Atelerix albiventris (Wagner, 1841)

Russell A. Mittermeier & Don E. Wilson, 2018, Erinaceidae, Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona: Lynx Edicions, pp. 288-330 : 321-322

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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).

Figures

1. West European Hedgehog (Erinaceus europaeus), 2. Northern White-breasted Hedgehog (Erimaceus roumanicus), 3. Southern White-breasted Hedgehog (Erinaceus concolor), 4. Amur Hedgehog (Erinaceus amurensis), 5. North African Hedgehog (Atelerix algirus), 6. Four-toed Hedgehog (Atelerix albiventris), 7. Somali Hedgehog (Atelerix sclateri), 8. Southern African Hedgehog (Atelerix frontalis), 9. Desert Hedgehog (Paraechinus aethiopicus), 10. Brandt's Hedgehog (Paraechinus hypomelas), 11. Indian Hedgehog (Paraechinus micropus), 12. Bare-bellied Hedgehog (Paraechinus nudiventris), 13. Common Long-eared Hedgehog (Hemiechinus auritus), 14. Indian Long-eared Hedgehog (Hemiechinus collaris), 15. Wang's Forest Hedgehog (Mesechinus wangi), 16. Small-toothed Forest Hedgehog (Mesechinus miodon), 17. Daurian Hedgehog (Mesechinus dawuricus), 18. Hugh's Hedgehog (Mesechinus hughi), 19. Long-eared Gymnure (Hylomys megalotis), 20. Dwarf Gymnure (Hylomys parvus), 21. Short-tailed Gymnure (Hylomys suillus), 22. Moonrat (Echinosorex gymnurus), 23. Shrew Gymnure (Neotetracus sinensis), 24. Hainan Gymnure (Neohylomys hainanensis), 25. Mindanao Gymnure (Podogymnura truei), 26. Dinagat Gymnure (Podogymnura aureospinula)

Distribution. Senegal E to Eritrea, Djibouti, and Somalia and S through East Africa to Zambezi River.