| DINGO Canis lupus familiaris/Canis familiaris |
Includes NEW GUINEA SINGING DOG OR NEW GUINEA
HIGHLAND WILD DOG C. dingo hallstromi/C. hallstromi
HB ♀ 70.3–101cm, ♂ 75–111cm; T 20–37cm; W ♀ 8–17kg, ♂ 7–22kg
The Dingo has been classified as a distinct species, Canis dingo, or a subspecies of the Grey Wolf, C. lupus dingo, i.e. derived from a regional population of Grey Wolves in Asia independently of the lineage leading to the domestic dog (C. l. familiaris/C. familiaris). The New Guinea Singing Dog (named for its distinctive melodic howl; also known as New Guinea Highland Wild Dog) has been classified as both a subspecies of the Dingo, C. dingo hallstromi, and as a full species, C. hallstromi. A recent and comprehensive review, including genetic analyses, concluded that the 2 forms are very closely related and together comprise an ancient breed (or breeds) of domestic dog that likely arose in Southeast Asia perhaps ~10,000 years after the effective genetic separation of domestic dogs from the ancestral Grey Wolf population. Accordingly, from an evolutionary perspective, the Dingo is most correctly regarded as an archaic domestic dog that later became established in the wild. Based on fossil evidence, the Dingo colonised Australia at least 3,500–4,000 years ago, probably in association with humans; there is limited, unresolved genetic evidence that this occurred without human assistance as early as 8,300 bce, when a land bridge connected New Guinea and Australia. Regardless, Australia is now the only place where the Dingo unequivocally lives wild. In New Guinea and Southeast Asia, it is primarily associated with humans, although often in a semi-feral state. Australian Dingoes are approximately the size of a Border Collie; New Guinea Singing Dogs have a shorter body length and shorter legs, giving them a smaller, stockier appearance. Usually tawny ginger; pale sandy, white (not albino) and black-and-tan variants occur. Tail tip and paws are usually white or cream. Sable (Alsatian-like), brindled and piebald coloration typically indicates hybridisation with modern (of European descent) domestic dogs.
Distribution and Habitat
Commensal with humans in S Myanmar, Thailand, Laos, Cambodia, Vietnam, Malaysia, Indonesia, Borneo, Philippines, Sulawesi and New Guinea; wild in Australia, where it inhabits desert, grassland, woodland savannah, wetland, alpine moorland and forest. Occurs in rural habitats, including extensive livestock production areas, and in peri-urban areas provided there is sufficient habitat and prey. Wild populations mostly avoid intensive agriculture, although extensive sugar-cane monocultures with associated bushland are important habitats for peri-urban Dingoes (Queensland).
Feeding Ecology
Very broad diet; 177 prey species recorded from Australia, with mammals comprising around 75% of the diet on average. At least 1 macropod – especially Red Kangaroo, Euro, and Swamp, Agile and Red-necked wallabies – features prominently in the diet across its range. Other important prey includes wombats, brushtail possums, bandicoots, introduced European Rabbit and Magpie Goose. Small rodents, mainly Spinifex Hopping-mouse, Sandy Inland Mouse, Long-haired Rat and introduced House Mouse, dominate the diet of Simpson Desert Dingoes during rodent population irruptions. A very wide variety of other items are opportunistically eaten, usually forming a small proportion of the diet, including birds, reptiles, crocodile and turtle eggs, invertebrates, fruits and seeds. Dingoes on Fraser Island (Queensland) eat mainly small to medium-sized mammals, especially Northern Brown Bandicoot, with relatively high proportions of seeds, fruits and large skinks, as well as marine species hunted and scavenged along the shore. After bandicoot, fish – almost all scavenged from people’s fishing waste – are the second-most common dietary item for Fraser Island Dingoes. They also consume crustaceans and molluscs, and scavenge stranded marine turtles, whales, dolphins, Dugongs and New Zealand Fur Seal. Occasionally cannibalistic. Kills sheep, goats and cattle calves. Foraging is mainly nocturno-crepuscular, but diurnal where the Dingo is free from persecution. Forages alone or socially; large prey such as kangaroos are usually hunted cooperatively in packs, which increases hunting success, e.g. from 5.5% (alone) to 19% (packs) when hunting Red Kangaroos. Scavenges, including from livestock carcasses, human-killed feral animals (especially pigs and deer), human refuse and recreational fishing waste. Peri-urban Dingoes in Queensland eat pet food, but only rarely; diet is mainly natural rather than from human sources.
Social and Spatial Behaviour
Free from persecution, lives in stable packs of 2–12 adults and their pups in enduring home ranges. Territorial, but often shares important resources such as waterholes with neighbouring packs. Breeding is often restricted to the alpha pair, and other pack members help raise pups by provisioning and guarding. Under persecution (around half of its Australian range), social structure may be fractured, so that packs are smaller and less stable. Individuals associate in loose ‘tribes’, sharing a range that is not actively defended, and foraging is mostly solitary. Size of pack territories: 4–55km2 (moist, cool forest) and 32–126km2 (Simpson Desert), to more than 300km2 in Australian desert areas generally.
Reproduction and Demography
Breeding is generally seasonal, most strongly in arid C Australia. Dingoes (including New Guinea Singing Dogs) differ from modern domestic dog breeds in having only a single breeding season per year. Mating period peaks April–June; births peak June–August. Gestation 61–69 days. Litter size 1–10, averaging 5. Most females first breed at around 24 months; males sexually mature at 12 months, but breeding is limited by social dynamics. Females can breed until 11 years (wild). MORTALITY Highly variable depending on the level of human persecution; on average, approximately two-thirds of Dingoes die before 2 years from natural causes (primarily starvation and intraspecific killing, including infanticide). Persecution by people is the next most significant cause of mortality, especially for adults. Occasionally killed by Water Buffalo, Red Kangaroo and Wedge-tailed Eagle (pups). LIFESPAN 13.3 years in the wild, 20 in captivity.
Status and Threats
Widespread in Australia, where it is protected in national parks, World Heritage sites and Aboriginal reserves. Elsewhere, legally regarded as a pest and persecuted, mainly in livestock-producing landscapes, including by state-sanctioned trapping and poisoning. Persecution is intense in some areas, but Dingo populations are very resilient and capable of rapid recovery. Hybridisation from modern (European) domestic dogs threatens the integrity of the ancient Dingo genotypes, worthy of conserving in their own right (even if only for what they might reveal about the early stages of dog domestication). Pure Dingoes are most common in C and N Australia, rare/possibly extinct in S and NE Australia, and probably extinct in SE and SW areas. Without intensive conservation effort, pure Dingo genotypes are unlikely to persist except perhaps on islands (e.g. Fraser Island) or in very remote areas (e.g. Tanami Desert). Red List VU, population trend Increasing overall (including hybrids), purebreds Decreasing.
Plate 43
| GREY WOLF Canis lupus |
TIMBER WOLF, ARCTIC WOLF, TUNDRA WOLF
HB ♀ 87–117cm, ♂ 100–130cm; T 35–50cm;
SH 66–81cm; W ♀ 18–55kg, ♂ 20–79.4kg
The world’s largest canid, with significant variation in size and coloration. Largest individuals (Alaska and Canada) are 3–6 times as heavy as Middle Eastern and S Asian wolves. Typically pale to dark grey, but highly variable, e.g. ginger in E to C Asia (‘Himalayan Wolf’ or ‘Tibetan Wolf’), brown in W to N Eurasia, and white, especially in N Canada (‘Arctic Wolf’). Black coloration is rare outside forested North America, and traces to interbreeding with early domesticated dogs 10,000–15,000 years ago. Grey Wolves hybridise with Coyotes, particularly in areas where Grey Wolves, and hence conspecific mates, are rare. Grey Wolf is the progenitor of the domestic dog, and they can interbreed. Dingo treated here as a domestic dog variant, is sometimes classified as a Grey Wolf subspecies. Twelve Grey Wolf subspecies are described, most on the basis of morphology, which is unreliable. Genetically, the most distinct lineages are the Mexican Wolf (C. l. baileyi); wolves of coastal SE Alaska, usually recognised as C. l. ligoni; and Himalayan Wolves (C. l. himalayensis; Himalayas to Tibetan Plateau), which some scientists treat as a separate species, C. himalayensis.
Distribution and Habitat
North America and Eurasia, formerly one of the most widespread mammals. Widespread in Canada and Alaska, extirpated in the lower USA except in the N Rockies and N Midwest, and relict in SW USA–Mexico, where reintroduction has established 100–110 Mexican Wolves. Widespread in Russia and C Asia, fragmented and reduced in SW Asia, the Middle East, and W and N Europe. Occupies many habitat types, including desert (to 50°C), open plains, steppe, mountainous areas, swamps, forest and Arctic tundra (to –56°C). Although tolerant of habitat modification, rarely inhabits agricultural and pastoral areas due to intense anthropogenic persecution.
Feeding Ecology
Highly opportunistic and proficient pack hunter. Diet varies extensively by region and season, but medium to large ungulates are usually the mainstay, e.g. Musk-ox, bison, Moose, Elk, Red Deer, Reindeer/Caribou, White-tailed Deer, roe deer, wild sheep, ibex and Wild Boar. Juvenile and debilitated individuals are most often killed, but even a single Grey Wolf is capable of killing healthy adults, especially during winter. When ungulates are less vulnerable (spring–summer), the diet is more diverse, with increased consumption of beavers, hares, rodents, waterfowl, fish and fruits. Kills smaller carnivores, especially Coyotes, and occasionally Pumas and young Black Bears (here), Brown Bears and (exceptionally) Polar Bears. Livestock is readily killed, especially during spring–summer, when wild prey disperses and stock occupies productive grazing areas; livestock (including semi-domestic Reindeer) is the most important prey for many Eurasian populations. Unprovoked attacks on humans are very rare and usually attributed to rabies, e.g. 2 fatalities by healthy wolves in North America since 1900. Hunting is cathemeral, with greater diurnalism where it is protected. Hunting is highly social; pack members exchange the lead in chases and cooperatively bring down large prey. Can reach 64km/h and has extraordinary endurance, maintaining pursuit to 8km (with an exceptional record of 21km). Hunting success rates 10–49% for packs (North America). Scavenges, including from refuse dumps and appropriated carcasses from other carnivores.
Social and Spatial Behaviour
Highly social and territorial, living in packs numbering up to 42 but typically 2–15. Nucleus of the pack is a mated adult pair, accompanied by adult offspring. Pack size fluctuates depending on dispersal of offspring, which is affected by food availability. Dispersal is low in productive years, producing large packs with up to 4 generations of grown offspring. Most offspring ultimately disperse and seek non-related adults to form new packs. Recruitment of unrelated individuals (especially to replace a lost breeder) or small groups of dispersers occurs occasionally in established packs. Packs occupy enduring ranges that are usually defended aggressively from other packs. Range size 33–4,335km2, averaging 69–2,600km2. Ranges increase locally during winter and with increasing latitude; largest ranges recorded are from Alaska and the Canadian Arctic. Wolves following migratory herds have massive ranges of 63,000–100,000km2 annually that are not defended. Density estimates 5/1,000km2 (NW Alaska) to 92/1,000km2 (Isle Royale, Canada), but rarely exceeding 40/1,000km2.
Reproduction and Demography
Seasonal. Mating January–April depending on latitude, with pups born March–June. Gestation 60–75 days, typically 62–65. Litter size 1–13, averaging 4–7. Pack’s dominant female is normally the only breeder, although multiple females (probably close relatives) occasionally breed under high prey availability. All pack members help raise pups by provisioning the mother and pups at the den, and by defending pups from predators. Weaning at around 8–10 weeks. Pups leave the den permanently and travel continually with the pack from 4–6 months by September–October. Dispersal is usually at 11–24 months, but is recorded at 5–60 months. Inter-litter interval typically 12 months; 2 litters a year produced on rare occasions. Sexual maturity at around 10 months for both sexes, although breeding opportunities rarely arise before 3 years. MORTALITY Annual pup mortality averages 34%, ranging from 9% (Denali NP, Alaska) to 61% (N Wisconsin). Adult mortality from 14%, to 44% where heavily exploited. Human hunting and trapping are often the main cause; in unexploited populations, starvation (mostly of pups) and aggression from other wolves are the main factors. Disease is an important cause of death, although population effects are poorly known. Wolves occasionally die in hunting accidents and are killed by bears, Puma and Amur Tiger. LIFESPAN 13 years in the wild, 17 in captivity.
Status and Threats
Widespread and stable in most of its northern range, especially in Alaska, Canada, Kazakhstan and Russia (total combined estimate 113,000–127,000 wolves). None the less, it has lost an estimated third of its historic range, mainly in the USA/Mexico, W Europe and S Asia, where it is mostly threatened or endangered. Chief threat is persecution by humans, often as part of state-sanctioned control programmes. Diseases, especially canine parvovirus, mange and rabies produce local declines. Reintroduction of 31 individuals to Yellowstone NP in 1995–96 has been highly successful, growing to approximately 1,900 animals in 6 states in 2016; reintroduction to the SW USA (Mexican Wolf subspecies) has been less successful. Legally hunted and trapped in at least 15 countries, with the largest numbers killed in Canada (4,000 annually), Russia (10,000–20,000 annually) and Mongolia. CITES Appendix I – Bhutan, India, Nepal and Pakistan, Appendix II – elsewhere; Red List LC, population trend Stable.
Plate 44
| COYOTE Canis latrans |
BRUSH WOLF, PRAIRIE WOLF
HB 74–95cm; T 26–46cm; W ♀ 7.7–14.8kg (exceptionally to 25kg), ♂ 7.7–18.1kg (exceptionally to 34kg)
Generally uniformly coloured, from frosted grey to rufous-brown, with pale underparts and often a greyish or ‘salt-and-pepper’ saddle. Tail is typically infused with dark brown to black hairs, and usually dark-tipped (rarely white-tipped). Varies very widely geographically and seasonally; northern and winter individuals are generally larger and paler/greyer. Rufous sides, outer legs and backs of ears are common in eastern and southern (including Central American) individuals. Melanistic, leucistic, brindled, sable (Alsatian-like) and blond individuals occur, particularly in eastern North America, where admixture with domestic dogs is most prevalent (‘Eastern Coyote’).
Distribution and Habitat
N Alaska and Canada (except NE Canada), throughout the USA and Meso-America to Panama. Recorded east of the Panama Canal for the first time in 2013; not confirmed from Colombia (as of 2017). At the time of European settlement, Coyotes were restricted to non-forested W and C North America. Habitat conversion, with a concomitant increase in White-tailed Deer and extirpation of Grey Wolves, fostered their expansion across virtually the entire North American continent and Meso-America in 200 years. Now inhabits virtually all habitat types, from Arctic tundra to deserts, tropical and montane forest, and heavily modified anthropogenic habitats, including all kinds of agricultural land and suburban–urban areas, even in large cities, e.g. Chicago and New York.
Feeding Ecology
Extremely opportunistic generalist that consumes virtually any edible food source. Vertebrate prey is most important, mainly rodents (chiefly squirrels, mice and voles), lagomorphs, juvenile ungulates and carrion. A wide variety of other small to medium-sized mammals are also taken incidentally, including armadillos, Virginia Opossums and mesocarnivores, e.g. Northern Raccoons, Striped Skunks, foxes and Bobcats (probably juveniles). Capable of killing adult large ungulates, including 4 recent records of 2–5 Coyotes killing Moose older than 20 months (Ontario, Canada), but large mammals are rarely killed, and debilitated individuals, such as those weakened in deep snow, are usually targeted. Scavenging of winter- and wolf-killed large ungulates is particularly important to northern populations, e.g. scavenged Moose is the main dietary item in Cape Breton Highlands NP, Canada. Mammalian prey is always supplemented by a wide variety of other food sources, especially fruits, seeds, vegetables, grains (including crops), eggs and invertebrates. Diet varies very widely regionally and seasonally. Coyotes adjust intake of different food types depending on local availability, for example, different fruits including blackberries, wild plum, wild grape, black cherry and persimmon peak at different times in the diet of Coyotes in Longleaf Pine habitat (Joseph W. Jones Ecological Research Center, Georgia) as the fruiting season progresses. Birds are typically ancillary prey; waders, shorebirds and waterfowl form a large part of the diet on protected islands off the coast of South Carolina. Coyotes take small livestock (goats, sheep and young calves) and poultry. Domestic pets, especially cats and less so small dogs, are eaten rarely; cats occur in 0–2% of scats collected in urban and suburban studies. Attacks on people are very rare and not as prey; attacks occur most often during the breeding and pup-rearing season, when adults are very defensive. There are 2 recorded human fatalities, a 3-year-old child (California, 1981) and an adult woman killed by a pair (Cape Breton Highlands NP, 2009). Foraging is mainly diurno-crepuscular, but generally nocturnal near humans, particularly where hunted. Usually forages alone, less often in pairs. Where large ungulates are available, Coyotes may hunt socially, usually as pairs or small family groups; typically the alpha pair is responsible for the attack, and younger animals play little part. Sometimes associates with foraging American Badgers, snatching rodents flushed from cover or burrows as the badger digs. Large groups, including non-relatives, congregate relatively amicably at large carcasses. Scavenges, including from carnivore (especially Grey Wolf) kills, human refuse, pet food and bird feeders.
Social and Spatial Behaviour
Sociality is extremely flexible, changing regionally and temporally depending on food availability. Basic social unit is a territorial monogamous pair that may breed for life. ‘Associate’ individuals, usually grown offspring of previous litters, remain with the pair under high food availability, forming packs numbering up to 10. Associates help raise pups and defend territories, although not to the extent of the breeding pair. Large packs occur where ungulates are the main prey, while pairs and trios are typical where prey is small. Some Coyotes never join packs and live as solitary nomads. Average resident territory size (excluding small breeding ranges) 2–3km2 (SW USA) to 42–61km2 (Minnesota). Territories in urban habitats range from 5–36km2 (resident) to 27–115km2 (transient). Density varies very widely, depending on habitat, season, protection and Grey Wolf density, e.g. 1–9/100km2 (Alberta) to 150–230/100km2 (Texas); urban populations reach extremely high densities, e.g. up to 600/100km2 in Chicago.
Reproduction and Demography
Seasonal. Mating January–March, with pups born April–June. Gestation 58–65 days. Maximum litter size 11, averaging 4–7. Pups weaned at 5–7 weeks; dispersal from 6 months. MORTALITY Humans are the main cause of death, mainly from shooting or trapping; collisions with vehicles comprise 35–62% of mortalities in urban areas. Grey Wolf is the chief natural predator. Annual mortality (adults and subadults) is lower in protected and urban populations (20–47%), compared to trapped and hunted populations (50–70%). LIFESPAN 15.5 years in the wild (rarely beyond 8), 18 in captivity.
Status and Threats
Extremely widespread, common and resilient. Coyotes are heavily hunted, trapped and persecuted across much of their North American range, but populations are very resistant to persecution. Official control actions, largely in response to complaints of livestock depredation, killed more than 77,000 Coyotes in the USA in 2016. Very tolerant of habitat modification; deforestation to open habitat actually favours Coyotes. Combined with extirpation of Grey Wolves (the main natural predator and competitor), such habitat conversion has enabled colonisation outside its historic range, e.g. E Canada, E USA and E Central America, which continues today. Red List LC, population trend Increasing.
Plate 45
| GREY WOLF–COYOTE HYBRIDS |
The canids depicted opposite have all been (and sometimes still are) classified as discrete species in eastern North America. Recent (2016) compelling genetic evidence indicates that all are hybrid populations of Coyote and Grey Wolf. The analysis suggests that Coyotes and Grey Wolves diverged from a common ancestor very recently, within the last 117,000 years, making them sufficiently closely related to interbreed occasionally and produce fertile offspring. The amount of admixture correlates with Grey Wolf declines in eastern North America following European colonisation; this opened up the large canid niche to Coyotes and allowed them to expand eastward into the void. Faced with reduced opportunities for reproduction, remaining Grey Wolf individuals mated with Coyotes, and their fertile hybrid pups formed part of an expanding Coyote population that rapidly colonised former Grey Wolf distribution. Accordingly, continuous, large populations of Alaskan Grey Wolves almost never hybridise, and average 8–8.5% Coyote ancestry. Wolves from Algonquin Provincial Park, Ontario, thought to be pure Eastern Wolf (see below), actually have 32.5–35.5% Coyote ancestry, while Red Wolves in CE USA (see below), where human pressures on Grey Wolves were early, intense and sustained, have 70–80% Coyote ancestry – they are now more Coyote than Grey Wolf.
Coyotes and Grey Wolves continue to exchange genetic material today, although hybridisation is most prevalent where and when wolf persecution is intense and Coyote populations expand; there is little evidence of ongoing hybridisation in some populations with mixed ancestry where persecution has relaxed. Wolves of the western Great Lakes region are recovering from historic low numbers around a century ago, when their Coyote ancestry (21.7–23.9%) is assumed to have largely arisen. Great Lakes Grey Wolves today are not limited in mating opportunities with other Grey Wolves (strictly speaking, mostly Grey Wolf, from a genetic perspective), and are more likely to kill Coyotes rather than breed with them.
Grey Wolf–Coyote hybrid populations not only reveal the intriguing challenges in defining species (see the box ‘How many species of carnivores?’); they also present a quandary for conservationists. Does having mixed ancestry mean they do not warrant conservation efforts? They are fascinating in their own right and have similar aesthetic and ecological value to ‘true’ species. They fulfil an important regulatory role as the major predators of super-abundant White-tailed Deer, a species that causes hundreds of human deaths and tens of thousands of injuries through road accidents, as well as an estimated US$3.5 billion in damage to crops, nursery plants and tree seedlings each year across eastern North America. Medium-sized wolf-like canids such as the Red Wolf and Eastern Coyote defy traditional categorisation as species, yet they are better suited to modern anthropogenic eastern North America than the original Grey Wolf occupants, which need vast wilderness areas with very large ungulate prey.
| EASTERN COYOTE Canis latrans var. (Canis ‘oriens’) |
See Coyote for species account
Advocates have proposed that the Coyote in NE North America warrants recognition as a unique species, Canis ‘oriens’ (often referred to as ‘Coywolf’), with little validity. Eastern Coyotes tend to be larger than western Coyotes, but there is continuous variation in size from west to east; they are not significantly genetically differentiated nor reproductively isolated from western or southeastern Coyotes. The Eastern Coyote is essentially a Coyote with some Grey Wolf genes that, in the absence of competing Grey Wolves, has undergone natural selection for a slightly larger ecotype to exploit White-tailed Deer more effectively. As well as having Grey Wolf ancestry, Eastern Coyotes have hybridised with domestic dogs. All sampled individuals are mainly Coyote, with some that have almost no Grey Wolf genes, but none is exclusively Coyote, and none completely lacks dog genes. The mix of genes gives rise to wide morphological variation, including coat colour variants such as brindled, sable (Alsatian-like) and blond (depicted opposite), rarely seen in other populations.
| RED WOLF Canis ‘rufus’ |
HB ♀ 99–120cm, ♂ 104–125cm; T 30–46cm; SH 66–76cm; W ♀ 16–30kg, ♂ 21–41kg
The Red Wolf’s status is controversial. There is ongoing debate over whether hybridisation was pre-historic (~10,000 bce) or since European colonisation; and some taxonomists suggest it is not a hybrid but instead evolved very recently from a common ancestor with the Coyote. Regardless, its biological profile – updated from the first edition of this book – is included here to provide ecological data broadly representative of the canid hybrid populations now inhabiting eastern North America.
Reddish fur, becoming pale ginger to cream on the lower limbs, and distinctive white throat and chest patches. Distribution and Habitat Extinct in the wild except for a reintroduced population occupying 6,000km2 in E North Carolina, USA, which inhabits pine forest–wetland mosaics, marshland and agricultural land with cover. Feeding Ecology Eats mainly White-tailed Deer, raccoons, Marsh Rabbit and small rodents. Livestock is eaten, although recent records are all of carrion. Hunting is nocturno-crepuscular, and usually in small packs or singly. Social and Spatial Behaviour Forms small family packs of 2–12 animals, comprising a dominant breeding pair and its offspring. Packs are territorial and occasionally kill unrelated intruders. Females disperse at higher rates than males, and some individuals never disperse, remaining in the pack as non-breeding helpers. Resident pack range size averages 68.4km2 (range, 25–190km2); ranges for transients are much larger, average 319km2 (range, 122–681km2). Reproduction and Demography Seasonal. Mating February–March; births April–May. Gestation 61–63 days. Litter size averages 3–5, exceptionally to 10. MORTALITY Rates low for canids: 32% (pups), 21% (yearlings) and 19% (adults) annually. Shooting and roadkills are the main causes; natural deaths account for a quarter of the total, mainly from intraspecific killing, sarcoptic mange and starvation (of pups). LIFESPAN 20 years in captivity. Status and Threats Treated as an endangered species by the US government and declared Extinct in the Wild by 1980. Reintroduced from captivity in 1987 and formerly numbered up to 130; there are currently fewer than 50 in the wild, including pups (2018). Humans are the main threat. Given recent genetic data, its conservation status is uncertain; it is still (2018) treated as a distinct species by the US Fish and Wildlife Service. Red List CR, population trend Decreasing.
| EASTERN WOLF Canis lupus var. (Canis ‘lycaon’) |
See Grey Wolf for species account
The Eastern Wolf has been variously classified as a distinct species, Canis ‘lycaon’, the same species as Red Wolf, a hybrid between Red Wolf and Grey Wolf, or (as in this book) a Coyote-admixed population of the Grey Wolf. Historically, its range was defined as S Quebec and Ontario, centred on Algonquin Provincial Park, although this is contiguous with Grey Wolf distribution across E Canada. Algonquin Provincial Park wolves have historically been heavily hunted by people (including official culling programmes as recently as the 1960s; they have been protected since), and wolves are legally hunted and trapped throughout the area considered Eastern Wolf range. Resulting hybridisation with Coyotes is typical of heavily hunted wolf populations; this declines along a cline north and west into more continuous and less hunted Grey Wolf range in Canada.
Plate 46
| EURASIAN GOLDEN JACKAL Canis aureus |
GOLDEN JACKAL, ASIATIC JACKAL, INDIAN JACKAL, COMMON JACKAL
HB ♀ 69–85cm, ♂ 70–90cm; T 20–38cm; SH 35–48cm; W ♀ 4.9–13.6kg, ♂ 6.7–14.5kg
The only jackal in Eurasia. Until 2011, it included the species now classified as the African Wolf. It is now uncertain whether the Eurasian Golden Jackal occurs anywhere in Africa; there is a possible hybrid zone from Egypt to Israel. Uniformly tawny grey to greyish brown, without discrete markings except a greyish mantle in some populations; bushy tail usually has a dark tip (never white). Melanism occurs very rarely and may be the result of hybridisation with domestic dogs. Distribution and Habitat Arabian Peninsula to SE Europe, and C Asia to Indochina; may occur in Egypt. Inhabits semi-desert, grassland, dry woodland, forest, and agricultural and semi-urban areas. Feeding Ecology Omnivorous, eating mainly rodents, hares, lizards, snakes, invertebrates, fruit, seeds, mast and vegetable matter, including crops. Small fawns e.g. of Chital and rarely to Sambar size, are taken. Birds are usually ancillary prey but dominate diet locally or seasonally, e.g. mostly migratory egrets, Garganey and coucals are the main winter prey in Patna Bird Sanctuary, India. Jackals inhabiting a mangrove island in the Gulf of Kachchh, India, eat almost exclusively crabs and fish. Takes small stock and poultry, these dominating the diet (kills and carrion combined) in some sites, e.g. Greece and Israel. Foraging is mainly nocturnal, and crepuscular/diurnal where protected. Hunts singly, in adult pairs or in small family groups. Caches surplus food in shallow holes, and scavenges: congregations of up to 18 adults are recorded at large carcasses and in dumps. Social and Spatial Behaviour Breeding pair is the main social unit, often accompanied by grown helpers from previous litters. Pairs are usually formed during the breeding season, but persist year-round under high food availability. Pairs defend a core territory centred on dens, and may defend larger territories under high food availability. Territories 3–30km2 (India). A collared female in Thailand occupied 151km2 in mostly agricultural habitat. Densities 0.1–1.5/km2 (Greece, Hungary, Romania) to 15/km2 (very high prey availability, with no large carnivores, Keoladeo NP, India). Reproduction and Demography Seasonal; births often coincide with peak food supply, such as birth flushes of ungulates or rodents, e.g. December–May (Israel) and April–June (India, C Asia). Gestation 63 days. Maximum litter size 8, typically 3–6. MORTALITY Poorly known; sometimes killed by large cats, Grey Wolf and domestic dogs. LIFESPAN 16 years in captivity. Status and Threats Generally common and widespread. Slowly declining outside protected areas in some countries, but expanding range in Europe (possibly due to historic Grey Wolf declines), most recently into Austria, Czech Republic, Denmark, Estonia, Germany, Italy, Latvia, Lithuania, Poland and Switzerland. Very tolerant of human activities and converted landscapes, but disappears under agricultural intensification (often with associated use of poisons) and urbanisation. Vulnerable to transmission of disease, especially rabies and distemper by feral dogs in anthropogenic landscapes. CITES Appendix II – India; Red List LC, population trend Increasing.
| AFRICAN WOLF Canis lupaster |
AFRICAN GOLDEN WOLF, GOLDEN WOLF
HB ♀ 74–100cm, ♂ 76–105cm; T 20–26cm; SH 38–50cm; W ♀ 6.5–14.5kg, ♂ 7.6–15.5kg
Prior to 2011, misclassified as a Golden Jackal subspecies. Morphological and genetic analyses has shown that the canid previously assigned to Golden Jackal in Africa is a distinct species, Canis lupaster (sometimes called C. anthus), which is more closely related to Grey Wolf. Western and northern African Wolves are larger, more variable in appearance, and tend to be more ‘wolf-like’ than E African individuals, which closely resemble Eurasian Golden Jackal. Yellow-grey to greyish brown, with pale underparts, breast shield and throat, often with a greyish mantle, and russet or ginger undertones on upperparts. Tail is usually interspersed with blackish hairs, especially dorsally, and has a black tip (never white). Distribution and Habitat Former distribution of Golden Jackal in Africa is now assumed to be African Wolf range: N Tanzania to Senegal, throughout C, W and N Africa. Very wide habitat tolerance; inhabits semi-desert, grassland, woodland, forest, and agricultural and semi-urban habitats. Penetrates deeply into very arid habitat in the Sahara, where it is associated with massifs, vegetation or the presence of livestock. Recorded from sea-level to 3,800m (Bale Mountains, Ethiopia). Feeding Ecology Omnivorous and highly opportunistic, with a broad diet. Capable of killing ungulates to the size of adult Thomson’s Gazelle, but most kills are of small vertebrates to the size of hares. Common prey includes mice, rats, mole rats, gerbils, cane rats, ground squirrels, hares, reptiles, birds and eggs. In Algeria, Wild Boar comprises 20% (Djurdjura NP) to 41% (Tlemcen HR) of diet by biomass, presumably mainly scavenged although piglets may be hunted. Barbary Macaque, Dama Gazelle, Red Fox and domestic cat are also recorded from scats in Algeria. Invertebrates, especially dung beetles, grasshoppers, locusts, termites and larvae, are readily consumed. Eats fruit, seeds and vegetable matter, including cultivated crops. Takes small stock; Ethiopian pastoralists in the Guassa highlands consider it to be the main livestock predator, mainly of sheep, although dietary analysis found sheep to be only rarely consumed. Foraging is mainly nocturnal, especially where persecuted, but frequently crepuscular/diurnal where protected. Forages singly or in small family groups; larger prey is more often taken by groups. Caches surplus food in shallow diggings, and scavenges from carcasses and human refuse; individuals in Ethiopian montane farmlands take rodents caught in human hunters’ traps. Congregates at large carcasses and in dumps. Social and Spatial Behaviour Known mainly from E Africa (especially Serengeti NP, Tanzania), when studied as Golden Jackal. Breeding pair is the main social unit, often accompanied by grown helpers from previous litters. Pairs typically form during the breeding season, but persist year-round in good conditions. They defend a core territory around dens, and may cooperate to establish larger territories, depending on food availability. Territory size 1.1–20km2, with densities of up to 4/km2 (Serengeti NP). Densities in Guassa highlands are 0.03/km2 (natural bushland) to 0.31/km2 (human dump site). Reproduction and Demography Seasonal, with births coinciding with peak food supply, e.g. December–April (E Africa). Gestation 63 days. Maximum litter size 8, typically 3–6. MORTALITY Poorly known; sometimes killed by Lions, Leopards and domestic dogs. Humans are likely the main cause of mortality for many populations; Ethiopian pastoralists (Guassa highlands) kill them by plugging den entrances with stones. LIFESPAN 16 years in captivity. Status and Threats Generally common, although declining outside protected areas, where persecution is intense. Very tolerant of human activities and prospers in livestock and farming areas despite often high levels of persecution. They have been extirpated where poisoning (usually illegal) occurs. Red List NE, population trend Unknown.
Plate 47
| DHOLE Cuon alpinus |
ASIATIC WILD DOG
HB 80–113cm; T 32–50cm; SH 42–55cm; W ♀ 10–17kg, ♂ 15–21kg
Superficially resembles a large jackal. Pale tawny brown to rich russet-brown with a dark-tipped tail. Northern temperate individuals are usually more reddish, with contrasting bright white underparts. Distribution and Habitat W and S China, Nepal, Bhutan, India, Southeast Asia, Sumatra and Java; patchily distributed in N China; uncertain in SE Russia and C Asia, where there are no confirmed records for >30 years. Inhabits forest, forest–grassland mosaics and montane scrubland. Avoids open habitat, and agricultural and pastoral areas. Feeding Ecology Pack hunter that mostly kills ungulates, with Chital, Red Muntjac and Sambar preferred across much of its range; also Blackbuck, Nilgai, Swamp Deer, Gaur, Asiatic Buffalo, Banteng, Blue Sheep, Markhor, Himalayan Tahr, gorals and Wild Pig. Juveniles are often selected, but packs are capable of killing adults of all but the largest species. Large Indian Civet was a preferred prey species during a short-term study in Salakpra WS, Thailand. Livestock is sometimes killed, especially when unguarded and where natural prey is depleted. People in SE Thailand blame Dholes for most losses of free-roaming poultry, possibly due to confusion with Eurasian Golden Jackals. Foraging is usually diurno-crepuscular and cooperative. Scavenges, including kleptoparasitism from other carnivores, e.g. Leopard. Social and Spatial Behaviour Lives in packs of 2–15 adults and their pups, exceptionally totalling 30 individuals. Packs have a dominant breeding pair and are biased towards males because females disperse more often. Packs occupy defined home ranges, though the extent of territorial defence is unknown. Pack range estimates (telemetry) 12–49.5km2 (mixed evergreen forest, Phu Khieo WS, Thailand) and 26.1–202.8km2 (dry forest, Pench NP, India). A pack of 6 in lowland forest used 100km2, as estimated by camera-traps (Khao Ang Rue Nai, Thailand). Reproduction and Demography Seasonal. Breeding October–April (India) and January–May (Java). Gestation 60–63 days. Litter size 4–12. Breeding usually restricted to the dominant pair, although multiple females occasionally breed and subordinate males sometimes mate with the alpha female. Pack members assist reproduction by guarding, provisioning at the den and regurgitating food to pups. A monitored pack in Baluran NP, Java, denned in burrows on steep slopes with dense vegetation and moved pups to new dens every 2 weeks on average. MORTALITY Poorly known; Tiger and Leopard are known predators. LIFESPAN 16 years in captivity. Status and Threats Endangered and declining, chiefly from persecution, habitat loss and prey declines due to human hunting. Extirpated from at least 75% of former range. Likely extinct in its former E and C Asian range, and in Southeast Asia restricted to large protected areas. S and C India is the species’ stronghold, where it reaches high densities in small protected areas; reasonably tolerated in non-protected areas, where it is generally not significantly involved in conflict with humans or hunted for illegal trade of body parts. CITES Appendix II; Red List EN, population trend Decreasing.
| ETHIOPIAN WOLF Canis simensis |
SIMIEN JACKAL, ABYSSINIAN WOLF, SIMIEN FOX
HB ♀ 84.1–96cm, ♂ 93–101.2cm; T 27–39.6cm; SH 53–62cm; W ♀ 11.2–14.2kg, ♂ 14.2–19.3kg
Rich tawny rufous with white underparts and bright white markings on the lower face, throat, chest and lower legs. Tail has a white base, darkening to a chocolate-brown tip. Hybrids with domestic dogs have a stockier build and lighter, duller coat. Despite its confusing array of common names, the species is most closely related to Coyote and Grey Wolf. Distribution and Habitat Restricted to 7 isolated populations at 3,000–4,500m in Ethiopia. Inhabits open highland habitats, especially montane grassland, heath and shrubland. Avoids agricultural areas, which reach 3,500–3,800m in parts of its range. Feeding Ecology Feeds almost exclusively on small diurnal mammals, especially mole rats, rats and Starck’s Hare. Infrequent prey includes Rock Hyrax, juvenile Grey Duiker, reedbucks and Mountain Nyala, as well as birds such as Blue-winged Goose juveniles and francolins, and their eggs. Foraging is largely diurnal and solitary, with most kills made by individual wolves stalking rodents or digging them from burrows. Small packs of 2–4 sometimes cooperatively pursue prey, especially hares and young antelopes. Rarely kills sheep lambs; does not kill cattle calves, and often forages among herds and Gelada Baboon troops, which may assist hunting by providing cover and flushing rodents. Appropriates kills from raptors and scavenges, including from livestock carcasses. Caches surplus food in shallow holes. Social and Spatial Behaviour Forms packs of 2–13 adults that defend small, stable territories from other packs. Pairs or small packs occur where prey availability is low. Males rarely disperse, so packs contain up to 8 related adult males, as well as 1–3 adult females that may or may not be related; some females remain in their natal pack, while others disperse for breeding opportunities. Average territory size ranges from 6km2 in productive habitat to 13.4km2 in poor habitat. Estimated densities include 0.1–0.25/km2 in poor habitat or unprotected areas, to 1–1.2/km2 in optimum protected habitat. Reproduction and Demography Seasonal. Mating August–November; births October–January. Gestation 60–62 days. Litter size 2–6. Reproduction is largely by the pack’s alpha pair, but the dominant female also mates with visiting males from neighbouring packs. All pack members provision pups at the den, and subordinate females sometimes assist in suckling (it is unclear if extra nursing females are pseudo-pregnant or absorb/abandon their own litters). Pups weaned from 10 weeks, and accompany the pack from 6 months. Sexual maturity at 18–24 months. MORTALITY Most mortality is anthropogenic and natural factors are poorly known; predation has not been observed, but may occur on pups by Spotted Hyaena, African Wolf and large eagles. LIFESPAN 12 years in the wild. Status and Threats Endangered, with approximately 360–440 adults (>1 year old), of which <250 are breeding adults in 7 disjunct populations. Extreme pressure on habitat for agriculture and livestock is the chief threat, combined with exotic disease, especially rabies and distemper, from domestic dogs. More than half of the remaining Ethiopian Wolves live in the Bale Mountains, where numbers have declined by ~30% since 2008 (to approximately 210 adults >1 year old) due to successive disease epidemics. Roadkills, persecution and hybridisation with domestic dogs (mainly in the Bale Mountains) are lesser threats. Red List EN, population trend Decreasing.
Plate 48
| AFRICAN WILD DOG Lycaon pictus |
PAINTED DOG, CAPE HUNTING DOG
HB 76–112cm; T 30–42cm; SH 61–78cm; W ♀ 17–26.5kg, ♂ 21–36kg
Africa’s largest canid, with coarse fur coloured a mottled patchwork of tawny, black and white. Tail has a conspicuous white tip, extending variably along the length; some individuals have an entirely white tail. Very rarely, the tail is tipped black. Coloration varies very widely within and between populations; southern African individuals tend to be tawnier, while E and NE African animals tend to be darker, but colour is not reliable for identifying origin. Coloration is unique to each individual, allowing identification for life. Little sexual dimorphism; males are slightly heavier than females and recognisable by a prominent penile sheath. There is little regional variation in size.
Distribution and Habitat
Sub-Saharan Africa, mainly in E and southern Africa, with scattered populations across Sahelian W and C Africa. Reaches highest densities in woodland savannah, but occurs widely in open grassland, semi-desert and scrubland. Absent from C African forest, but inhabits forest patches in E Africa. Penetrates deeply into true desert, but cannot permanently colonise very arid areas. Tolerant of habitat modification, but rarely inhabits pastoral landscapes due to intense persecution.
Feeding Ecology
Highly efficient pack hunter capable of taking prey as large as adult zebras and female African Buffaloes, but mostly kills medium-sized antelope species; each population focuses on 1–2 of the most common locally available species. Typical prey includes Impala, Nyala, Red Lechwe, Thomson’s Gazelle and Blue Wildebeest (usually to subadult size). Largest preferred prey is Greater Kudu, in which adult females (135kg) are commonly killed (Namibia). Able to switch to smaller prey, especially where large species are absent or in low numbers, e.g. outside protected areas in N Kenya, Kirk’s Dik-dik (3–7kg) comprises 70% of prey. Bushbuck, duikers, Steenbok and Warthog are also important prey in some areas. Opportunistically kills smaller prey, including hares, small carnivores (e.g. Bat-eared Fox) and reptiles, but these form an insignificant proportion of the diet. Kills small livestock, but depredation is rare where wild prey is available, even when it is heavily outnumbered by stock. No records of predation on humans. Hunting is almost always diurnal and highly social, often preceded by a frenzy of greeting between pack members. Reaches speeds of 66km/h and has terrific endurance, with chases extending for 2km. Hunts are highly coordinated and cooperative; pack members fan out and run in relays to maximise opportunities for capture, sometimes yielding more than 1 kill per hunt. Prey is killed cooperatively, usually by many pack members after capture by 1 dog; large and dangerous prey, e.g. Warthog and wildebeest, is often restrained by the head while other dogs disembowel and dismember it. Although this appears cruel (contributing to pastoralists’ hatred for the species), the prey usually dies within 2–4 minutes. Hunts have high success rates, 42–70%, which increase with the number of adults present. Occasionally scavenges, including appropriating carcasses from other packs, Leopard, Spotted Hyaena and (very rarely) Lion.
Social and Spatial Behaviour
Intensely social, with pack members in almost constant association. Packs form around the dominant breeding pair, with up to 28 adults, but normally average 5–10 adults accompanied by yearlings and pups; the pack size occasionally exceeds 50, including pups old enough to travel. Same-sex adults in the pack are usually related to one another but not to opposite-sex adults. Packs usually form through interchange of same-sex subgroups, typically dispersing littermates that join opposite-sex subgroups. Both sexes may disperse, with females dispersing sooner but settling nearer to their natal range than males. Occupies enduring defined ranges that are often very large and overlap those of other packs. Active territorial defence is infrequent, but occurs in overlapping areas and around den sites, where inter-pack encounters are aggressive and sometimes fatal. Range size 150–2,460km2, averaging 423–1,318km2 and dropping to 50–260km2 when young pups are in the den. Due to its wide-ranging behaviour, the species naturally occurs at low densities that fluctuate significantly depending on pup survival, e.g. 5/1,000km2 (semi-arid savannah, N Botswana), 2.8–22.5/1,000km2 (dry savannah, N Kruger NP), 16–24/1,000km2 (Selous GR, Tanzania) and 19–39/1,000km2 (mesic woodland, S Kruger NP).
Reproduction and Demography
Weakly seasonal. Pups born year-round, but most litters coincide with peak prey availability, e.g. March–June (Serengeti NP, Tanzania) and April–September (Kruger NP). Gestation 69–73 days. Litter size typically 10–11, exceptionally to 21. Pack’s dominant female is usually the only breeder. All pack members cooperate to help raise pups by provisioning the mother at the den, regurgitating food to the pups and guarding the den. Helpers are essential for raising litters, and small packs (<4 adults) rarely reproduce successfully. Subordinate females occasionally breed, but their pups survive only when prey availability is high (they are sometimes ‘stolen’ and raised by the dominant female); otherwise, they are frequently killed by the dominant female or die from starvation brought about by harassment of subordinate mother(s). Weaning is at around 8 weeks, and pups emerge from the den at around 12–16 weeks, after which they travel with the pack. Dispersal occurs most often at around 21–22 months for females (range 13–31), and 28 months for males (range 17–43). Inter-litter interval averages 12–14 months. Sexual maturity is at around 2 years for both sexes, but breeding usually occurs with social dominance at around 4–5 years. MORTALITY 25% (Selous GR) to 65% (Kruger NP) of pups die in their first year. Adult mortality 23–28% for prime adults, rising to around 50% for older adults. Main natural causes are Lions, other African Wild Dogs and infectious diseases such as rabies and canine distemper. LIFESPAN 11 years in the wild, 16 in captivity.
Status and Threats
The species has undergone a drastic loss of range and is now extinct in at least 25 of 39 original range countries. Total numbers are estimated at 6,600 adults in 39 subpopulations, of which only 1,400 are mature individuals. African Wild Dogs were actively destroyed by wildlife managers until the 1970s; they are now protected throughout their range, although anthropogenic factors overshadow natural deaths, even in protected populations. Persecution, incidental killing in snares, roadkills and exotic disease transmitted by domestic dogs are the main causes of death. The species now persists predominantly in areas with large parks. The main strongholds are the Okavango–Kaudom–Hwange ecosystem, Kafue NP and Luangwa Valley, Zambia; Selous GR–Mikumi NP and Ruaha–Rungwa ecosystems, Tanzania; and Kruger NP, South Africa). Red List CR in W Africa and N Africa (Algeria only, if not already extinct there), EN elsewhere, population trend Decreasing.
Plate 49
| BLACK-BACKED JACKAL Canis (Lupulella) mesomelas |
SILVER-BACKED JACKAL
HB ♀ 66–85cm, ♂ 69–90cm; T 27–38cm; SH 38–48cm; W ♀ 5.9–10kg, ♂ 6.4–11.1kg
Recent molecular analyses show that the 2 African jackal species are not closely related to Eurasian Golden Jackal, as formerly thought. Indeed, they diverged early (estimated ~3.5 million years ago) from the lineage that gave rise to the Canis clade of large, wolf-like species (see here) and may warrant being grouped separately within a distinct genus, Lupulella (adopted by some authorities). African jackals are most closely related to the African Wild Dog, and are not closely related to other Canis species. Black-backed Jackal is recognisable by its dark-edged silver-grey saddle on a buff to rufous-brown body, and grizzled, dark bushy tail. Pups show adult coloration, including saddle and dark tail, from a very young age (compare with Side-striped Jackal pups). Melanism is not recorded. Usually the most conspicuous jackal where it occurs, due to preference for open areas and aggressive dominance over other similar-sized canids. Distribution and Habitat Two disjunct populations, in southern and E Africa. Occurs in true desert, grassland, montane meadows, arid to mesic woodland savannah, and agricultural habitats. Recorded from sea-level to 3,660m (Mt Kenya, Kenya). Feeding Ecology Generalist omnivore. Recorded killing adult Springbok, Thomson’s Gazelle and Impala, but typical prey includes rodents, Springhare, hares, and young or small ungulates, especially ‘hider’ species such as Bushbuck and Grey Duiker. Juvenile Cape Fur Seals are readily killed by coastal-living jackals near seal colonies. Birds, reptiles, eggs, invertebrates and carrion are also important food items. Readily takes poultry and small livestock. Foraging is mainly nocturnal, especially where it is persecuted, but often crepuscular/diurnal where it is protected. Usually forages alone or in pairs, but up to 12 adults cooperate to kill large prey, and large congregations gather relatively amicably at carcasses and in food-rich patches like seal colonies. Readily scavenges, including from large carnivore kills and human refuse; caches food for later. Social and Spatial Behaviour Monogamous and territorial, forming breeding pairs that may endure for life and defend territories from other pairs. Up to 3 helpers accompany a pair and help to raise pups, including regurgitating food for them and the mother. Coastal Namibian jackals commute up to 20km from their territories to feed at a fur seal colony (Cape Cross Seal Reserve) along shared ‘highways’. Commuters avoid and are tolerated by territorial residents as they travel, and residents mostly abandon territorial defence at the colony itself except during the denning period. Territories range from an average of 1.0km2 (Hwange NP, Zimbabwe) to 24.9km2 (coastal Namibia). Densities peak during the breeding season, e.g. 0.5–0.8/km2 (non-breeding) to 0.7–1/km2 (breeding; Hwange NP). Density estimates 40/100km2 (Giant’s Castle GR, South Africa), 62/100km2 (Kalahari, possibly applying only to riverine areas) and as high as 220/100km2 (Cape Cross Seal Reserve, representing density only in the area of the seal colony). Reproduction and Demography Seasonal. Mating May–August; births June–November. Gestation ~60 days. Maximum litter size 6, typically 3–4. Pups weaned at 8–9 weeks. Sexually mature at 11 months. MORTALITY Predators include Leopards, Lions, Spotted Hyaenas and, occasionally, Cheetahs and African Wild Dogs. Very susceptible to domestic dog diseases (especially rabies) near settlements, although populations recover quickly. LIFESPAN 14 years in captivity. Status and Threats Widespread and common in protected and pastoral areas. Persecuted intensely on farmland (especially in southern Africa), but is very resilient, and anthropogenic population declines appear to be only temporary. Red List LC, population trend Stable.
| SIDE-STRIPED JACKAL Canis (Lupulella) adusta |
HB ♀ 65–76cm, ♂ 66–81cm; T 30–41cm; SH 41–48cm; W ♀ 6.2–10kg, ♂ 5.9–12kg
Grizzled buff-grey, with a characteristic pale stripe (often outlined with a dark border) along the flanks and a distinctive white-tipped dark tail. Pups are typically uniformly buff-grey without obvious body markings, except for obvious white-tipped tail, until >5–6 months old. Melanism is not recorded. Distribution and Habitat Southern, W and C Africa; replaced by Black-backed Jackal in arid southern Africa, and African Wolf in arid N Africa. Inhabits wooded grassland, woodland savannah, marshland, montane areas and forest edges. Uses more open natural habitats, e.g. open grasslands, in the absence of Black-backed Jackals and African Wolves. Avoids very open habitat but can utilise agricultural areas with cover, and occurs in peri-urban and urban areas near major cities. Feeding Ecology The most omnivorous and least predatory jackal species, rarely killing prey larger than gazelle fawns. Diet varies widely locally and seasonally; mainly includes a wide range of fruits, seeds and crops, as well as small rodents, hares, Springhare, small birds, insects and carrion. Free-range poultry is sometimes killed. More strictly nocturnal than other jackals, perhaps to reduce competition with Black-backed Jackal, but becomes crepuscular when unmolested. Forages alone, but family groups gather at rich patches such as termite nests, and up to 12 from various families congregate to scavenge from carrion or dumps. Social and Spatial Behaviour Monogamous, forming mated pairs that may be lifelong. Resident pairs maintain exclusive use of a core territory, with edges shared by neighbouring pairs. Yearling offspring often remain in the territory, sometimes forming groups of up to 7 with the resident pair, although it is unclear if they act as helpers. Range size 0.15–0.56km2 outside breeding, expanding to 0.55–1.6km2 during breeding. Density estimates 7/100km2 (depleted savanna, Niokolo-Koba NP, Senegal) and 54–79/100km2 (W Zimbabwe) in non-breeding season, to 97/100km2 (W Zimbabwe) in breeding season. Reproduction and Demography Seasonal. Mating June–July; births August–November. Gestation 57–60 days. Litter size 4–6. Pups weaned at 8–10 weeks. Sexually mature at 11 months. MORTALITY Preyed on by larger carnivores, including domestic dogs near human habitation. Vulnerable to diseases such as rabies and canine distemper. LIFESPAN 10 years in captivity. Status and Threats Widespread and common. Tolerant of habitat conversion, and persists in suburban and agricultural habitats. Persecuted in human-dominated areas as a livestock predator, and many are killed by snares and vehicles, but human-caused deaths probably produce only local declines unless they are associated with poisoning or disease outbreaks. Thought to be expanding in some areas, e.g. NE South Africa, where Black-backed Jackals are heavily persecuted. Red List LC, population trend Stable.
Plate 50
| ARCTIC FOX Vulpes lagopus |
POLAR FOX, WHITE FOX, BLUE FOX
HB ♀ 50–65cm, ♂ 53–75cm; T 25–42.5cm; W ♀ 3.1–3.7kg, ♂ 3.6–6.7kg
The northernmost canid and the only canid species to change colour seasonally. Two distinct colour morphs: white winter morph moults to grey-brown with cream underparts in summer; blue morph is pale bluish brown in winter and dark grey-brown in summer. White morph is more common, but blue dominates on islands and in coastal areas. Distribution and Habitat Circumpolar in the Arctic. Restricted to Arctic and tundra habitats, mostly north of the treeline in Canada, the USA (Alaska), Greenland, Russia, Finland, Norway, Sweden and Iceland. Inhabits most Arctic islands and winter sea ice to within 60km of the North Pole. Feeding Ecology Small rodents, chiefly lemmings and voles, are critical, especially to inland populations, which fluctuate with rodent ‘boom-bust’ cycles. Other food includes fruits, eggs, birds to the size of geese, ground squirrels, Arctic Hare and infrequent kills of Reindeer/Caribou neonates. Coastal foxes also eat molluscs, crabs, fish, seabirds, and the carcasses of seals and whales; sometimes kills Ringed Seal pups. Blamed for killing domestic sheep lambs, although these are probably scavenged. Foraging is largely nocturno-crepuscular (light during the Arctic summer) and solitary, but congregates at large carcasses. Caches surplus food; 1 larder contained more than 500 eggs. Scavenges from bear and Grey Wolf kills, winter-killed Reindeer and human refuse. Social and Spatial Behaviour Usually solitary and monogamous, forming tight-knit breeding pairs that are territorial near the den. Pairs usually separate after raising pups, but remain in the territory year-round, pairing up again each spring to breed. Helpers from previous litters sometimes linger, forming extended family groups that persist if food is plentiful. Coastal home ranges are generally smallest (5–21km2), with high densities (4–8/100km2) due to more reliable food availability; inland ranges 15–60km2 and densities 0.09–3/100km2. Individuals may wander spectacular distances, possibly driven by rodent crashes, e.g. 2,300km over 3 years by an Alaskan male. Reproduction and Demography Seasonal. Mating February–May; births April–July. Gestation 52–54 days. Litter size typically 3–11, exceptionally reaching 19 in rodent booms. Pups weaned at 7–8 weeks, reaching independence at 12–14 weeks. Most disperse in autumn, but they may overwinter on their parents’ range; some (mainly females) become helpers. Both sexes mature at 10 months, but most individuals do not breed until their third year. MORTALITY Survival is tied to rodent abundance; up to 50% of adults and 80% of pups die in poor years, and populations collapse by up to 80%. Predators include Red Fox (especially of pups), large raptors, Wolverine, Grey Wolf and bears. Domestic dogs kill Arctic Foxes near human settlements and may also transmit disease. LIFESPAN Maximum 10–11 years in the wild, typically under 5; 15 in captivity. Status and Threats More than 100,000 Arctic Foxes are trapped annually for their dense fur, but populations appear to tolerate hunting if pressure is relaxed during poor food years. In Iceland, legally killed as pests by sheep farmers and eider-down collectors. Northwards expansion of Red Fox due to climate warming and human subsidies is causing Arctic Fox declines and range contraction in lowland Fenno-Scandinavia, possibly more widely. Protected in Finland, mainland Norway and Sweden, where historic population crashes have not recovered; unprotected elsewhere. Red List LC, population trend Stable.
| RED FOX Vulpes vulpes |
CROSS FOX, SILVER FOX, COMMON FOX
Includes NORTH AMERICAN RED FOX V. fulva
HB ♀ 45–68cm, ♂ 59–90cm; T 28–49cm; W ♀ 3.4–7.5kg, ♂ 4–14kg
The world’s most widespread and abundant wild carnivore. Typically various shades of rich red-brown, but highly variable, including platinum-tipped black (‘silver fox’), an intermediate morph called ‘cross’ and pale silvery blond. All morphs (except albinos) have black-backed ears, and white-tipped tails are typical but not ubiquitous. Sometimes classified as 2 species based on genetic analyses showing divergence ~209,000 years ago: Eurasian Red Fox (V. vulpes) and North American Red Fox (V. fulva). Distribution and Habitat North America, Eurasia and N Africa. Occurs in virtually all habitats to 4,500m, including farmland, suburbs and cities north of the Tropic of Cancer, except the northernmost Arctic and deserts of the SW USA. Introduced to Australia in the mid-1800s, where it has serious impacts on native wildlife. Feeding Ecology Highly opportunistic, eating mainly reptiles, birds and small mammals to the size of hares; also eggs, amphibians, fish, invertebrates, fruits, acorns, fir cones, sedges, fungi and tubers. Kills juvenile small stock and poultry, and consumes crops such as wheat and corn. Hunting is terrestrial (although it is recorded occasionally climbing to 8m) and mostly nocturno-crepuscular, often more diurnal in winter and where it is undisturbed. Usually forages alone, but aggregates in food-rich patches, e.g. shorebird nest colonies and dumps. Caches surplus food for later use, and has an excellent memory for larder sites. Readily scavenges: ungulate carcasses (including of domestic livestock) can be especially important in winter, and scavenges from human refuse, bird feeders and compost heaps. Social and Spatial Behaviour Usually solitary and monogamous. Mated pair is the main social unit, but sociality during breeding is very flexible. With sufficient food, may form groups comprising 1 male and up to 5 vixens (probably related). Group-breeding females may den alone or together; younger females are mostly non-breeding helpers from previous litters. Range size is resource-dependent, from 0.2km2 (Oxford, UK) to 50km2 (Oman). Density estimates 0.1/km2 (Arctic tundra), 1–3/km2 (temperate forest, Canada and W Europe) and exceptionally reaching 30/km2 with abundant food, e.g. urban areas where foxes are subsidised. Reproduction and Demography Seasonal. Mating is in winter, usually December–February, earlier in southerly latitudes (Australia: June–October). Births usually March–May. Oestrus 1–6 days; gestation 49–55 days. Litter size reaches 12 depending on food availability, typically 3–8. Pups weaned at 6–8 weeks, and most disperse from age 6 months before the next breeding season. Some young females remain as helpers. Sexually mature at 9–10 months. MORTALITY First-year mortality reaches 80% and averages 50% for adults, mainly due to humans. Large raptors, other carnivores and domestic dogs kill Red Foxes, although humans are overwhelmingly their main predator. Major vector for rabies, with outbreaks causing intermittent population crashes. LIFESPAN Maximum 9 years in the wild, typically <5; 15 in captivity. Status and Threats Remarkable adaptability and resilience enables it to tolerate intense persecution, with some 1–2 million wild individuals killed annually for the fur trade, and perhaps the same amount again by sport hunters and pest control. Unprotected in most of its range; exports of furs from India are restricted. CITES Appendix III – India; Red List LC, population trend Stable.
Plate 51
| KIT FOX Vulpes macrotis |
HB 45.5–54cm; T 25–34cm; W ♀ 1.6–2.2kg, ♂ 1.7–2.7kg
Smallest fox on mainland North America. Tawny grey with ochre sides, neck and legs. Closely related to Swift Fox; hybrids occasionally occur in a narrow band of overlap in Texas and New Mexico. Distribution and Habitat W USA and N Mexico. Inhabits semi-arid to arid desert scrub and grassland. Can occupy urban and agricultural areas. Feeding Ecology Eats mainly mice, rats, kangaroo rats, ground squirrels, prairie dogs, lagomorphs and insects; also small birds, reptiles, carrion, wild cactus fruits and crops, e.g. tomatoes and almonds. Foraging is mainly nocturnal and solitary. Scavenges from livestock carcasses and human refuse. Social and Spatial Behaviour Monogamous, normally mating for life. Helpers, usually grown daughters, often remain with the pair. Ranges are stable, with exclusive core denning areas. Range size (both sexes) averages 2.5–11.6km2. Densities fluctuate widely depending on prey oscillations: 10–170/100km2, averaging 44/100km2. Reproduction and Demography Seasonal. Mating December–January; births February–March. Gestation 49–55 days (estimated). Litter size 1–7. Weaning at around 3 months and independence at 5–6 months. MORTALITY Average annual mortality 65% (juveniles) and 45% (adults), mainly from Coyote predation and starvation during prey shortages. LIFESPAN 7 years in the wild, 20 in captivity. Status and Threats Secure but has undergone significant local declines, e.g. California and Mexico, from habitat conversion and eradication of prey colonies, e.g. prairie dog and kangaroo rat towns. Threatened in California and Oregon, endangered in Colorado, vulnerable in Mexico. Red List LC, population trend Decreasing.
| SWIFT FOX Vulpes velox |
HB 47.5–54cm; T 25–34cm; W ♀ 1.6–2.3kg, ♂ 2–2.95kg
Very similar to Kit Fox, with more rounded ears and a shorter tail. Distribution and Habitat Mid-western North America, from Alberta to New Mexico. Restricted to short- to medium-grass prairies and grassland. Tolerates dry land agricultural areas. Feeding Ecology Diet dominated by prairie dogs, ground squirrels, mice and lagomorphs, plus wild fruits, seeds, insects, small birds, reptiles, eggs and carrion. Foraging is solitary and mainly nocturno-crepuscular. Social and Spatial Behaviour Mated pairs are typical, but sociality is flexible; trios and extra-pair breeding occur. Pairs/groups maintain exclusive core areas with overlapping edges; neighbours are often related. Range size (both sexes) 7.6km2 (Colorado) to 25.1km2 (W Kansas). Reproduction and Demography Seasonal. Mating December (Oklahoma) to March (Canada); births March–May. Gestation 50–55 days. Litter size 3–6, exceptionally to 8. MORTALITY Rates average 67–95% (juveniles) and 36–57% (adults), mainly from Coyote predation. LIFESPAN 8 years in the wild, 14 in captivity. Status and Threats Extirpated from about 60% of its historic range, from massive prairie conversion and intense persecution of prey. Extirpated from Canada by 1938; now present as a small reintroduced population. Red List LC, population trend Stable.
| INDIAN FOX Vulpes bengalensis |
BENGAL FOX
HB ♀ 46–48cm, ♂ 39–57.5cm; T 24.5–32cm; W ♀ 2–2.9kg, ♂ 2.3–3.6kg
Slender, fine-featured fox with a narrow face, yellow-grey to silvery-grey fur and a black-tipped tail. Distribution and Habitat Endemic to the Indian subcontinent. Inhabits hot, semi-arid grassland, plains, scrub and open, dry forest. Occurs on agricultural land. Feeding Ecology Omnivorous, eating mainly small mammals and insects, supplemented by birds, reptiles, eggs, fruits, seeds and fresh shoots. Foraging is solitary and nocturno-crepuscular; often diurnal on cool or overcast days. Scavenges from carrion, but rarely since feral dogs dominate carcasses. Social and Spatial Behaviour Monogamous in mated pairs. Other adults sometimes associate with pairs, but do not help raise pups; their role and relatedness is uncertain. Range estimates average 1.6km2 (♀s) and 3.1km2 (♂s). Densities fluctuate due to rodent cycles and disease outbreaks, usually 1–15/100km2, reaching 150/100km2 in ideal conditions. Reproduction and Demography Seasonal. Mating November–January; births January–May. Gestation 50–53 days. Litter size 2–4. Both parents provision and guard pups. MORTALITY Disease is implicated in population crashes. LIFESPAN 8 years in captivity. Status and Threats Relatively widespread, but occurs at low densities in habitats that are under strong development pressure. Naturally vulnerable to population crashes, exacerbated by hunting pressure from people for food. CITES Appendix III – India; Red List LC, population trend Decreasing.
| CORSAC FOX Vulpes corsac |
CORSAC, STEPPE FOX
HB ♀ 45–50cm, ♂ 45–59.5cm; T 19–34cm; W ♀ 1.6–2.4kg, ♂ 1.7–3.2kg
Medium-sized fox with pale, tawny-grey fur (silky and frosted in winter) and a black-tipped tail. Distribution and Habitat C Asia, from W Russia to NE China. Inhabits steppes, grassland, shrubland, semi-desert and desert. Feeding Ecology Omnivorous and opportunistic, exploiting seasonal fluctuations in food, especially of rodents, e.g. lemmings, voles, ground squirrels, gerbils and jerboas. Also consumes birds, reptiles, insects and carrion. Foraging is solitary and usually nocturnal; diurnalism increases when feeding young pups and during food shortages. Scavenges, including from human refuse, Grey Wolf kills and winter-killed livestock. Social and Spatial Behaviour Forms monogamous breeding pairs, although it is unknown if they are permanent or territorial, and 2 females sometimes den together with their pups. Juveniles may become helpers. Range size (pairs) 3.5–11.4km2 (C Mongolia), occasionally to 35–40km2 in poor habitat. Reproduction and Demography Seasonal. Mating January–March; births mid-March–May. Litter size 2–10, averaging 5–6. Gestation 52–60 days. Both parents provision and guard pups; helpers sometimes assist. MORTALITY Average adult mortality 34% (protected reserve, C Mongolia), mainly from human hunting and Red Fox predation. LIFESPAN 9 years in captivity. Status and Threats Widespread and locally common, but heavy hunting pressure for furs and rodent-poisoning campaigns (especially in China) are pervasive. Extirpated in many areas, especially in Russia and its former republics. Red List LC, population trend Decreasing.
Plate 52
| BLANFORD’S FOX Vulpes cana |
KING FOX, ROYAL FOX, AFGHAN FOX
HB ♀ 34–45cm, ♂ 38.3–47cm; T 26–36cm; W 0.8–1.6kg
Very small fox with distinctive dark ‘tear’ lines along the muzzle. Brown-grey fur is interspersed with long black guard hairs, and very bushy tail usually has a black (rarely white) tip. Distribution and Habitat E Egypt, Arabian Peninsula and C Asia. Inhabits semiarid to arid rocky desert and mountainous habitats. Independent of water. Feeding Ecology Largely insectivorous and frugivorous, eating mainly beetles, crickets, grasshoppers, ants, termites, scorpions, wild capers, olives, grapes and melons. Small rodents, birds and (rarely) reptiles are also hunted; newborn ibex records are probably carrion. Foraging is usually solitary and nocturnal, with increased crepuscularity in winter. Social and Spatial Behaviour Forms territorial monogamous pairs that cooperate to raise pups, but are fairly solitary outside the breeding period. Non-breeding yearling females (possibly from previous litters) are often tolerated by resident pairs. Range size (both sexes) 0.5–2km2, averaging 1.6km2. Density estimates 0.5–2/km2 (Israel) to 8/km2 (high-quality habitat with abundant carrion, Jordan). Reproduction and Demography Seasonal. Mating January–February (Israel); births late February–May. Gestation 50–60 days. Litter size 1–3. Males groom, guard and accompany pups (2–4 months) on foraging excursions. MORTALITY Rabies and old age appear to be the main mortality factors. Red Fox is a confirmed predator. LIFESPAN <5 years in the wild, 6 in captivity. Status and Threats Fairly widespread and common. Locally threatened by habitat development, especially in coastal areas, and incidentally killed by poison set for other species. CITES Appendix II; Red List LC, population trend Stable.
| PALE FOX Vulpes pallida |
PALLID FOX, AFRICAN SAND FOX
HB 38–55cm; T 23–28.5cm; W 1.2–3.6kg
Very small fox, uniformly pale sandy cream except for dark-tipped tail (distinguishing it from Rüppell’s Fox). Ears medium-sized, in proportion to the head and lacking the oversized appearance in Rüppell’s Fox and Fennec. Distribution and Habitat Sub-Saharan Africa, in a narrow band from W Senegal–Mauritania to Eritrea and Ethiopia. Inhabits very arid, sandy and stony deserts, and dry savannah. Occurs near human settlements. Feeding Ecology Primarily insectivorous, eating mainly weevils, scarabs, grasshoppers and scorpions; small mammals, especially gerbils and jerboas, are also taken. Robust molars suggest fruits, seeds and plant matter are consumed, but these appeared very rarely in scats collected in SE Niger. Largely nocturnal. Social and Spatial Behaviour Mostly seen in pairs and small groups, probably mated pairs with offspring, suggesting patterns similar to those of other small foxes. Range estimates average 5.6km2 with minimal overlap, suggesting territoriality (Termit and Tin Toumma National Nature and Cultural Reserve, Niger) Reproduction and Demography Probably seasonal. Gestation (captivity) 51–53 days. Litter size 3–6. MORTALITY Poorly known; occasionally killed by domestic dogs. LIFESPAN Unknown. Status and Threats One of the least known canids; status unknown. Occasionally killed near settlements, and used locally for traditional medicines, e.g. Sudan, for asthma. Red List LC, population trend Unknown.
| RÜPPELL’S FOX Vulpes rueppellii |
SAND FOX
HB 35–56cm; T 25–39cm; W ♀ 1.1–1.8kg, ♂ 1.1–2.3kg
Small, fine-featured fox with large ears and a slender face. Body colour near white to greyish brown, often with a silvery sheen due to dark guard hairs. Long, bushy tail has a white tip. Distribution and Habitat N Africa (Sahara), the Arabian Peninsula and C Asia. Independent of water and inhabits semi-arid to very arid habitats, including sandy and stony deserts, rocky steppes, massifs, scrub and vegetated watercourses. Occurs near human habitation. Feeding Ecology Omnivorous. Eats small mammals, birds, lizards, insects and plant matter, including wild fruits (especially dates), desert succulents and grass (probably an emetic). Foraging is solitary and nocturno-crepuscular. Most hunting is terrestrial; climbs palm trees for dates. Scavenges from carrion and human refuse. Social and Spatial Behaviour Monogamous, forming mated territorial pairs, but larger aggregations of up to 15 suggest more complex sociality. Territories are largely exclusive, although large ranges overlap significantly. Female range sizes 13.2km2 (Saudi Arabia) to 53.8km2 (Oman); male ranges 20.9–84.4km2. Density estimates 0.7–1.05/km2 (fenced reserve, Saudi Arabia). Reproduction and Demography Seasonal. Mating November–February (Saudi Arabia); births March–May. Gestation 52–56 days. Litter size 2–6. Pups independent at 4 months and disperse at 6–10 months. MORTALITY Rabies, distemper and predation are the main factors; Steppe Eagle and Pharaoh Eagle-owl are confirmed predators. LIFESPAN 7 years in the wild, 12 in captivity. Status and Threats Widespread and quite common. Killed indiscriminately during poisoning campaigns, and absent in heavily grazed areas. Expanding Red Fox range (in association with human settlements) displaces Rüppell’s Fox; it is almost extinct in Israel as a result. Red List LC, population trend Stable.
| TIBETAN FOX Vulpes ferrilata |
TIBETAN SAND FOX, SAND FOX
HB 49–70cm; T 22–29cm; W ♀ 3–4.1kg, ♂ 3.2–5.7kg
Very distinctive stocky fox with a long, narrow muzzle, small ears and small, wide-set eyes on a broad face. Body grizzled grey with white underparts, and a rufous cape, head and lower legs. Distribution and Habitat Restricted to the Tibetan Plateau in Ladakh (India), N Nepal, and through S and C China; presence in N Bhutan is unconfirmed. Inhabits remote, cold, semi-arid to arid steppes, meadows, grassland and slopes at 2,500–5,200m. Tolerates ambient temperatures of –40°C to 30°C. Feeding Ecology Eats mainly small mammals, especially pikas (with which it is closely associated), mice and zokors. Also eats hares, marmots, birds, lizards, insects and berries. Eats carrion, including scavenging Grey Wolf kills, and follows foraging Brown Bears to mop up flushed rodents. Hunts alone and appears diurnal, possibly reflecting the activity patterns of pikas. Social and Spatial Behaviour Poorly known. Monogamous breeding pairs are typical, but adult trios with pups are recorded. Pairs are found in close proximity, especially in food patches like pika colonies, suggesting the species is not strongly territorial. Densities are naturally low, reaching 2–4/km2 with abundant prey and low hunting pressure. Reproduction and Demography Seasonal. Mating December–March; births February–May. Litter size 2–5. Gestation 50–55 days. MORTALITY and LIFESPAN Unknown. Status and Threats Widespread and inhabits remote areas, somewhat insulating it from human threats. Given its strong association with pika colonies, the gravest threat is the state-sanctioned rodent poisoning affecting most of the Tibetan Plateau. Red List LC, population trend Unknown.
Plate 53
| FENNEC Vulpes zerda |
FENNEC FOX
HB 33.5–39.5cm; T 12.5–23cm; W 0.8–1.9kg
Smallest canid, with proportionally the largest ears in the family. Pale cream to sandy red, with lighter underparts, a dark-tipped tail and a dark caudal spot. Soles of the feet are fully furred for traversing hot, loose sand. Distribution and Habitat Restricted to N Africa; reports from the Arabian Peninsula are unverified. Extremely well adapted to deserts; water-independent. Prefers stable sand dunes for burrows, but occupies all semi-arid/arid habitats. Feeding Ecology Eats very small prey, chiefly small rodents, lizards, geckos, birds to the size of sandgrouse, invertebrates and their larvae, eggs, fruits (especially dates) and tubers. Reportedly raids poultry coops, although evidence is anecdotal. Forages alone, largely at night, becoming more crepuscular/diurnal in winter. Prodigious digger, catching most prey by speedy excavation after locating it with its extremely sensitive hearing. Caches surplus food in little excavations, and sometimes enters human settlements to scavenge. Social and Spatial Behaviour Poorly known. Thought to form territorial breeding pairs in enduring ranges; observed groups with up to 10 individuals suggest extended families with helpers. Density is unknown but they are often the most common mammal (aside from rodents) recorded in surveys. Reproduction and Demography Thought to be seasonal (but breeds year-round in captivity). Mating January–February; births March–April. Gestation 50–52 days. Litter size 1–4. Pups weaned at 61–70 days. MORTALITY Putative predators include large owls, African Wolf and domestic dogs. LIFESPAN 13–14 years in captivity. Status and Threats Status uncertain, but probably secure by virtue of inhabiting very remote regions. Main threat is trapping for the tourist pet and domestic fur trades, which drives local extinctions around settlements. CITES Appendix II; Red List LC, population trend Stable.
| CAPE FOX Vulpes chama |
HB 45–61cm; T 25–40.6cm; W 2–3.3kg
The only small, light-coloured fox in southern Africa. Body fur grizzled silver-grey, blending into pale tawny-reddish limbs, neck and head. Long, bushy tail has a characteristic dark tip. Distribution and Habitat SW Angola, Namibia, Botswana and South Africa; possibly Swaziland and Lesotho. Favours semi-arid/arid habitats extending into moderately mesic scrub habitat in South Africa and Botswana. Occurs on farmland. Feeding Ecology Eats chiefly mice and gerbils; also invertebrates, small birds, reptiles and fruits. Largest kills are hares and Springhare. Very rarely kills newborn sheep and goats; most livestock consumed is scavenged. Forages alone, usually at night, with activity peaks after dusk and before dawn. Most prey is captured after prolonged listening at burrows or holes, ending with frenzied digging. Surplus food is cached. Eats carrion, although rarely visits large carnivore kills. Social and Spatial Behaviour Usually solitary and monogamous. Breeding pairs are typical, with both parents raising pups. Helpers occur rarely, and adult females (possibly related) occasionally den together. Solitary outside breeding, but pairs share the same range year-round. Range size of pairs 1–32.1km2. Density estimates 0.05–0.3/km2. Reproduction and Demography Broadly seasonal, weakly so in some regions. Seasonal populations mate from June; most births August–December. Gestation 51–52 days. Litter size 1–6, typically 2–4. Pups begin hunting at around 15–16 weeks, reaching independence at 5 months. MORTALITY Adult mortality 26–52% (South Africa), depending on densities of Black-backed Jackal, the most important predator. Other predators include large raptors, owls and large carnivores, including domestic dogs. Contracts rabies, but is apparently less susceptible than other canids. LIFESPAN <7 years in the wild. Status and Threats Many thousands of Cape Foxes are killed, mainly in South Africa and Namibia, for perceived livestock losses, deliberately and as ‘by-catch’ in trapping and poisoning campaigns targeting jackals and Caracal. Despite this, the species often prospers on farmland that lacks larger predators (especially jackals) and has expanded its range in some areas. Red List LC, population trend Stable.
| BAT-EARED FOX Otocyon megalotis |
HB 46.2–60.7cm; T 23–34cm; W 3.4–5.4kg
Grizzled, smoky grey, with black legs and a bushy black-edged tail. Unique dentition, with a range of 46–50 total teeth, the most for any placental land mammal. Distribution and Habitat Two disjunct populations in southern and E Africa. Favours arid or semiarid grassland and open woodland savannah. Inhabits farmland and degraded habitat provided insecticide use is limited. Feeding Ecology Almost entirely insectivorous, eating particularly 2 termite genera with which its distribution overlaps almost completely. Also eats other invertebrates, small reptiles, rodents and some fruits, especially in the dry season. Does not kill livestock or poultry, but eats insect larvae in livestock carcasses, sometimes incurring blame for depredations. Forages mainly by sound, typically in family groups. Feeding aggregations of up to 15 individuals from different families occur when termites are abundant. Foraging is chiefly nocturnal, but shifts diurnally in winter, reflecting changes in termite activity. Does not cache food and rarely scavenges. Social and Spatial Behaviour Monogamous pair is typical, but sometimes forms extended groups with 1 male and 2–3 related females that breed communally and allo-suckle pups. Insectivorous diet precludes female provisioning at the den, so the male is essential for guarding pups and allowing mothers to forage. Ranges overlap and territorial defence is limited to the den area. Range size 1–8km2. Densities fluctuate depending on food and season, e.g. 2.3/km2 (non-breeding) to 9.2/km2 (breeding; Mashatu GR, Botswana), and 0.7–14/km2 (Kalahari, South Africa). Reproduction and Demography Seasonal. Pairs mate for life. Mating July–September; births October–December. Gestation 60–75 days. Litter size 1–6, averaging 3. Pups weaned at 10–15 weeks, but remain with the parents until the following June–July; many disperse, although some stay as helpers. MORTALITY Adult mortality 25–30% (South Africa), mainly from predation by Black-backed Jackal and disease episodes (rabies and canine distemper). Other predators include large raptors, owls and larger carnivores, but groups vigorously mob predators, often deterring predation. LIFESPAN 9 years in the wild, 13 in captivity. Status and Threats Reasonably widespread and tolerant of habitat conversion, but mistakenly persecuted as a livestock predator and cannot persist where insecticide kills off prey. Disease outbreaks associated with domestic dogs sometimes trigger severe local die-offs. Red List LC, population trend Stable.
Plate 54
| ISLAND FOX Urocyon littoralis |
ISLAND GREY FOX, CHANNEL ISLANDS FOX
HB 45.6–63.4cm; T 11.5–32.2cm; W 1.07–2.7kg
Smallest North American canid, very similar to Grey Fox but much smaller. An insular dwarf form of Grey Fox, separated for 10,400–16,000 years, and genetically distinct. Distribution and Habitat Restricted to 6 Channel Islands off the California coast, USA. Occupies all island habitats, including grassland, chaparral scrub, woodland, coastal scrub and dunes. Avoids degraded areas such as overgrazed pasture. Feeding Ecology Omnivorous, focusing on the most abundant food source, which differs by habitat and island. The most important food types are small mammals (especially deer mice), insects and fruits (especially prickly pears, and also berries). Also eats seeds, acorns, birds, nestlings, eggs and crustaceans. Cathemeral, with nocturnal activity peaks. Forages on the ground, climbs shrubs for fruits and birds’ nests, and scales cliffs for seabird eggs and chicks. Social and Spatial Behaviour Monogamous and territorial, living as mated pairs with offspring, which usually remain until their second year; adult offspring are often tolerated even after they are living independently. Resident males chase and fight with other males, mainly during the breeding season. Ranges are among the smallest for any canid: 0.15–0.87km2. Reproduction and Demography Seasonal. Mating January–March; births February–May (peaking April). Gestation 50–53 days. Maximum litter size 5, typically 1–3. Pups weaned at 6–8 weeks and forage with adults from 2 months. MORTALITY Recent Golden Eagle colonisation on northern Channel Islands led to hyperpredation, prompting population crashes. Exotic disease (especially canine distemper) reduced the Santa Catalina Island population by 95% in 1998–2000. LIFESPAN 10 years in the wild, 15 in captivity. Status and Threats Restricted to small, isolated populations that are greatly vulnerable to random events. Golden Eagle predation and exotic disease (and less so roadkill) produced calamitous declines on 4 of the 6 islands, reducing the total population from 6,000, to 1,500 in 2002. Aggressive conservation interventions (especially captive breeding and reintroduction, vaccination against canine diseases and relocation of Golden Eagles) have since reversed declines; by 2011, total population had increased to 5,500 (including >4,000 mature adults) and annual survival rates to >85%. Red List NT, population trend Increasing.
| GREY FOX Urocyon cinereoargenteus |
NORTHERN GREY FOX, TREE FOX
HB ♀ 52.5–58cm, ♂ 56–66cm; T 28–44.3; W ♀ 2–3.9kg, ♂ 3.4–7kg
Grizzled grey body colour, bordered by a sharp edge of rufous on the neck, sides and legs, changing to white underparts, chest and cheeks. Long, bushy tail has a black dorsal stripe and tip. Distribution and Habitat Extreme SE Canada, most of the USA, Meso-America, extreme N Colombia and N Venezuela. Inhabits temperate and tropical forests, woodland, brush, semi-arid scrubland, agricultural habitats and peri-urban areas. Avoids very open areas such as grassland and prairie. Feeding Ecology The most omnivorous North American fox. Hunts mainly rodents and rabbits in winter, but greatly expands its diet in other seasons as different foods become available. Can be almost exclusively insectivorous in summer, while fruits, seeds and nuts comprise up to 70% of its diet in autumn. Also eats small herptiles, birds, eggs and carrion. Foraging is mainly nocturnal and solitary. The most arboreal of canids, and forages both terrestrially and in trees up to 18m. Social and Spatial Behaviour Monogamous and territorial, living as mated pairs with offspring. It is unclear if pair bonds are permanent or if offspring remain as helpers. Territory size differs little between sexes, ranging from 0.13km2 (Wisconsin) to 27.6km2 (Alabama), averaging 1–6.7km2. Density estimates 0.4/km2 (California) to 1.5/km2 (Florida). Reproduction and Demography Seasonal. Mating January–April; births March–early June. Gestation 60–63 days. Maximum litter size 10, typically 3–5. Pups weaned at 6–8 weeks and accompany foraging adults from 3 months. MORTALITY Most important predator is Coyote, which suppresses populations in some areas. Distemper and rabies produce local crashes. LIFESPAN 4–5 years in the wild, 14 in captivity. Status and Threats Widespread and common. Legal trapping is the main source of mortality; trapping is not regarded as a threat, although it probably contributes to localised population impacts in combination with disease epidemics. Red List LC, population trend Stable.
| RACCOON DOG Nyctereutes procyonoides |
TANUKI
HB 49–70.5cm; T 15–23cm; W 2.9–12.5kg
Grizzled dark grey to buff-grey, with a black facial mask, chest, legs and feet. White (leucistic) individuals occur. An ancient canid lineage with no close relatives, and classified in its own genus. Some authors consider the Japanese population a distinct species, N. viverrinus, based on limited genetic and morphological evidence. Distribution and Habitat Japan, SE Russia, W Mongolia, E China, Korean Peninsula and extreme N Vietnam. Introductions and escapes from fur farms have established populations throughout N, E and W Europe. Inhabits a variety of forest types, shrubland, farmland and urban areas. Feeding Ecology Omnivorous. Rodents are the mainstay, supplemented by small herptiles, birds, eggs, fish, crustaceans and carrion. Fruits, including berries, and seeds are important in late summer–autumn before hibernation. New leaves and flowers are consumed mainly in spring, and insects peak in the diet in summer. Eats crops, including oats, corn, maize, watermelon and fruits, and urban populations exploit garden fruits such as gingko and persimmon, e.g. Japan; rarely raids poultry. Foraging is mostly terrestrial and nocturnal. Adult pairs forage together, but often some distance apart. It is the only canid that hibernates, in November–March in areas with severe winters. Social and Spatial Behaviour Strictly monogamous, forming permanently mated pairs that share a territory, usually moving and denning together. Territory cores are exclusive, especially while breeding, but range edges overlap. Territory sizes 0.07km2 (urban, Japan), 6.1km2 (subalpine habitat, Japan) and to 20km2 (SE Russia, introduced). Reproduction and Demography Seasonal. Mating February–March; births April–June. Gestation 59–70 days. Litter size averages 4–9, exceptionally reaching 19. Pups weaned at 5 weeks, and forage with adults shortly thereafter. MORTALITY Sarcoptic mange, distemper and rabies cause population declines, but impacts appear temporary. LIFESPAN 5 years in the wild, 13 in captivity. Status and Threats Abundant in much of its original range, and widely considered a pest where it has been introduced. Very tolerant of suburban and agricultural habitats. Up to an estimated 370,000 are killed annually on roads in Japan, apparently without producing population declines. Fur farms in China maintain a staggering 10 million Raccoon Dogs (2009), presumably mostly captive-bred. Red List LC, population trend Stable.
Plate 55
| CHILLA Lycalopex griseus |
ARGENTINE GREY FOX, SOUTHERN GREY FOX
HB 50.1–66cm; T 11.5–34.7cm; W 2.5–5kg
Small, pale, grizzled grey fox, with rufous-buff lower legs and a rufescent head tinged with grey. Lower thighs have a distinctive dark patch, and the lower jaw is conspicuously dark. Distribution and Habitat S Peru to Tierra del Fuego on both sides of the Andes; Peruvian population may be disjunct, as the species is thought to be absent from the Atacama Desert. Inhabits grassland, shrubland, steppes and Nothofagus thickets in lowlands and Andean foothills (rarely to 4,000m). Tolerates ranching, agriculture and plantations. Feeding Ecology Eats mainly small rodents, European Hare and carrion. Where small mammal availability declines, the diet includes more fruits, arthropods, reptiles and birds. Kills domestic poultry and (very rarely) lambs. Foraging is largely nocturno-crepuscular, and solitary or as loosely associated pairs with offspring during breeding. Buries excess food. Social and Spatial Behaviour Forms monogamous breeding pairs, occasionally with female helpers, which help raise pups. Pairs associate loosely outside breeding season. Range size 1.4–2.8km2. Reproduction and Demography Seasonal. Mating August–September; births October. Gestation 53–58 days. Litter size 4–6. Males help raise pups and provision mother at den. Two mothers (possibly related) may cooperate to raise litters. MORTALITY Killed by domestic dogs and canine diseases, but impacts are unknown. LIFESPAN Unknown. Status and Threats Widespread and locally common, especially in its southern range. Once heavily hunted for fur, which has declined, leading to population recovery in some areas. Killed for depredation and in the false belief that it transmits disease to livestock. CITES Appendix II; Red List LC, population trend Stable.
| DARWIN’S FOX Lycalopex fulvipes |
HB 48–59.1cm; T 17.5–25.5cm; W ♀ 1.8–3.7kg, ♂ 1.9–4kg
Small, stocky fox, dark grey-brown with contrasting white underparts, rufous extremities and a dark bushy tail. Distribution and Habitat Endemic to Chile, on Chiloé Island and on the mainland coastal range. Mainland population formerly thought to be restricted to Nahuelbuta NP, but recently confirmed from a much larger area. Relies on dense southern temperate (Valdivian) forest, but tolerates forest mosaics with dunes, beaches and pasture patches. Feeding Ecology Eats mainly small vertebrates, insects, crustaceans, fruits and seeds. Largest recorded prey, Southern Pudú (10kg) and Magellanic Penguin, is probably scavenged. Foraging is solitary, but congregates at food patches. Cathemeral; foraging mainly nocturnal in Nahuelbuta, perhaps to avoid larger Chilla (absent from Chiloé). Scavenges, including from fishing waste, pet food and carrion. Social and Spatial Behaviour Solitary. Forms monogamous pairs, which associate mainly for breeding on Chiloé but persist year-round in Nahuelbuta. Range size (both sexes) averages 1.5–3km2. Density estimates 0.95–1.14/km2. Reproduction and Demography Poorly known. Seasonal. Observed litters occur October–January. Gestation unknown. Litter size estimated at 2–3. Both parents help raise pups. MORTALITY Rates in Nahuelbuta are 7–16% annually (juveniles and adults combined). Puma is a confirmed predator (mainland only); no natural predators on Chiloé. Killed by domestic dogs at both sites. LIFESPAN Unknown. Status and Threats Formerly thought to number <250 adults in 2 populations, but recent range extensions significantly increase the estimated population to ~640–900. None the less, Darwin’s Foxes are still strongly threatened by habitat loss to logging, and widespread occurrence of feral domestic dogs and associated disease. CITES Appendix II; Red List EN, population trend Decreasing.
| SECHURAN FOX Lycalopex sechurae |
SECHURA DESERT FOX, SECHURA FOX
HB 50–78cm; T 27–34cm; W 2.6–4.2kg
Slender head with a narrow, long muzzle, large rufous-backed ears and a rufous ring around the eyes. Distribution and Habitat Coastal NW Peru and extreme SW Ecuador. Inhabits desert, associated beaches and sea cliffs, dry forest and Andean foothills to 1,000m. Occurs in agricultural areas. Feeding Ecology Omnivorous and opportunistic. Fruits and seeds comprise much of the diet; also small rodents, birds, reptiles, insects, scorpions and carrion. Coastal individuals eat crabs, and seabirds (probably scavenged) and their eggs. Occasionally takes poultry and domestic guinea pigs; blamed for killing goats (unlikely). Foraging is primarily nocturnal and solitary; congregates in small groups at large carcasses. Social and Spatial Behaviour Poorly known. Sightings are largely of single adults or females with pups. Reproduction and Demography Poorly known. Births probably peak October–January. MORTALITY Roadkills common in N Peru; impacts unknown. LIFESPAN Unknown. Status and Threats Very limited distribution; status poorly known. Appears to tolerate rural and agricultural areas. Vulnerable to persecution, and religious ceremonial uses as amulets and handicrafts. Red List NT, population trend Unknown.
| CULPEO Lycalopex culpaeus |
ANDEAN FOX
HB 44.5–92.5cm; T 31–49.5cm; W ♀ 3.9–10kg, ♂ 3.4–13.8kg
Largest South American fox, powerfully built and with a robust head. Light to dark grey, with tawny extremities and pale underparts. Distribution and Habitat Extreme S Colombia to southern tip of South America. Inhabits all Andean habitats, from dry desert to temperate rainforest, and from coasts to 4,800m (higher than other South American canids). Occurs on ranchland. Feeding Ecology More carnivorous and predatory than other South American foxes. Eats mainly rodents, introduced and native lagomorphs, and ungulates (wild and domestic), mostly as carrion but recorded killing young Alpacas, Vicuñas, Llamas and Guanacos (up to 33kg); Northern Pudú (5–10kg) and Little Red Brocket (8–14kg) dominate the diet in Podocarpus NP, Ecuador. Also consumes fruits, birds, invertebrates and herptiles. Kills small livestock, generally young lambs but occasionally adults, e.g. a 3.6kg juvenile fox killed a 24kg goat by suffocation. Generally forages alone and is cathemeral; strictly nocturnal where hunted. Scavenges. Social and Spatial Behaviour Solitary and territorial. Forms pairs for breeding; male assists in caring for pups. Range estimates (both sexes) typically 4.5–8.9km2, but as large as 800km2. Reproduction and Demography Seasonal. Mating August–October; births October–late December. Gestation 55–60 days. Litter size 3–8, averaging 5. MORTALITY Rates for juveniles 20% (not hunted) to 92% (hunted), and for adults 31% (not hunted) to 49% (hunted). Chief predators are domestic dogs and Puma. LIFESPAN 11 years in the wild. Status and Threats Widespread, common and resilient. Benefits from pastoral/agricultural conversion with introduction of exotic lagomorphs. Intensely persecuted as a livestock pest and extirpated from extensive sheep-ranching areas. CITES Appendix II; Red List LC, population trend Stable.
Plate 56
| HOARY FOX Lycalopex vetulus |
HOARY ZORRO, SMALL-TOOTHED DOG
HB 49–71cm; T 25–38cm; W 2.5–4kg
Small, slender fox, grizzled grey with buff lower legs and underparts, and with a less crisply contrasting chest and throat patches than in similar foxes. Smallest canid in its range. Distribution and Habitat Endemic to Brazil. Inhabits mainly open cerrado savannah, and occasionally associated woodland, forest and floodplains. Occurs in pastoral, agricultural and plantation habitats. Feeding Ecology Largely insectivorous, with a diet dominated by harvester termites, dung beetles and grasshoppers. Also eats small rodents, birds, reptiles, wild fruits and grasses. Evidence for poultry depredation is equivocal. Foraging is largely nocturnal, singly or as pairs with offspring. Most prey is taken on the soil surface; flips cattle dung for termites and dung beetles. Social and Spatial Behaviour Forms monogamous breeding pairs that inhabit a defined range and cooperate to raise pups. Unclear whether pairs are permanent, and there is no evidence of extended family groups with helpers. Range size 3.8–4.6km2. Reproduction and Demography Seasonal. Mating late May–June; births July–August. Gestation 50 days (estimated). Litter size 2–5. Weaning at around 3 months. Males help groom, guard and chaperone pups on foraging excursions. MORTALITY Maned Wolves are putative predators. In C Brazil in 2007–13, 30 of 70 foxes monitored in non-protected habitat were killed by people (including road mortalities) and their dogs. LIFESPAN 8 years in captivity. Status and Threats Widespread, common and tolerant of some habitat conversion to pasture and agriculture. Persecuted (probably mistakenly) for poultry depredation, and roadkills are frequent. Rabies and sarcoptic mange are confirmed, although population impacts are unknown. Red List LC, population trend Unknown.
| PAMPAS FOX Lycalopex gymnocercus |
AZARA’S FOX
HB ♀ 50.5–72cm, ♂ 60–74cm; T 25–41cm; W ♀ 3–5.7kg, ♂ 4–8kg
Medium-sized fox, grizzled grey with reddish ears, neck and lower limbs. Throat, chest and lower hind limbs creamy white. Smaller than sympatric Culpeo. Distribution and Habitat Extreme SE Brazil, E Bolivia, NE Argentina, Paraguay and Uruguay. Optimal habitat is pampas grassland, but also inhabits scrub, open woodland, pasture and agricultural land. Feeding Ecology Adaptable omnivore with a diet that shifts depending on availability. Most important food items are small rodents, European Hare, grassland birds, insects and fruits. Eats carrion, especially livestock carcasses. Rarely kills newborn lambs. Foraging is solitary, but congregates at large carcasses. Cathemeral, becoming largely nocturnal where persecuted. Scavenges. Social and Spatial Behaviour Solitary. Forms monogamous pairs, but appears to associate only during the breeding season. Range estimates 0.4–1.8km2. Reported densities typically 1–3/km2, peaking at 5.85/km2 in optimal pampas habitat free of persecution. Reproduction and Demography Seasonal. Mating July–August; births September–December. Gestation 55–60 days. Maximum litter size 8, typically 3–4. Pups weaned at 2 months. Both parents guard pups, and males provision females and pups at the den. MORTALITY Predators include Puma and domestic dogs. LIFESPAN 14 years in captivity. Status and Threats Widespread, common and tolerant of agricultural/pastoral conversion, but legal control and bounties for perceived livestock depredation kill many tens of thousands, resulting in local population declines. CITES Appendix II; Red List LC, population trend Stable.
| CRAB-EATING FOX Cerdocyon thous |
CRAB-EATING ZORRO
HB 57–77.5cm; T 22–41cm; W 4.5–8.5kg
Medium-sized fox, with coarse, grizzled dark grey fur, giving it a dark bristly appearance. Muzzle and lower limbs usually conspicuously dark. Distribution and Habitat E Panama (a recent range expansion since 1999–2000 into agriculture-dominated landscapes), N Colombia, Venezuela, E and S Brazil, E Bolivia, Paraguay, N Argentina and Uruguay. Occupies all kinds of forest, woodland, grassland and marshland to 3,690m (Colombia), as well as pastoral and agricultural habitats. Feeding Ecology Omnivorous, with a catholic diet, especially fruits, insects and small mammals. Also eats birds, reptiles, eggs, amphibians, land crabs, insects and carrion. A pair was filmed killing a 2.5m Boa Constrictor (Mato Grosso do Sul, Brazil). Raids poultry and kills small lambs (rarely). Foraging is mainly nocturno-crepuscular and as pairs with pups, or alone. Congregates in larger groups on turtle-nesting beaches and at carcasses. Social and Spatial Behaviour Monogamous, forming mated pairs occupying exclusive territories. Yearling offspring often remain with the resident pair, forming family groups of up to 7. Dispersers may settle near their parents and interact amicably as adults. Range size for adults 0.5–10.4km2, averaging 2.2–5.3km2. Density estimates 0.55/km2 (Brazilian scrub savannah) to 4/km2 (Venezuelan Llanos). Reproduction and Demography Possibly seasonal, with dry-season breeding peaks, although births occur year-round in some areas. Gestation 52–59 days. Litter size 3–6. Pups weaned at 12 weeks. MORTALITY Killed by larger carnivores like domestic dogs; common as roadkill. LIFESPAN 9.2 years in the wild, 11.5 in captivity. Status and Threats Widespread, common and adaptable; deforestation to agricultural mosaics actually favours the species and it has expanded its range in some areas. CITES Appendix II; Red List LC, population trend Stable.
| SHORT-EARED DOG Atelocynus microtis |
SMALL-EARED DOG
HB 72–100cm; T 25–35cm; SH c.35cm; W 9–10kg
Medium-sized canid with short, sleek, uniformly coloured fur ranging from very dark brown to rufous-grey. Face long and slender, and ears conspicuously small. Classified in its own genus; most closely related to Bush Dog. Distribution and Habitat Restricted to W lowland Amazonia, in Colombia, Ecuador, Brazil, Peru and Bolivia. Inhabits undisturbed primary forest and is usually associated with rivers. Avoids human presence and disturbed habitats. Feeding Ecology Poorly known; fish dominate the diet in Cocha Cashu, Peru. Other food includes agoutis, small marsupials, rodents, birds, crabs, frogs and fruits. Has been observed hunting in waterholes and swimming after prey; its elongated shape, sleek fur and partially webbed toes may be adaptations for semi-aquatic hunting. Social and Spatial Behaviour Poorly known. Most sightings are of individuals, suggesting it is solitary, but adult pairs have been observed foraging together. Reproduction and Demography Poorly known. Observed litters cluster in the dry season (May–December). MORTALITY Unknown. LIFESPAN 11 years in captivity. Status and Threats Naturally rare and relies on intact forest. Amazon deforestation is a serious threat, and domestic dog diseases were implicated in an apparent population decline in Peru. Red List NT, population trend Decreasing.
Plate 57
| BUSH DOG Speothos venaticus |
HB 57.5–75cm; T 11–15cm; SH c.30cm; W 5–8kg
Small, stocky dog unlike any other canid, with a long body, short legs and short, bushy tail. Broad head is bear-like with small eyes and short, rounded ears. Body colour varies from blond to dark brown; usually tawny blond on the neck and head, and dark brown on the legs and tail. Distribution and Habitat N Argentina and N Paraguay, through much of Brazil into N South America, N Ecuador to Panama–Costa Rica border; first confirmed in Costa Rica (Las Tablas Protected Zone, Talamanca Mountains) in 2016. Closely tied to intact forest and forest savannah, including well-vegetated cerrado and pampas grassland. Occurs on ranchland and in agricultural areas, but is dependent on forest fragments in disturbed habitats. Feeding Ecology Eats mainly small mammals up to its own weight, especially pacas, agoutis, Nine-banded Armadillo, Brazilian Rabbit, opossums and rats. Large terrestrial reptiles, e.g. tegu lizards, and birds, e.g. tinamous, are also taken. Packs supposedly kill large prey, including Capybara, brocket deer and rhea, by biting the legs until the quarry tires: a report exists of a pack of 6 harassing and badly wounding an adult Brazilian Tapir. Sometimes eats Cecropia fruits. Local people report that it occasionally takes chickens. Foraging is mostly diurnal, and it sleeps in burrows overnight; there are a handful of observations of nocturnal hunting. Hunts socially, mainly by prolonged pursuit through thick vegetation, assisted by its long, squat body shape. Readily takes to water during pursuits, and enters and excavates burrows in pursuit of prey. Social and Spatial Behaviour Most social of small canids, living in small packs of 2–12 (generally 2–6) that are in constant close contact, including sleeping in groups. Pack composition is thought to comprise a monogamous breeding pair with its grown offspring. Pack members help raise pups by provisioning mothers at the den during nursing. Spatial behaviour is poorly known; the few recorded range sizes are very large for such a small canid: a pack inhabiting mainly native cerrado savannah forest covered 141km2, compared to 709km2 for a pack occupying mainly ranchland and agriculture with cerrado fragments (both packs in Mato Grosso, Brazil). Reproduction and Demography Aseasonal, although birth peaks possibly occur in the wet season. Gestation averages 67 days (65–83 days in captivity). Litter size 3–6, exceptionally to 10. Pups weaned from 4 weeks. Sexual maturity at 10 (♀s) to 12 (♂s) months. MORTALITY Poorly known; predation by large cats is reported by local people. LIFESPAN 10.4 years in captivity. Status and Threats Widespread but nowhere abundant, and appears to be naturally rare. Conversion of forested habitat for livestock and agriculture is the main threat. Occasionally killed on roads and as a perceived predator of poultry. Captive individuals are vulnerable to canid diseases such as distemper and parvovirus. CITES Appendix I; Red List NT, population trend Decreasing.
| MANED WOLF Chrysocyon brachyurus |
HB 95–115cm; T 28–50cm; SH 70–74cm; W 20.5–30kg
Tall, very long-legged canid with tawny-rufous fur, dark on the neck and shoulders, with black socks and a white-tipped bushy tail. Face fox-like, with very large ears, a white throat and a black muzzle. Melanism is very rare, with a single record from Veredas do Acari Sustainable Reserve, SE Brazil. Despite its common name, it is not closely related to wolves and belongs in its own genus; its closest relative (but distant none the less) is thought to be Bush Dog. Distribution and Habitat C and S Brazil, E Bolivia, E Paraguay and NE Argentina; occasional records in extreme SE Peru and N Uruguay. Inhabits primarily cerrado and pampas savannah, woodland–savannah mosaics, shrub forest and seasonally flooded wetland such as the Pantanal. Occurs on ranchland and in agricultural areas with cover. Feeding Ecology Omnivorous, with a very broad diet of both plant and animal matter; at least 102 fruits and 157 animal species are recorded in Brazil alone, although small vertebrate prey is most important by weight. Common prey includes mice, spiny rats, cavies and small armadillos, as well as small birds, arthropods and (less so) reptiles, including venomous snakes. Occasionally kills tamandua and brocket deer; one record of predation on an adult Pampas Deer, which was killed by a throat bite. Wolf Apple is consumed year-round and is the most important fruit in the diet. Diet expands in the wet season to include more fruits, such as bell peppers, coffee and papaya, as well as new grass. Readily kills chickens, and possibly takes juvenile small stock, although reports are equivocal. Foraging is cathemeral and solitary. Scavenges, including from roadkill carcasses and human refuse. Social and Spatial Behaviour Basic social unit is a monogamous pair that shares and defends a common territory from other pairs. Most behaviour, including hunting, is solitary; pairs occasionally rest and travel together, but longer associations occur only when breeding. Territories are stable, marked constantly with urine and faeces, and used equally by both pair members. No difference in range size between males and females, except that female ranges decrease during breeding. Resident pair territory size averages 75km2 (cerrado, Noel Kempff Mercado NP, Bolivia) and 80.2km2 (cerrado, Emas NP, Brazil); individual non-territorial ranges in Emas NP average 188km2. Densities are naturally low, 1.6–8/100km2 in Brazilian and Bolivian protected cerrado and Pantanal. Reproduction and Demography Seasonal. Mating April–June; most births occur in the dry season, June–September. Gestation 56–67 days. Litter size 1–7, averaging 3. Weaning at around 15 weeks, but pups may accompany the mother from 7 weeks. Role of the male in raising pups is unclear; captive males regurgitate food and groom pups, and wild males often accompany mothers and young pups. Sexual maturity at 12 months, but first breeding is probably during the second year. MORTALITY Annual adult and subadult mortality 35% and 36%, respectively (Emas NP). Most mortality is anthropogenic; Puma and domestic dogs are confirmed predators. LIFESPAN 15 years in captivity. Status and Threats Lives at naturally low densities, and its habitat is under intense pressure from agricultural development. In Brazil, roadkills are one of the leading causes of death, and domestic dogs on ranches often kill Maned Wolves; dogs may also be a source of exotic disease, although effects on populations are unknown. Frequently killed for folkloric medicine, especially in Bolivia, and in retribution for depredation on poultry. CITES Appendix II; Red List NT, population trend Unknown.
Plate 58
