hedgerows and use ofpesticides have reduced cover and amounts ofinvertebrate prey. It may move ontoirrigatedfields in dry years. Food and Feeding. The West European Hedgehog is omnivorous but primarily eats invertebrates such as beetles, larvae, slugs, worms, andspiders. It also eats fruits, fungi, and small vertebrates including frogs, snakes, lizards, young birds, eggs, mice, and carrion. It has been reported to congregate around gardens and vineyards to feed on snakes. It commonly eats millipedes and carabid beetles, which use distasteful and noxious substances in defense. Some other distasteful species such as woodlice, staphylinid beetles, and some ladybird beetles seem to be avoided. It enjoys bread and milk, pet food, and kitchen scraps offered by humans. When foraging, it generally progresses slowly and pauses briefly to sniff the air or ground. It also will forage in a more rapid andlinear style whereit covers much more ground but still searches for suitable prey. The West European Hedgehog always is alert and constantly sniffing and listening. Prey likely are detected mainly by their odor, but sound also is important in finding prey. Prey items usually are snapped up rapidly, anditems buried in litter, grass, or soil are quickly rooted out. Males tend to travel faster than females while foraging, and both sexes travel fastest in short grass and progressively slower as density of cover increases. Breeding. The West European Hedgehog is believedto be polygynous, but multiple paternities oflitters indicate that at least some individuals practice polyandry. There usually are two breeding seasons each year, and they vary geographically. In Scandinavia, breeding occurs in May to mid-July, and in England, pregnancies occur in May-October, with peaks in May-July and September. Later mating generally occurs ifthe first litter is lost or aborted. Rate ofgrowth ofyoung borntolate litters is faster than for young bornto early litters. There is no difference in rate of growth between sexes. Sexual cycle of males varies regionally. During breeding season, reproductive organs of males increase to 10%ofbody weight. Gestation lasts 31-35 days, but some estimates are up to 42 days. Litters have 4-7 young (range 1-11). Reports of conjoined twins are rare. Young are born in nests ofleaves. Neonates are nearly hairless, eyes are closed, length is ¢.70 mm, and weights are 8-25 g depending onsize oflitter. Young are born with soft whitish spines that harden and begin to be replaced by adult-type spines at 2-7 days old. By about three weeks old, spines look like those of adults. At four weeks, young begin following their mother and are weaned at 4-6 weeks old. Males do not participate in rearing young. Sexual maturity is achieved at 9-12 months. In Sweden, females do not breed until their second year. Maximum body size is attained at 2-3 years old. Maximumlife expectancy is ¢.6-8 years in the wild and ten years in captivity. Activity patterns. The West European Hedgehog is predominantly nocturnal, resting by day under piles of brush or in leaf nests in rocky crevices and burrows. Activity during midday often indicates that an individual is sick, but healthy individuals can be active during the day, particularly in autumn. Some lactating females can be almost as active during day as at night during summer, some individuals tend to emerge before sunset, and some retire to their nest well after dawn. Usual gait is slow, meandering walk that frequently is punctuated by short bursts of speed up to 50-120 m/minute. Males usually move at an average speed of 3-7 m/minute, females at 2:2 m/minute, and subadults at 2:2 m/minute. The West European Hedgehog lives on the surface ofthe ground without extensively burrowing or climbing. It is a good climber, and spines, besides serving as protection from predators, act as a cushion when it falls or drops from a height. It is a good swimmer but might not be able to escape from steepsided garden pools. It hibernates from about October to early April in colder parts ofits distribution, but individuals become active, emerge, and move to another nest 2-3 times in winter. Hibernation bouts last 5-27 days (average nine days). Some nests are repaired and re-used, and others are abandoned. Nests may be occupied, then abandoned for up to four months, and later re-occupied. Others are empty at first and occupied later, suggesting there are spare nests. Movements, Home range and Social organization. Although primary foraging area of a West European Hedgehog can be ¢.200-300 m from its day-shelter, average minimal nightly distances traveled are 868-1785 m for adult males, 570-1006 m for adult females, and 1188 m for subadults. Some males travel up to 3 km/night. Most individuals are active all night. The West European Hedgehog is solitary except during the breeding season or for a mother with her young. It usually is non-territorial, but it is intolerant of conspecifics. Rather than maintaining a defended area, it uses mutual avoidance to allow non-simultaneous use ofthe same area and thus avoid direct competition. Individuals rarely meet, and adults ofthe same sex usually do not approach to within 20 m ofeach other. Adults usually are together only for mating or when attracted to a localized food source where fights sometimes occur; otherwise fights are rare. Presumably as time to hibernate approaches, they vigorously defend territories of1:8-2.5 ha, but in spring after emergence from hibernation, individuals do not stay in these defended areas. At that time, they wander over large areas looking for food and mates, and there may be intensive fighting between males. Home ranges overlap, often completely, irrespective ofsex. In England, average home range of adult males is larger (32 ha; range 16-42 ha) than that of adult females (10 ha; range 6-12 ha) and subadults (12 ha; range 10-15 ha). In Sweden, adult males have larger home ranges (47 ha; range 25-68 ha) than females (20 ha; range 8-30 ha), and in Russia, home ranges are 0-5—11-2 ha for males and 0-8-3-5 ha for females. There is a tendency for both sexes to occupy the same home range for years. Young have a dispersal phase in the first few months oflife; they establish their future home range before, or shortly after, their first winter. Nesting sites usually are shaded by vegetation or other objects. In summer and winter, nests are mostly aboveground, under brambles or other vegetation, and often are constructed ofdry, broad leaves. Burrows are simple, blind, and c.1 m long, with a small chamber at the end. Such burrows are excavated by hedgehogs or by rabbits. The West European Hedgehog engages in self-anointing with frothy saliva. Reasons for self-anointing are unknown, but it might be used to attract mates, as a defense from predators, or by juveniles seeking attention from their mothers. Self-anointing is dependent on gender, age, and season. First-year independent young self-anoint more than adults, and males self-anoint more than females. In adults, selfanointing peaks in summer. Typical defensive posture is to roll into a ball, covering vulnerable underside, face, and limbs with spines. Status and Conservation. Classified as Least Concern on The IUCN Red List. The West European Hedgehog is common, widespread, and generally well-known. It consumes pests in gardens, is popular in folklore and books, and is eaten by humans, although that is becoming uncommon. Starvation and collisions with motor vehicles are major causes ofdeath. In Belgium, vehicles kill at least 230,000-350,000 West European Hedgehogs each year, and throughout Europe, c¢.1-6 ind/kmofroads arekilled each year. Populations have decreasedsignificantly: 15-22%in Sweden, 26%in Switzerland, and 30% in the Netherlands. Bibliography. Allanson (1934), Beer (2003), Berthoud (1980), Boitani & Reggiani (1984), Boukheroufa et al. (2015), Brodie (1977), Bunnell (2009), Corbet & Southern (1977), D'Havé, Scheirs, Verhagen & De Coen (2005), Dickman (1988), Dmi'el & Schwarz (1984), Gropp et al. (1969), Haigh et al. (2012), Harrison Matthews (1952), Hayssen et al. (1993), Herter (1965), Holsbeek et al. (1999), Huijser & Bergers (2000), Hutterer (2005a), Jackson, D.B. (2007), Karaseva et al. (1979), Kompanje (2005), Kral (1967), Kristiansson (1981, 1990), Lay (1967), Mathias et al. (1998), Moran et al. (2009), Morris (1977, 1985, 1986, 1988, 2006), Morris & Tutt (1996), Mouhoub Sayah et al. (2009), Nowak (1999), Ortowski & Nowak (2004, 2006), Parkes (1975), Pavlinov & Lissovsky (2012), Philchagov (1988), Reeve (1982, 1994), Reeve & Morris (1985), Skoudlin (1981), Smith (1992), Stone (1995b), Sykes & Durrant (1995), Thomas (1911b), Warwick (2014)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 319, DOI: 10.5281/zenodo.6639332

10. Brandt's Hedgehog Paraechinus hypomelas French: Hérisson de Brandt / German: Brandt-Igel / Spanish: Erizo de Brandt Other common names: Brandt's Steppic Hedgehog, Long-spined Hedgehog Taxonomy. FErinaceus hypomelas Brandt, 1836, “pays des Turcomans [= country of the Turkmens].” Interpreted by D. R. Frost and colleagues in 1991 as “roughly equivalent to the Turkmen, Uzbek, Kazak, and Karakalpak regions of the [former] U.S.S.R.”” and by R. Hutterer in 2005 as “somewhere in S Kazakhstan.” Five subspecies are recognized. Subspecies and Distribution. P.h.hypomelasBrandt,1836—mostof Turkmenistan,SEUzbekistan,SWTajikistan,andIran,includingKhargI. P.h.blanfordiJ.Anderson,1878—EIran,Afghanistan,andPakistan. P.h.eversmanniOgnev,1927—EofCaspianSeainSWKazakhstan,WUzbekistan,andNWTurkmenistan. P.h.sabaeusThomas,1922—ArabianPeninsula. P. h. seniculus Thomas, 1922 — Tunb I, Persian Gulf, Iran. Descriptive notes. Head—body 136-290 mm, tail 18-38 mm, ear 36-55 mm, hindfoot 30-46 mm. No specific data are available for body weight. Typically, Brandt's Hedgehog has striking dark brown or black pelage, with no white band across forehead, but white (not albino) individuals have been documented. Dorsal pelage has spines, with rough surfaces and strong grooves along their length; they are up to 36-37 mm long. Median parting of spines occurs on crown of head. Spines have black tips, often with entire distal one-third being black. Underside, legs, and face are covered with mix of black and gray hairs. Generally, middle part of forehead has more black hairs, with admixture of gray hairs on sides of face. Throat has mix of creamy-white hairs. Legs are long, large ears protrude beyond adjacent spines, and hallux is somewhat reduced but consistently well developed. Habitat. Dry deserts, desert steppes, and other arid places. In Pakistan, Brandt's Hedgehog appears to prefer dry gravelly slopes or rocky areas, and it survives and thrives in barren arid areas. It also can occur in lowland areas and will venture into cultivated fields. Food and Feeding. Brandt's Hedgehog is a voracious feeder and will tackle almost any prey from venomous snakes to beetles. Although it primarily is insectivorous, it also eats scorpions, other invertebrates, small vertebrates, eggs of ground-nesting birds, and fruits. In years when the desert locust (Schistocerca gregaria) is plentiful, it forms the bulk of the diet. It has a fondness for ripe melons that are an important summer crop. It also feeds on fallen mulberries and silverberries (Elaegnus, Elacagnaceae). It may be cannibalistic, more so in captivity than in the wild. Breeding. In non-mountainous areas where there is some monsoon influence, Brandt's Hedgehog possibly breeds throughout the year. Litters usually are born in late spring and early summer (April-May). Average litters have 3—4 young (range 1-6). Eyes open in 21-23 days, and teeth appear in 26-28 days. There is no evidence that more than onelitter is produced each year. A captive Brandt's Hedgehog lived seven years and two months. Activity patterns. Brandt's Hedgehog is terrestrial and nocturnal; it forages most often at night. It has been observed day-resting in burrows 1m deep in sandy embankments of railway tracks. Movements, Home range and Social organization. Brandt's Hedgehog is solitary and rather nomadic, and it does not use the same shelter continuously throughout the year. Day-resting sites can be under overhanging ledges, in crevices between rocks, and occasionally burrows. Its legs are relatively longer and claws shorter and sharper compared with other species of hedgehogs. These physical traits might be a disadvantage in burrowing and might explain why it is not a very active burrower. It exhibits self-anointing behavior. Status and Conservation. Classified as Least Concern on The IUCN Red List. Increasing desertification in its distribution is resulting in fragmentation of populations. Bibliography. Anderson (1878), Beer (2003), Brodie (1977), Corbet (1988), Frost et al. (1991), Habibi (2004), Hutterer (2005a), Nader (1991), Nowak (1999), Ognev (1928), Reeve (1994), Roberts (1997), Stone (1995b), Thomas (1922a), Walton & Walton (1973), Zherebtsova (1992)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 324, DOI: 10.5281/zenodo.6639332

28. Least Shrew Tenrec Muicrogale pusilla French: Petit Microgale / German: Gnomkleintenrek / Spanish: Tenrec musarana minima Other common names: Lesser Shrew Tenrec Taxonomy. Microgale pusilla Major, 1896, “Forest of the Independent Tanala of Ikongo, in the neighbourhood of Vi- nanitelo,” Madagascar. Restricted by R. D. E. MacPhee in 1987 to “50 km SE of Fianarantsoa town and 10 km SSE of Vo- hitrafeno town, western margin of eastern forest; Fianarantsoa, Fianarantsoa [Province, Madagascar]; 21°45’S, 47°17L. Further clarified by M. D. Carleton and D. F. Schmidt in 1990 and P. D. Jenkins and Carleton in 2005 to “21°43’S, 47°16’E.” Recent comprehensive molecular phylogenetic analysis showed that M. pusilla is sister to a clade of M. jenkinsae, M. longicaudata, M. majori, and M. principula. Monotypic. Distribution. Central Highlands, E & SE Madagascar. Descriptive notes. Head—body 51-63 mm,tail 62-85 mm, ear 8-13 mm, hindfoot 11-13 mm; weight 3-5 g. The Least Shrew Tenrec is very small, and tail is long, 130-160% of head-body length. Dorsal pelage is soft and grizzled reddish brown, and venteris gray, with a buffy wash. Habitat. Humid forests (tolerant of some disturbance) and wet grasslands to marshy habitats at elevations of 530-1670 m (but recently documented at higher elevations). Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Least Shrew Tenrec is widely distributed and occurs in several protected areas. It uses some disturbed habitats, including urban rice fields. Its overall population appears to be stable, but major threats could be habitat degradation and use offire for forest clearing. Bibliography. Carleton & Schmidt (1990), Everson et al. (2016), Goodman etal. (2013), Jenkins (2003), Jenkins & Carleton (2005), MacPhee (1987), Olson & Goodman (2003), Stephenson et al. (2016m)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 171-172, DOI: 10.5281/zenodo.6808230

18. Hugh's Hedgehog Mesechinus hughi French: Hérisson de Hugh / German: Hugh-Igel / Spanish: Erizo de Hugh Other common names: Central Chinese Hedgehog, Shanxi Hedgehog Taxonomy. Erinaceus hughi Thomas, 1908, “Paochi, Shen-si.” Interpreted by D. R. Frost and colleagues in 1991 as “Baoji, Shaanxi Sheng, China, 34°23’N, 107°09°E.” This species is monotypic. Distribution. S Gansu, S Shaanxi, Shanxi, NE & SE Sichuan, and S Henan in C China. Descriptive notes. Head-body 148-232 mm, tail 12-24 mm, ear 16-33 mm, hindfoot 30-47 mm; weight 112-750 g. Hugh’s Hedgehog is very dark. Dorsal pelage has spines that are smooth and have minute, subterminal, and pale band just before dark tips. Ears are no longer than adjacent spines. Ventral pelage is pale. Habitat. Semiarid and dry steppe habitats and subalpine and low-elevation coniferous forests. Food and Feeding. No information. Breeding. No information. Activity patterns. There is no specific information for this species, but Hugh’s Hedgehog is terrestrial. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Hugh's Hedgehog occupies relatively small area of suitable habitat thatis shrinking because of human encroachment. It has been regionally Red Listed in China as vulnerable. Research is needed to better understand population status, biology and ecology, habitat status, and conservation threats. Bibliography. Ai Huaisen et al. (2018), Beer (2003), Corbet (1988), Frost et al. (1991), Hutterer (2005a), Kong Fei, Wu Jiayan & Guo Jianmin (2016a), Nowak (1999), Reeve (1994), Smith & Yan Xie (2013), Stone (1995b), Thomas (1908b)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 327, DOI: 10.5281/zenodo.6639332

15. Lesser Long-tailed Shrew Tenrec Mucrogale longicaudata French: Microgale a longue queue / German: Kleiner Langschwanz-Kleintenrek / Spanish: Tenrec musarana de cola larga menor Taxonomy. Microgale longicaudata Thom- as, 1882, “Ankafana forest, Eastern Bet- sileo,” Madagascar. Restricted by R. D. E. MacPhee in 1987 to “Ankafina... 10 km S of Ambohimahasoa and 3 km W of Tsarafidy town, on extreme western margin of eastern rain forest; Fianarantsoa, Fianarantsoa [Province, Madagascar]; 21°12’S,47°13’E.” Analysis of mtDNA revealed several cryptic taxa within M. longicaudata including M. major. Subsequent comprehensive molecular phylogenetic analysis showed that M. longicaudata issister to M. jenkinsae and forms a clade with M. jenkinsae, M. majori, and M. principula. Microgale prolixacaudata was potentially one of the cryptic taxa revealed by the mtDNA analysis but could not be defined at that time, and despite its subsequent recording as a distinct species, it still awaits accurate definition. Microgale longicaudata might contain cryptic species, but taxonomy requires reassessment. Here treated as monotypic. Distribution. N, Central Highlands, E & SE Madagascar. Descriptive notes. Head—body 59-80 mm, tail 136-158 mm, ear 10-16 mm, hindfoot 12-18 mm; weight 6-11 g. The Lesser Long-tailed Shrew Tenrec is small to mediumsized, with very long, partially prehensile tail; tail is greater than twice head-body length. Dorsal pelage is reddish brown; venteris gray, with reddish buff or buff wash; and tail is gray-brown above, clearly distinguished from reddish buff below. Digits of forefeet and hindfeet are elongated, and fifth hind digit is only slightly shorter than second. Distal portion of tail is naked, with broad transverse scales on dorsal surface. Habitat. Humid forests (tolerant of some disturbance) at elevations of 530-2500 m. Food and Feeding. Diet of the Lesser Long-tailed Shrew Tenrec contains species of Orthoptera, Coleoptera, Hymenoptera, Arachnida, and Amphipoda. In one captive study, insects up to 6 cm long were eaten, and some species of insects were preferred when an individual was offered a choice. Hindfeet and distal portion of long tail were used for grasping branches and stabilization when chasing insects and feeding. Breeding. One wild-caught lactating Lesser Long-tailed Shrew Tenrec had two placental scars in November. Activity patterns. Lesser Long-tailed Shrew Tenrecs are presumably terrestrial, scansorial, and semi-arboreal based on morphology and do not appear to hibernate. In a short study, one captive individual was nocturnal, spending the day in a nest in the leaf litter c.5 cm in diameter, becoming active at 18:30-19:00 h on the ground and on small branches, and spending rest periods on branches. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Lesser Long-tailed Shrew Tenrec is widely distributed and occurs in several protected areas. Its overall population appears to be decreasing, and major threats are habitat loss and fragmentation due to logging activities, use of fire for land clearing, and conversion to agricultural land. Bibliography. Everson et al. (2016), Goodman & Jenkins (1998, 2000), Goodman, Jenkins & Pidgeon (1999), Goodman, Soarimalala et al. (2013), Hilgartner (2005), Jenkins (2003), Jenkins et al. (1996), MacPhee (1987), Olson & Goodman (2003), Olson, Goodman & Yoder (2004), Soarimalala & Goodman (2011), Stephenson et al. (2016p)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 168, DOI: 10.5281/zenodo.6808230

21. Short-tailed Gymnure Hylomys suillus French: Petit Gymnure / German: Kurzschwanz-Rattenigel / Spanish: Gimnuro de cola corta Other common names: Lesser Moonrat, Short-tailed Moonrat Taxonomy. Hylomys suillus S. Miller, 1839, “Java en het andere van Sumatra.” Re- stricted by H. J. V. Sody in 1933 to “Java,” Indonesia. Seven subspecies are recognized. Subspecies and Distribution. H.s.swillusS.Muller,1839—Java. H.s.dorsalisThomas,1888—NBorneo(fromMtKinabaluStoKelabitHighlands). H.s.maxiSody,1933—SThailand,MalayPeninsula,andWSumatra. H.s.microtinusThomas,1925—SWChina(Yunnan),Vietnam,andCambodia. H.s.peguensisBlyth,1859—EMyanmarandadjacentSWChina(Yunnan). H.s.siamensisKloss,1916—SEMyanmar,N&WThailand,andLaos. H. s. tionis Chasen, 1940 — Tioman I, off SE Malay Peninsula. Descriptive notes. Head—body 90-156 mm, tail 10-30 mm, ear 12-21 mm, hindfoot 20-28 mm; weight 45-80 g. The Short-tailed Gymnure is small, and its short tail distinguishes it from all other gymnures. Snoutis pointed, ears and tail are short, and overall appearance is ratlike. Pelage is soft and dense. Upper parts are rusty brown, sometimes with faint black nape stripe or dorsalstripe, and underparts are grayish, brownish, or yellowish. Microstructures of hairs and vibrissae of the Short-tailed Gymnure are similar to those in the Long-eared Gymnure (H. megalotis) and the Shrew Gymnure (Neotetracus sinensis); there are spearshaped lead hairs and zig-zag guard hairs, but there are no downy hairs. Dental formulais13/3,C1/1,P4/4,M 3/3 (x2) = 44. Habitat. Usually confined to dense, damp, forested areas, especially hilly, mountainous, and lowland forests with thick undergrowth. In China, the Short-tailed Gymnure occurred in secondary forests that had been clear-cut 13 years previously but was not documented in nearby tropical monsoon forests. In Thailand, it occurred under logs in a grove of wild bananas. It has been recorded at elevations of 90 m on mainland South-east Asia to 3000 m on Sumatra and 1000-3400 m on Borneo. Its presence has been verified along edges of a disturbed montane rainforest at 1200-1500 m in Peninsular Malaysia, in lower montane rainforest at 1700 m on Java, and in lower and upper montane rainforests at 1500-3000 m on Borneo. Food and Feeding. Diet of the Short-tailed Gymnure is mainly invertebrates such as insects and earthworms, but it also eats fruits and vegetation. It forages by searching underleaf litter and in upperlayers of humus with its long, mobile snout, which tosses aside leaves and twigs with jerks of head and shoulders. Breeding. The Short-tailed Gymnureis polygynous, and it probably breeds throughout the year. A pregnant female with two embryos was observed in February and another with two embryos in March. Litters have 3-6 young. Gestation lasts 30-35 days. Lifespan probably does not exceed two years. Activity patterns. The Short-tailed Gymnure is mainly diurnal but can be active at any time of day or night, singly or in small groups. It is mainly active aboveground, beneath litter, and under logs, but sometimes it climbs on low bushes. It builds nests of leaves in rock crevices, burrows, and hollows among roots of trees. It does not hibernate. Movements, Home range and Social organization. Short-tailed Gymnures are primarily solitary; however, there have been observations of groups of 2-3 individuals that might have been females with young. Although capable of climbing, Short-tailed Gymnures generally stay on the ground, moving quickly and nimbly with short leaps. It appears to use definite paths or runways on the forest floor, often passing under logs that are slightly off the ground. When threatened, the Short-tailed Gymnure bounds away with considerable speed. It emits shrill squeaks that are not loud. In Malaya, average densities in favorable habitats are 3—4 ind/ha. Home ranges are ¢.30-40 m in diameter. The Short-tailed Gymnure has a strong odor. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Shorttailed Gymnure has a wide distribution and relatively stable overall population. Bibliography. Beer (2003), Chernova et al. (2015), Hutterer (2005a), Jenkins & Robinson (2002), Lekagul & McNeely (1988), Medway (1983), Nowak (1999), Phillipps & Phillipps (2016), Rudd (1980), Ruedi et al. (1994), Smith & Yan Xie (2013), Sody (1933), Stone (1995b), Wu Delin et al. (1996)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 328, DOI: 10.5281/zenodo.6639332

26. Dinagat Gymnure Podogymnura aureospinula French: Gymnure de Dinagat / German: Dinagat-Rattenigel / Spanish: Gimnuro de Dinagat Other common names: Dinagat Moonrat, Dinagat Wood Shrew, Golden-spined Moonrat, Spiny Moonrat Taxonomy. Podogymnura aureospinula Heaney & Morgan, 1982, “Plaridel, Albor Municipality, Dinagat Island, Surigao del Norte Province, Republic of the Philip- pines.” Interpreted by D. R. Frost and colleagues in 1991 as “10°12°N, 125°35’E.” This species is monotypic. Distribution. Dinagat and Bucas Grande Is, S Philippines. Descriptive notes. Head-body 190-211 mm, tail 59-73 mm, ear 22-26 mm, hindfoot 39-42 mm. No specific data are available for body weight. The Dinagat Gymnure has spiny pelage and is second in size only to the Moonrat in the subfamily Galericinae. Dorsal pelage is short, spinous, golden-brown overall, and composed of three types of hairs: slate-gray underfur ¢.5 mm long; stiff, bristly, or spiny hairs c.15 mm long with black bases, others are golden-yellow, and many have black tips; and black, spiny hairs, c¢.12 mm long. Golden spines are twice as abundant as black hairs in middle of back. Black spines are densest at mid-dorsum, decreasing in abundance laterally, and disappearing on sides. Black-tipped golden spines are especially common on mid-dorsum and disappear on sides. Only golden spines are present on sides. Overall color of dorsum is golden-brown, with black spines and black-tipped golden spines, adding black speckling. Ventral pelage lacks spines and grades evenly from dorsal color to brownish gray over most of venter. Ventral hairs are of two types: soft, gray underfur c.5 mm long and slightly coarser guard hairs ¢.9 mm long, gray at bases, and tipped with pale brown. Pelage of rostrum and around eyes is short, dense, and spiny. Upper and lowerlips are covered in short, moderately dense, white, or pale brown fur. Vibrissae are dark at bases, very pale for most of their lengths, and up to 55 mm long. Rhinarium is long, naked, and distinctly bilobed, with nostrils opening laterally. Ears are relatively large and appear naked, but they have sparse covering of extremely short, white hairs. Forefeet and hindfeet have moderate coverings of short, white, or pale brown hairs dorsally and are nearly naked ventrally. Hindlegs appear almost naked distal to knee. Dorsal base oftail is sparsely furred, nearly naked, and 15 mm in diameter. Tail has sparse covering of short hairs. Dental formula is I 3/3, Cl1l/1,P3/3,M 3/3 (x2) =40. Habitat. Primary and secondary forests. The Dinagat Gymnure is known from a logged area in rolling low hills where there were many patches of remnant dipterocarp forests. These forests contained at least twelve families of plants including Dipterocarpaceae, which dominated the habitat with Anisoptera, Dipterocarpus, Hopea, and Shorea. Undergrowth was mainly rattans (Calamoideae) and ferns. Food and Feeding. Diet of the Dinagat Gymnure contains invertebrates. Breeding. No information. Activity patterns. The Dinagat Gymnure is nocturnal. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Endangered on The IUCN Red List. The Dinagat Gymnure is restricted to patches of forested habitats that are being destroyed by logging and slash-and-burn agriculture. Bibliography. Beer (2003), Frost et al. (1991), Heaney & Morgan (1982), Heaney & Rabor (1982), Heaney, Balete et al. (1998), Heaney, Dolar et al. (2010), Hutterer (2005a), Jenkins & Robinson (2002), Nowak (1999), duPont & Rabor (1973), Reeve (1994), Stone (1995b).

13. Common Long-eared Hedgehog Hemiechinus auritus French: Hérisson a oreilles longues / German: Eigentlicher Langohrigel / Spanish: Erizo de orejas largas dorado Other common names: Afghan Hedgehog, Long-eared Steppe Hedgehog Taxonomy. Erinaceus auritus S. G. Gmelin, 1770, “in regione Astrachanensi.” Inter- preted by D. R. Frost and colleagues in 1991 as “Astrackhan’skaya Oblast’, [former] U.S.S.R., 46°21’N, 48°03’E,” now southern Russia. Three subspecies are recognized. Subspecies and Distribution. H.a.auritusS.G.Gmelin,1770—NLibya,NEgypt,MiddleEast,EUkraine,SWRussia,ECaucasus,extremeE&SAnatolia(Turkey),andN&WIran.IntroducedintoCyprus. H.a.albulusStoliczka,1872—Kazakhstan,SSiberia(Tuva),W&SMongolia,andNW&NCChina. H. a. megalotis Blyth, 1845 — Turkestan, SW & E Iran, Afghanistan, and W Pakistan. Descriptive notes. Head-body 160-278 mm, tail 10-55 mm, ear 24-60 mm, hindfoot 30-55 mm; weight 230-400 g (but some weigh 1-1 kg). Unlike species of Erinaceus and Paraechinus, the Common Long-eared Hedgehog and the Indian Long-eared Hedgehog (H. collaris) have no central parting of spines on crown of head. The Common Long-eared Hedgehog has long limbs, short barely visible tail, well-developed hallux on hindfeet, and five toes on forefeet. Ears are long and conspicuous,slightly pointed, flexible and mobile, considerably longer than adjacent spines, and apparently adapted for radiation of desert heat. If ears are pressed forward, they cover eyes. Dorsal pelage has dark rough spines, with numerous papillae, many shallow longitudinal grooves, whitish bases, two wide black bands separated by pale band, pale brownish tip, and average length of ¢.25 mm. Spines on lower back and flanks are c.34 mm long. Spines are sharp and flexible, and they act as shock absorbersif a fall occurs from a high place. Head,tail, and limbs generally are white to pale brown. There is no dark face mask. Ventral pelage generally has soft white hairs, but populations in Afghanistan and Pakistan (megalotis) are uniformly brown on their undersides. In Pakistan, the Common Long-eared Hedgehog tends to be rather reddish. Hair on limbs, underside, tail, and lower part of head is reddish brown or auburn. Upper part offace, inside of ears, and top of crown has some white hairs that produce rusty color. Entire body tends to look dark reddish brown, and spines give an impression of lying in a rather smooth and regular pattern because of brown tips and their lengths. Dental formulais 13/2, C1/1, P3/2,M 3/3 (x2) = 36. Chromosomal complement has 2n = 48-50 and FN = 94-98. Habitat. Dry riverbeds, dunes, oases, valleys with shrubs, semiarid country, and steppes over most of its distribution, including coastal semi-desert scrublands with dense vegetation, semiarid grasslands, and high-elevation steppes with subzero temperatures in winter. The Common Long-eared Hedgehog is not desert adapted, but it is moderately tolerant of heat and drought, and its large ears might be important in thermoregulation. It is rare in sparsely vegetated areas. It favors mesic sites such as suburban parks and gardens, olive groves, and agricultural lands and is commonly associated with humans. In Pakistan,it is plentiful above elevations of 900 m and can occur in valleys up to 2500 m. In Iran, it spends days in shallow burrows, beneath mud walls, or in piles of hay. In Egypt, it often is in buildings, crevices in walls, piles of bricks and stones, fissures in rocks, small caves, graveyards, and burrows of the Fat Sand Rat (Psammomys obesus) in salt-marsh habitats. It selects soft ground such as sandy dunes where it can dig relatively deep burrows that slope at a slight angle. Food and Feeding. The Common Long-eared Hedgehog is omnivorous,it is a voracious feeder, and it can survive for up to ten weeks without eating or drinking. Diet primarily is insects (bugs, beetles, grasshoppers, termites, lepidopterans, ants), but it also eats other invertebrates (worms, millipedes, and snails), small vertebrates (mammals, birds, snakes, and lizards), eggs, carrion, fruits, and seeds. It seems immune to poisons and stings of insects and other invertebrates and will hunt and eat scorpions, venomous spiders, and stinging hymenopterans. It will kill and eat small snakes, some venomous, by rolling into a ball after grabbing the lower body of the snake so that the snake strikes in vain against the spines, often damaging itself. The Common Longeared Hedgehog feeds on grapes, mulberries, and watermelons, which it hollows out. It does not eat food discarded by humans, and captive individuals show a tendency toward cannibalism. Even with its protective spines,it is an important source of food for carnivores. Breeding. The Common Long-eared Hedgehog is polygynous. Breeding probably occurs immediately after emerging from hibernation in early spring (mid-April) and continues into late summer (September). Courtship behavior is well developed and includes sniffing, licking genitalia, and flank rubbing. Male copulates by standing almost vertically on his hindlegs, while the female lies on her abdomen with hindlegs extended. Gestation lasts 28-42 days. Births peak in July-September. Before giving birth, females excavate nest chamber in their burrow. Apparently, they clear all debris from the floor of the nest chamber, they do not line the chamber, they give birth on the smooth floor, and they lick neonates until they are clean. Litters vary geographically but usually have 3-5 young (range 1-7). Neonates are born naked, except for some scattered spines. They weigh 7-13 g, body length is 47-61 mm, hindfoot length is 7 mm, and ear length is 3-4 mm. Their backs are thickly covered with spines by 14 days old, eyes open at 15-20 days, solid food is eaten at 21-28 days, teeth appear at 23-24 days, weaning is at ¢.31-40 days, and young leave nest at ¢.30-40 days. There is no evidence that more than one litter is produced each year. Young can reach sexual maturity as soon as six weeks of age. Longevity is 6-7 years. Activity patterns. The Common Long-eared Hedgehog is nocturnal and begins foraging early in the evening. Day-resting sites usually are in burrows that are under small shrubs or bushes. Burrows are 20-150 cm long and have a single opening. Rest sites occur in caves and crevices and under piles of rocks and bricks. Long limbs enable it to walk and run quickly. It is an active digger and will excavate its own burrow when abandoned dens made by rodents or other mammals are not available. Generally, it excavates burrows under large rocks and can dig in sandy, clay, or rocky soils. Digging employs broad-clawed forefeet to loosen and throw soil backward; snout is not used in digging. The Common Long-eared Hedgehog periodically moves backward up its burrow and kicks out accumulated soil using hindfeet. In firm soil, 30-38cm burrows are completed in 3-4 hours. In soft soil,it can dig a 10cm hole in five minutes and can dig several burrows each night. It enters and exits burrows headfirst. Females widen ends of burrows to accommodate young. During early spring when insects and other invertebrates are plentiful, it builds up a layer offat. It can enter torpor in any season during droughts, shortages of food, and cold or hot weather, and it can remain partially awake, or sufficiently conscious, to react quickly to any disturbance. Even during short periods of torpor and estivation, respiration is slow and shallow, and heart rate decreases. Body temperature usually remains within 1-2°C of ambient temperatures. In regions where temperatures in winter are as cold as —6-5°C, body temperature during hibernation can approach 0°C. In North Africa, it can enter torpor during the cold season (December—February) and remain in its burrow 5-40 days. In northern India, it might hibernate up to 3-5 months in winter, and in the mountains of Pakistan, hibernation occurs in October-March and can be 5-5-6 months. In China, it enters torpor in October and emerges in spring. During hibernation, the Common Longeared Hedgehog wakes up periodically and might emerge briefly to forage or move only within its nesting burrow. In warmer areas, there is no prolonged hibernation in winter. Movements, Home range and Social organization. Common Long-eared Hedgehogs can walk more than 17-20 m/minute. Estimates indicate that the Common Longeared Hedgehog can travel 7-9 km in a 7-9hour activity period. Average home ranges are 4-9-5 ha (range 1-6-10 ha) for adult males, 2-9 ha (range 0-9—4-2 ha) for nonlactating females, and 1:3-3-5 ha (range 0-2-4-3 ha) for non-lactating and lactating females. The Common Long-eared Hedgehog is solitary. It usually is non-aggressive, but when fighting, it is very noisy; it growls when alarmed. Adults only nest singly, but they use burrows of other hedgehogs in their absence. It performs self-anointing; practically any substance will induce this behavior. This activity can go on nearly continuously for up to 20 minutes. Reasons for this behavior are unknown; it might be a sexual signal because it occurs only in adults during the breeding season or, because substances that initiate this behavior are either novel or irritating, it might serve as protection against predators. The Common Long-eared Hedgehog also rubs toads against its spines suggesting that it might be making defensive use of toxins produced by the toads. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Common Long-eared Hedgehog is common in much ofits broad distribution. It can damage agricultural crops, especially ripe melons, but it is mostly beneficial because of the insects, reptiles, and other pests that it eats. Humans sometimes eat Common Longeared Hedgehogs. Bibliography. Al-Behbehani & Ibrahim (2007), Al-Khalili (1990), Arslan et al. (2009), Atallah (1977), Batsaikhan et al. (2010), Beer (2003), Bhatnagar & El-Azawi (1978), Blanford (1888), Blyth (1845), Brodie (1977), Colak et al. (1997 1998), Corbet (1988), Dmi‘el & Schwarz (1984), Flower (1932), Frost et al. (1991), Happold (2013d), Harrison (1964), Harrison & Bates (1991), Hassinger (1973), Hayssen et al. (1993), Herter (1965), Hoogstraal (1962), Hufnagl (1972), Hutterer (2005a), Ivanitskaya & Malygin (1985), Jones (1982), Karatas et al. (2007), Kefelioglu (1997), Krishna & Prakash (1955), Kumar & Pandey (1995), Lay (1967), Nader (1968), Niethammer (1973), Nowak (1999), Ognev (1928), Osborn & Helmy (1980), Poduschka & Poduschka (1986), Prakash (1953, 1954, 1960), Qumsiyeh (1996), Reading et al. (2010), Reeve (1994), Roberts (1997), Schoenfeld & Yom-Tov (1985), Smith & Yan Xie (2013), Stoliczka (1872), Stone (1995b), Zherebtsova (1982, 1992)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, pp. 288-330 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 325-326, DOI: 10.5281/zenodo.6639332

Moniliformis cryptosaudi n. sp. (Moniliformidae) is an acanthocephalan described from the long-eared hedgehog Hemiechinus auritus (Gmelin) (Erinaceidae) in Iraq as an incipient cryptic species of Moniliformis saudi Amin, Heckmann, Mohammed, Evans, 2016 described from the desert hedgehog Paraechinus aethiopicus (Ehrenberg) (Erinaceidae) in Saudi Arabia. Microscopical studies demonstrate that the two species are morphologically indistinguishable with practically identical measurements and counts but differed significantly in their energy dispersive X-ray analysis (EDXA) of metal composition of hooks. Hooks of specimens of the new species appeared to be of collagen material with very low levels of phosphorus and calcium unlike those of M. saudi and Moniliformis kalahariensis Meyer, 1931 that had high levels of calcium and phosphorus. Using 18S rDNA and cox1 genes, M. Saudi and M. kalahariensis were shown to be molecularly distinct but the molecular profiles of M. saudi and M. cryptosaudi were more similar. The molecular profile of M. kalahariensis collected from the South African hedgehog Atelerix frontalis Smith (Erinaceidae) in South Africa is reported for the first time and is studied only for comparative purposes. Moniliformis saudi and M. kalahariensis had comparable EDXA metal analysis that was distinct from that of M. cryptosaudi.

Medulloblastoma, the most common pediatric malignant brain tumor, is divided into at least four molecular subtypes. The Sonic Hedgehog (SHH) subtype, accounts for approximately 30% of all cases and has an intermediate prognosis (5-year overall survival of 60%). Current treatments involve surgery, adjuvant radiotherapy for older children, and chemotherapy, however, lead to devastating morbidity, including decline in cognition and intellect, endocrine deficiencies, and secondary malignancies. More recently, targeted treatment options that inhibit the Hedgehog pathway have demonstrated clinical efficacy, including the Smoothened inhibitor, Vismodegib, however result studies have also shown secondary toxicities including effects on bone development. Here we investigated the efficacy of Vismodegib packaged in a fucoidan-based nanoparticle (Fi-Vis) that targets P-selectin, a protein overexpressed on vascular endothelial cells and is induced by 2Gy ionizing radiation (XRT) in a time- and dose-dependent manner. This p-selectin targeting nanoparticle drug delivery system shows selectivity toward tumor vasculature and not normal brain vasculature in a genetically relevant SHH-driven medulloblastoma mouse model as assessed by ex vivo infrared imaging. Quantitative RT-PCR analysis on SHH medulloblastoma tumor tissue following 2Gy XRT and Fi-Vis single dose treatment (as low as 10mg/kg) showed a synergistic inhibition of Gli1 expression (up to ~90% target inhibition). Furthermore, we demonstrate that lower single fraction XRT doses as low as 1Gy also induce P-selectin expression within medulloblastoma tumor vasculature. These data suggest applicability of combined XRT and tumor vasculature-targeted nanotherapeutic dose de-escalation strategies for medulloblastoma and potentially other pediatric brain tumors.

The classic locality of Hasznos (Middle Miocene, Mid Badenian, Hungary) has yielded a diverse assemblage of insectivores that is dominated by Erinaceidae and Soricidae, with three species each. The Talpidae are represented by one species, Desmanodon aff. crocheti. Prior to this record of D. crocheti in Hasznos, this species has only been recorded in the Late Badenian of the North Alpine Foreland Basin. The high similarity of the assemblage to that of Samsonhaza 3 confirms that Hasznos and Samsonhaza are very close in age. The new data are discussed in their biostratigraphical and biogeographical contexts.

Hedgehogs (erinaceid insectivores) are a common element in Miocene small mammal faunas of Pakistan, but little material has been formally described. Here, we report on extensive collections from numerous localities across Pakistan, most from the Potwar Plateau, Punjab, and the Sehwan area in Sindh. The dominant erinaceid is Galerix, which is also known from Europe, Turkey, and East Africa. We document a new early species of Galerix, Galerix wesselsae, in sites from Sehwan, the Zinda Pir Dome, the Potwar Plateau, and Banda Daud Shah ranging in age from about 19 to 14 Ma. This species briefly co-occurs with and is then replaced by the smaller Galerix rutlandae, previously based on a few teeth from Daud Khel. G. rutlandae is a common element of Chinji Formation faunas through about 11.5 Ma. Scanty material of an apparently new species of Schizogalerix succeeds Galerix. After 10.2 Ma, no galericines occur in Pakistan, but rare erinaceines are later faunal components.

In the Early Pleistocene locality of Cueva Victoria, the insectivores are represented by several fossil teeth, mandibles and skull fragments. In the present work, we describe and provide measurements of these fossils for the first time. Insectivores show a rather low taxonomic diversity in Cueva Victoria, with only two different species. Considering the environmental requirements for their extant relatives, the presence of only one hedgehog (Erinaceus cf. praeglacialis Brunner, 1933), and one shrew (Crocidura kornfeldi Kormos, 1934) points to the existence of dry and warm conditions in the surrounding area of Cueva Victoria, unfavourable to other contemporary species of insectivores.

International audience; Background: In Europe, hard ticks of the subgenus Pholeoixodes (Ixodidae: Ixodes) are usually associated with burrow-dwelling mammals and terrestrial birds. Reports of Pholeoixodes spp. from carnivores are frequently contradictory, and their identification is not based on key diagnostic characters. Therefore, the aims of the present study were to identify ticks collected from dogs, foxes and badgers in several European countries, and to reassess their systematic status with molecular analyses using two mitochondrial markers. Results: Between 2003 and 2017, 144 Pholeoixodes spp. ticks were collected in nine European countries. From accurate descriptions and comparison with type-materials, a simple illustrated identification key was compiled for adult females, by focusing on the shape of the anterior surface of basis capituli. Based on this key, 71 female ticks were identified as I. canisuga, 21 as I. kaiseri and 21 as I. hexagonus. DNA was extracted from these 113 female ticks, and from further 31 specimens. Fragments of two mitochondrial genes, cox1 (cytochrome c oxidase subunit 1) and 16S rRNA, were amplified and sequenced. Ixodes kaiseri had nine unique cox1 haplotypes, which showed 99.2-100% sequence identity, whereas I. canisuga and I. hexagonus had eleven and five cox1 haplotypes, respectively, with 99.5-100% sequence identity. The distribution of cox1 haplotypes reflected a geographical pattern. Pholeoixodes spp. ticks had fewer 16S rRNA haplotypes, with a lower degree of intraspecific divergence (99.5-100% sequence identity) and no geographical clustering. Phylogenetic analyses were in agreement with morphology: I. kaiseri and I. hexagonus (with the similar shape of the anterior surface of basis capituli) were genetically more closely related to each other than to I. canisuga. Phylogenetic analyses also showed that the subgenus Eschatocephalus (bat ticks) clustered within the subgenus Pholeoixodes. Conclusions: A simple, illustrated identification key is provided for female Pholeoixodes ticks of carnivores (including I. hexagonus and I. rugicollis) to prevent future misidentification of these species. It is also shown that I. kaiseri is more widespread in Europe than previously thought. Phylogenetic analyses suggest that the subgenus Pholeoixodes is not monophyletic: either the subgenus Eschatocephalus should be included in Pholeoixodes, or the latter subgenus should be divided, which is a task for future studies.

The Late Miocene giant erinaceid Deinogalerix from Scontrone and Gargano (Italy) is associated with many other vertebrates in deposits of a past island, the “Abruzzo-Apulia Platform”. At Gargano, Deinogalerix is accompanied by the moderately endemized Galericini Apulogalerix. This first extensive cladistic analysis is aimed at defining the relationships of Deinogalerix with characteristic members of the tribe Galericini. The analysis was performed on a matrix of 30 characters and 19 taxa and identified some smaller clades, nested within three major ones. The latter include: (i) a pentatomy of Galerix species, (ii) a polytomy of “transitional” Galerix–Parasorex species and (iii) a large clade with Parasorex, Schizogalerix and Gargano representatives. Galerix and Parasorex proved to be paraphyletic and Schizogalerix monophyletic. Based on the results of the analysis, Deinogalerix and Apulogalerix have distinct origins, which supports an asynchronous colonization of the island. The line of Deinogalerix possibly stemmed from some eastern species transitional between Galerix and Parasorex around Mammal Neogene (MN) zone 2. Conversely, the line of Apulogalerix originated from a primitive Parasorex ibericus, or a close relative, around MN 9–10. Another important result was detecting an impressive early Miocene (MN 2?) radiation of Galericini. Moreover, Schizogalerix and Parasorex originated from eastern Galericini morphologically transitional between Galerix and Parasorex.

We sequenced and characterized the complete mitogenome (KX964606) of the Amur hedgehog Erinaceus amurensis to provide more data for comparative mitogenomics of the genus Erinaceus (Erinaceidae). The mitogenome of E. amurensis is a circular molecule 16,941 bp long, consisting of a control region and a conserved set of 37 genes containing 13 protein-coding genes, 22 tRNA genes, and two rRNA genes (12S rRNA and 16S rRNA). The mitogenome of E. amurensis is AT-biased, with a nucleotide composition of 33.9% A, 21.1% C, 32.6% T, and 12.4% G. The mitogenomes of E. amurensis and the closely related hedgehog species E. europaeus, excluding the control region (66.7%), share over 90% sequence similarity. According to the inter-generic relationship based on six mitogenomes described from five genera of Erinaceidae, the subfamilies Erinaceinae and Galericinae are strongly supported as monophyletic groups, with each genus well placed within its own subfamily. Within the subfamily Erinaceinae, E. amurensis is a sister species to E. europaeus, and the relationship between Hemiechinus and Erinaceus is strongly supported. Within the subfamily Galericinae, the clade of Hylomys + Neotetracus was sister to that of Echinosorex, with clades supported by high values. Our findings will help to understand the codon usage pattern and molecular evolution of E. amurensis, and provide insight into inter-generic relationships within the family Erinaceidae. In future studies, the inclusion of mitogenomes from other genera would greatly enhance our understanding of higher phylogeny within the Erinaceidae.

In the present study, light and scanning electron microscopy (SEM) were used to further study the detailed morphology of Physaloptera clausa Rudolphi, 1819, based on the material collected from the Amur hedgehog E. amurensis Schrenk in China. The results revealed a few previously unreported morphological features and some morphological and morphometric variability between our specimens and the previous studies. The present supplementary morphological characters and morphometric data could help us to recognize this species more accurately.

Methods of the diagnosis of pathophysiological processes in the skulls of hedgehogs are provided. They are based on the identification of functional relationships between the development of the individual parts of skull and evaluation of the potential danger for the changes. Swelling of frontal bones is not a diagnostic feature for diagnostics of the Eastern hedgehog, but a sign (indicator) of pathology. Swelling of frontal bones is not a diagnostic feature, but a sign (indicator) of pathology. Age changes of the hedgehogs skull arch’s inner surface are analyzed.

Recent advances in understanding the major bottlenecks in derivation of engraftable HSCs and lymphoid cells from pluripotent stem cells (PSC), have identified deficiencies in NOTCH and HOXA signaling as contributing factors to the observed functional deficits of PSC-derived hematopoietic progenitors. However, little is known about the mechanisms that are essential for establishing these pathways during PSC differentiation. Here, we revealed the critical role of SOX17 in linking HOXA and NOTCH-mediated arterial programs in hemogenic endothelium (HE) and specification of definitive lympho-myeloid hematopoiesis. Using SOX17-knockout (SOX17-/-) and SOX17 DOX-inducible (iSOX17) hESCs, we found that SOX17-deficiency substantially reduces formation of CD144+CD43-CD73-DLL4+CXCR4+/- arterial HE and definitive lympho-myeloid hematopoiesis, while SOX17 upregulation at mesodermal stage of development causes the opposite effect. Molecular profiling of HE generated from iSOX17 hESCs in DOX+ and DOX- conditions using RNAseq, SOX17 ChIPseq and ATACseq, revealed that SOX17 overexpression upregulates 522 genes enriched in NOTCH, TGFb, HEDGEHOG and WNT signaling, including DLL1, DLL4, NOTCH4, LFNG, WNT5a, WNT5b, GLI3, and genes associated with HSC development, CXCR4,KITLG and ALDH1A2. In addition, we noted significant upregulation of HOXA7,HOXA9, HOXA10, HOXB8, HOXC4 and CDX2 homeobox genes in SOX17-induced cultures, with no expression of HOXA genes observed in HE from SOX17-/- cells. ChIPSeq analysis revealed DOX+ specific SOX17 binding at transcriptional start sites (TSS) of 316 significantly upregulated genes, including ALDH1A2, CDX2, DLL1, DLL4, HEY1, HOXA7, HOXB8, HOXC4 and KITLG, suggesting that upregulation of these genes could be explained by their direct activation by SOX17. Since ALDH1A2 and CDX2 are known to play a role in the activation of HOXA genes, we investigated whether SOX17's effect on HOXA expression could also be mediated by ALDH1A2 and CDX2. We found that adding ALDH1 inhibitor to DOX+ cultures had no effect on arterial HE development and HOXA expression. In contrast, transfection of iSOX17 hPSCs cultures with CDX2 shRNA significantly decreased arterial HE formation and downregulated HOXA7, HOXA9, and HOXA10 expression. Overall, our studies indicate that SOX17 plays a critical role in the activation and integration of arterial and HOXA programs in HE, which is mediated by CDX2. These findings will be important for designing a strategy for direct HSC fate programming from hPSCs. Disclosures Uenishi: Casebia Therapeutics: Employment. Slukvin:Cynata Therapeutics: Consultancy, Other: Founder and Stockholder.

Sonic Hedgehog (SHH) medulloblastoma originates from the cerebellar granule neuron progenitor (CGNP) population that depends on Hedgehog signaling for its expansion. While SHH tumors are characterized by an overall deregulation of this pathway, they also exhibit aberrations that are specific for patient age. To investigate if the developmental stage of the tumor cell-of-origin can account for these age-specific lesions, we compared transcriptomes from developing mouse CGNPs and observed highly dynamic gene expression changes as function of age. Cross-species comparison with a cohort of SHH medulloblastoma showed partial maintenance of these patterns in the different medulloblastoma patient-age groups. In particular, we found low primary cilia expression in early CGNPs and infant medulloblastoma, which coincided with insensitivity to Smoothened manipulation. Together, these findings can explain the absence of SMOOTHENED mutations in infant patients and suggest that drugs targeting the SHH pathway downstream of SMOOTHENED will be most appropriate for infant patients.

The complete mitochondrial genome of the small-toothed forest hedgehog Mesechinus miodon (Eulipotyphla: Erinaceidae) has been reconstructed from Illumina sequencing data. The circular genome is 16 842 bp long, comprising 22 transfer RNAs (tRNAs), 13 protein-coding genes (PCGs), two ribosomal RNAs (rRNAs), and one control region. All PCGs are initiated with ATR (ATA/ATG) codons, except for nad4 with GTG as its initiation codon. Two PCGs (cox3 and nad4) harbor an incomplete termination codon T, while the others are terminated with TAA (atp6, cox1, cox2, nad1, nad2, nad3, nad4l, nad5, and nad6), TAG (atp8) or AGA (cytb). The base composition is highly biased (34.9% A, 19.7% C, 11.9% G, and 33.5% T for the light strand) with an overall A + T content of 68.4%. Phylogenetic analysis indicated that M. miodon is more closely related to the consubfamilial long-eared hedgehog Hemiechinus auritus than to those within the order Eulipotyphla.

Programs that control early lineage fate decisions and transitions from embryonic to adult human cell types during development are poorly understood. Using human pluripotent stem cells (hPSCs), in the present study, we reveal reduction of Hedgehog (Hh) signaling correlates to developmental progression of hematopoiesis throughout human ontogeny. Both chemical- and gene-targeting–mediated inactivation of Hh signaling augmented hematopoietic fate and initiated transitions from embryonic to adult hematopoiesis, as measured by globin regulation in hPSCs. Inhibition of the Hh pathway resulted in truncation of Gli3 to its repressor, Gli3R, and was shown to be necessary and sufficient for initiating this transition. Our results reveal an unprecedented role for Hh signaling in the regulation of adult hematopoietic specification, thereby demonstrating the ability to modulate the default embryonic programs of hPSCs.

The Shanyangzhai Cave is located in Qinhuangdao, Hebei Province, and has yielded abundant Middle Pleistocene mammalian fossils. This is a report on the Erinaceus fossils found from this cave. The morphological character of the Erinaceus fossil species ranges between Erinaceus olgae and Erinaceus europaeus but is closer to the latter. Its p4 paraconid, located in the front of the crown, is thick and tall. The protoconid is behind and linked with the metaconid to form a horizontal ridge (protocristid). The entoconid is linked with the hypoconid by a basal ridge. The protocristid does not occur in the fossil species as E. olgae , while it is evident in the present species. The fossil species from Shanyangzhai have different protocristid from the modern E. europaeus as well. The fossil species E. europaeus should be placed between the extinct species E. olgae and the modern E. europaeus . The E. olgae may be the ancestor species of modern E. europaeus .

Although Togo is a relatively small country in West Africa, it is characterized by a wide variation of vegetation zones ranging from moist forests to arid savannahs, including the “Dahomey Gap”. There has been no comprehensive documentation of the native mammal fauna of Togo since 1893. Our review of the extant and extirpated mammals ofT ogo includes 178 species, with Chiroptera (52 species) and Rodentia (47 species) being the most speciose groups. This number does not include additional species recorded along the borders ofT ogo, and whose presence inside the country is not verified. Seven species of mammals are presumably extinct in the country but we confirmed that two species of large ungulates, reputed to be extinct, survive in remote forest habitats. Ecological Zone IV, sustaining the moist forest areas, and Ecological Zone I, inclusive of all the relatively undisturbed dry savannahs of the extreme North of the country are the most important regions for mammal diversity and conservation.