Lycaon pictus ( Temminck, 1820 )

Bucci, Melanie E, Nicholson, Kerry L & Krausman, Paul R, 2022, Lycaon pictus (Carnivora: Canidae), Mammalian Species 54 (1017), pp. 220-240 : 221-225

Lycaon pictus ( Temminck, 1820)

African Wild Dog

Canis aureus : Thunberg, 1811:302. Not Canis aureus Linnaeus, 1758 .

Hyaena picta Temminck, 1820:54 , plate XXXV. Type locality “á la côte de Mosambique” (= coast of Mozambique).

Hyaena venatica Burchell, 1822: 456 . Type locality “ Northeast of Asbestos Range, upper Orange River.”

Canis pictus : Desmarest, 1822:538. Name combination.

Lycaon tricolor Brookes, 1827:151 . Type locality “ Cape of Good Hope.”

Cynhyaena picta : F. Cuvier, 1829:454. Name combination.

Lycaon typicus A. Smith, 1833:91 . Type locality “ South Africa.”

Lycaon pictus : A. Smith, 1833:91. First use of the current name combination.

Hyenoides picta : Boitard, 1842:214. Name combination.

Kynos (Canis) pictus : Rüppell, 1842:163. Name combination.

Lycaon venaticus : Gray, 1868:497. Name combination.

Lycaon pictus venaticus : Thomas, 1902:438. Name combination.

Lycaon pictus lupinus Thomas, 1902:439 . Type locality “Nyuki River Swamp, in the Rift Valley on Equator.Altitude 2000 m.”

Lycaon pictus somalicus Thomas, 1904:98 . Type locality “Central Somaliland.”

Lycaon pictus zuluensis Thomas, 1904:98 , footnote. Type locality “Pongola River,” Zululand, KwaZulu-Natal, South Africa.

Lycaon pictus sharicus Thomas and Wroughton, 1907:375 . Type locality “Maui, Shari River Chari] River,” Chad.

Lycaon pictus prageri Matschie, 1912:311 . Type locality “vom Nzoia-Flusse auf dem Guasso-Ngischu-Hochlande südlich des Elgon” (= Nzoia River on Guasso Ngischu Uplands south Elgon).

Lycaon lalandei Matschie, 1915:313 . Type locality “aus der Gegend zwischen der Algoa-Bucht und dem Keiskama- Flusse” [= between Algoa Bay and the Keiskama River],” Cape Province, South Africa.

Lycaon rüppelli Matschie, 1915:315 . Type locality “der Umgebung der Bajuda-Steppe stammt” [= area around Bajuda steppe],” northeastern Africa.

Lycaon lademanni Matschie, 1915:315 . Type locality “Mbarangandu in der Nähe der Stasse Songea-Liwale in Deutsch-Ostafrika ” (= Mbarangandu near the Songea-Liwale road in German East Africa [= Tanzania]).

Lycaon ssongaeae Matschie, 1915:317 . Type locality “bei Ussangire, 14 km westsudwestlich von Ssongae im Quellgebiete des Rovuma, Deutsch-Ostafrika (= near Ussangire, 14 km west-southwest of Songae in the headwaters of the Ruvuma River, German East Africa [= Tanzania]).

Lycaon hennigi Matschie, 1915:320 . Type locality “Makangaga, ungefähr 40 km westsüdwestlich von Kilwa (= Makangaga, about 40 km west-southwest of Kilwa [= Tanzania]).

Lycaon stierlingi Matschie, 1915:323 . Type locality “Rijuni, einem Nebenflusse des oberen Rovuma im Bezirke Songea, Deutsch-Ostafrika, ungefähr 30 km von den Quellen des Mbarangandu-Luwegu” (= Rijuni, a tributary of the upper Ruvuma River in Songea district, German East Africa [= Tanzania], about 30 km from the springs of Mbarangandu-Luwegu).

Lycaon huebneri Matschie, 1915:326 . Type locality “Kibwesi an der Uganda-Bahn [= Kibwesi on the Uganda Railway],” Kenya.

Lycaon ruwanae Matschie, 1915:328 . Type locality “der Ruwana-Steppe nordöstlich von der Speke-Bucht des am Viktoria -Nyansa in Deutsch-Ostrafrika” (= Ruwana-steppe northeast of Speke Bay in Lakeon the Victoria Nyanzain German East Africa [= Tanzania]).

Lycaon kondoae Matschie, 1915:331 . Type locality “Bubu bei Kondoa-Irangi in Deutsch-Ostafrika ” (= Bubu River at Kondoa-Irangi in German East Africa [= Tanzania]).

Lycaon langheldi Matschie, 1915:333 . Type locality “Njawa in Ussomgo, nordöstsüdllich von Ussongo Tabora in der westlichen Wembaeäre-steppe, Deutsch-Ostafrika ” (= Najwa in Ushomgo, south northwest of Ussongo Tabora in the western Wembaeäre-steppe, German East Africa [= Tanzania]).

Lycaon dieseneri Matschie, 1915:336 . Type locality “Myombo- Walde zwischen Usinsa und OstUssuwi im Süden am des Viktoria Nyansa in Deutsch-Ostafrika ” (= Myombo forest between Usinsa and east Ussuri on the south of Lake Victoria in German East Africa [= Tanzania]).

Lycaon gansseri Matschie, 1915:340 . Type locality “Bezirke Tabora [Tabora districts],” Tanzania.

Lycaon taborae Matschie, 1915:344 . Type locality 1/2 stunde von Tabora ” [= one-half hour south of Tabora],” Tanzania.

Lycaon wintgensi Matschie, 1915:347 . Type locality “m Luegere- Fluss an der Kungue-Bucht des Ostufers de Tanganjika in Deutsch-Ostafrika ” (= on the Luegere River on Kungue Bay on the east bank of the [Lake] on Tanganyika in German East Africa [= Tanzania]).

Lycaon richteri Matschie, 1915:350 . Type locality “ Daressalam [= Dar es-Salaam],” Tanzania.

Lycaon styxi Matschie, 1915:352 . Type locality “Mpapua in Deutsch-Ostafrika [ German East Africa],” Tanzania.

Lycaon luchsingeri Matschie, 1915:355 . Type locality “Edaballa, Mulla, nahe dem Hauasch im Danakil-Lande [=Edaballa, Mulla, near the Awash River in the Danakil Desert],” Ethiopia.

Lycaon zedlitzi Matschie, 1915:358 . Type locality “Scetel-Flusse in der Buschsteppe, ungefähr 800 m über dem Meere erlegt … östlichsten Quellfluss des Barka südöstlich von Agordat” (= Scetel River in the bush steppe, about 800 m above sea level … easternmost source river of the Barka [River] southeast of Agordat [ Eritrea]).

Lycaon takanus Matschie, 1915:360 . Type locality “Landschaft Taka [= Taka Mountain landscape, Sudan, northeastern Africa].

Lycaon manguensis Matschie, 1915:364 . Type locality “Djannaga nördlich vou Sansanne Mangu unweit des Oti in Togo ” (= Djannaga north of Sansanneé-Mango near Oti in Togo).

Lycaon mischlichi Matschie, 1915:366 . Type locality “nördlichen Teile des Bezirkes Kete-Kratschi in Togo, wahrscheinlich bei Bimbila, der Hauptstadt von Nanumba, zwischen dem Oti und Daka [= northern part of district Kete-Kratschi in Togo, possibly near Bimbila, the capital of Nanumba, between Oti and Daka].”

Lycaon ebermaieri Matschie, 1915:369 . Type locality “aus der Grasländern am Tschad-See, wahrscheinlich aus der Gegend von Dikoa” (= grasslands of Lake Chad, probably from Dikwa area [= northeastern Nigeria]).”

Lycaon fuchsia Matschie, 1915:371 . Type locality “Rio Cubal in Benguella” (= Cubal River in Benguella [= western Angola]).

Lycaon cacondae Matschie, 1915:373 . Type locality “Caconda in Benguella” (= Caconda in Benguella [= western Angola]).

Lycaon gobabis Matschie, 1915:373 . Type locality “Gobabis in Deutsch Südwestafrika ” (= Gobabis of German Southwest Africa [= Namibia]).

Lycaon krebsi Matschie, 1915:376 . Type locality “Bavians Revier- Fluss im Kaplande” (= Baviaans territory in [Eastern] Cape [ South Africa]).

Lycaon windhorni Matschie, 1915:378 . Type locality “Rustenburg, in Transvaal [now North West Province, South Africa].”

CONTEXT AND CONTENT. Context as for genus. Recent DNA analysis and research revealed there are no subspecies of Lycaon pictus , only geographical variants ( Sillero-Zubiri 2009).

NOMENCLATURAL NOTES. The generic name Lycaon can be traced to Greek mythology. Lycaon was a king who was turned into a wolf because he insulted a god. Lycaon , however, is not the word for “wolf” in Greek; it is “lycos” ( Borror 1960). The specific epithet pictus is from Latin meaning “painted or variegated,” aptly describing the painted coat of L. pictus . Other common names include Cape hunting dog ( Skinner and Chimimba 2005) and painted hunting dog ( Sillero-Zubiri 2009:355).

DIAGNOSIS

Lycaon pictus is the only extant species in the genus ( Nowak 1991; Girman and Wayne 1997). Compared with the other canids within its geographic distribution (African golden wolf [ Canis lupaster ], side-striped jackal [ Lupulella adustus ], black-backed jackal [ L. mesomelas ], and bat-eared fox [ Otocyon megalotis ]— Burgin et al. 2020), L. pictus is the largest, and lightly built with large, rounded ears ( Nowak 1991). Lycaon pictus has four toes on each foot, differentiating it from other canids, and legs are long and slender (van Lawick 1970; Nowak 1991). The most distinguishing feature of L. pictus is its unique coat coloration of irregularly mottled black, yellow-brown, and white that can be used to identify individuals ( Fig. 1 ; Wilson and Mitttermeier 2009).

GENERAL CHARACTERS

Lycaon pictus has a head–body length of 76–112 cm, tail length of 30–41 cm, shoulder height of 61–78 cm, and body weight of 17–36 kg ( Kingdon 1977, 2015). The second and third molars are reduced and smaller than the paracone (antero-external cusp) of the first molar. Palatal width between carnassials is greater than one-half the palate length measured from anterior edge of the canines ( Meester and Setzer 1971). The skull is large (length> 180 mm) and robust, and teeth are adapted for holding and slicing rather than grinding. Zygomatic arch is heavily built with a width> 120 mm ( Fig. 2 ; Meester and Setzer 1971; Skinner 2005). The pollex (i.e., big toe) is absent ( Meester and Setzer 1971).

Morphological and genetic differences occur in populations of L. pictus from eastern and southern Africa, indicating genetic differentiation and population structure throughout its geographic distribution. Although sizes and weights vary among populations ( Haltenorth and Diller 1980; Girman et al. 1993), genetic differences indicate recent (<100 years) interrupted gene flow between eastern and southern populations but not enough to identify distinct clades corresponding to subspecies ( Girman and Wayne 1997). There are no noticeable differences between sexes other than a 3–7% size difference in skeletal and musculature characteristics ( Girman et al. 1993; Creel and Creel 2002). Differences in back length and chest girth of males and females are positively correlated with prey density (back length: males, r = 0.33, P <0.001; females, r = 0.46, P = 0.002 and chest girth: males, r = 0.23, P = 0.026; females, r = 0.24, P = 0.123— McNutt and Gusset 2012).

Pelage of L. pictus is irregularly mottled black, yellow-brown, and white ( Van der Merwe 1959; Meester and Setzer 1971; Nowak 1991; Wilson and Mitttermeier 2009) and is unique to individuals ( Skinner and Chimimba 2005). Tip of the tail is almost always white, forehead is generally light, and muzzle is dark ( Kingdon 1977, 2015; Nowak 1991; Skinner and Chimimba 2005). Ears are large, round, and covered with short dark hairs with tufts of light hair on the inside ( Nowak 1991; Skinner and Chimimba 2005). Hair on the body is short and sparse, and there is no undercoat ( Skinner and Chimimba 2005).

DISTRIBUTION

Lycaon pictus was once widespread throughout sub-Saharan Africa ( Skinner and Chimimba 2005), inhabiting nearly all environments except rain forests ( Fanshawe et al. 1991) and the driest deserts ( Schaller 1972). Lycaon pictus has been reported from Mount Kilimanjaro ( Thesiger 1970) to deserts in central Sahara ( Monrod 1928; Lhote 1946). Historically, a relict population in Algeria suggests that arid lands might have been inhabited by L. pictus ( Fanshawe et al. 1997) . It has been virtually extirpated from western Africa and reduced in central and northeastern Africa. Botswana, Tanzania, and Zimbabwe provide habitat for nearly one-half of the remaining wild L. pictus . Other ( Fanshawe et al. 1997). A survey of 43 African countries reported 17 countries with viable or potentially viable populations of L. pictus ( Angola, Bein, Botswana, Burkina Faso, Central African Republic, Chad, Ethiopia, Kenya, Malawi, Mozambique, Namibia, Senegal, South Africa, Sudan, Tanzania, Zambia, and Zimbabwe), 11 countries with extirpated populations ( Burundi, Cameroon, Egypt, Eritrea, Eswatini, Gabon, Gambia, Ghana, Mauritania, Rwanda, and Sierra Leone), seven countries with nearly extirpated populations ( Democratic Republic of the Congo, Côte d’Ivoire, Guinea-Bissau, Mali, Nigeria, Togo, and Uganda), six countries having no verifiable records of L. pictus ( Djibouti, Equatorial Guinea, Lesotho, Liberia, Mauritania, and Western Sahara), and two countries with uncertain presence ( Algeria and Guinea — Fanshawe et al. 1997; Woodroffe and Sillero-Zubiri 2020).

FOSSIL RECORD

The fossil record for Lycaon pictus is sparse, and the earliest known indisputable fossil of L. pictus is from the mid- Pleistocene in Africa ( Savage 1978). Identification of early L. pictus fossils has been difficult because of their similarity to the early Pleistocene wolf, Canis africanus ( Malcolm 1979) . Whether late Pleistocene ( Kurtén 1968) and European L. pictus fossils are Canis is debated ( Thenius 1972). Lycaon pictus probably proliferated in Africa 2–3 million years ago ( Savage and Russell 1983), with major radiations occurring north and south of the miombo woodland belt ( Creel and Creel 2002). populations occur in Central African Republic, Ethiopia, Kenya, Mozambique, Namibia, South Africa, Sudan, and Zambia, with potential populations of <100 individuals in Cameroon, Shad, Senegal, and Somalia ( Fig. 3 ; Wilson and Mitttermeier 2009).

There are about 6,700 L. pictus , with about 1,400 mature adults, distributed throughout Africa ( Woodroffe and Sillero-Zubiri 2012, 2020). Populations throughout northern, eastern, and western Africa are low density; they are patchy in central Africa, and the majority of individuals occur in southern Africa

FORM AND FUNCTION

Feet of Lycaon pictus are digitigrade and reduced, with the four toes on each foot allowing for improved cursorial ability ( Girman et al. 1993). The only common pelage characteristic generally shared by all individuals is a white tip on the tail that might serve to keep individuals together when the pack is moving through dense, tall vegetation ( Fig. 4 ; Estes and Goddard 1967). Dental formula is i 3/3, c 1/1, p 3/4, and m 3/3, total 42; m3 is vestigial ( Van Valkenburgh 1989).

The preputial sebaceous glands of male L. pictus secrete their contents directly into the preputial lumen or the hair shaft, which indicates urine is mixed with glandular secretions before released. Secretions from the anal sacs and sebaceous glands (that occur over the entire body) might be useful in scent marking, with deposition of the scent accomplished through anal-dragging behavior ( Van Heerden 1981). During nonmating periods, subordinate females had higher levels of estrogen and estrogen:progesterone ratios indicating ovulation suppression ( Creel et al. 1997b), and subordinate males had increased glucocorticoid secretion ( Van Heerden and Kuhn 1985; Creel et al. 1996).

Hematological and biochemical values of the peripheral blood in six captive male and seven female L. pictus from a zoo in the Czech Republic did not differ at the 5% level except for the erythrocyte count (P = 0.031) and sodium level (P = 0.044— Pospíšil 1987). Mean erythrocyte counts (1012 /l) were 9.86 for females and 8.45 for males, and mean leukocyte counts (109 /l) were 13.03 for females and 12.88 for males ( Pospíšil 1987).

ONTOGENY AND REPRODUCTION

Ontogeny.— Newborn Lycaon pictus are altricial and weigh about 300 g at birth. Eyes open after about 13 days, but young remain in the den for up to 3 weeks when they begin to take solid food regurgitated by other pack members. Young begin following the pack by 3 months of age and can kill prey by 11 months, although they remain inefficient at hunting until 12–14 months after birth. Social restrictions can mask the age at which L. pictus is first capable of copulating, but 21-month-old males and 22-month-old females have copulated successfully ( Frame et al. 1979).

Reproduction. —Estrus in Lycaon pictus gradually builds during the weeks prior to copulation, at which time the vulva becomes swollen with a bloody discharge. The bond between the alpha pair develops and becomes more overt ( Creel and Creel 2002). Early attempts by the male to mount the female are rejected until she is ready and stands “firmly with her tail averted to one side as the male mounts” ( Creel and Creel 2002:204). Copulations occur over 3–7 days but not all attempts end in copulation. The copulatory lock (i.e., bulbis glandis of the penis enlarges preventing the pair from disengaging—van Lawick 1970; Reich 1981) can last 5–6 min (van Lawick 1970; Frame et al. 1979; Malcolm 1979; Creel 2002) during which the pair is highly vulnerable ( Skinner and Chimimba 2005).

Gestation lasts 71–73 days, and young are born in underground dens, usually an abandoned burrow ( Sillero-Zubiri 2009). Females have 12–14 mammae (van Lawick 1970). Young suckle for up to 12 weeks. In one case, pack members raised a litter of 5-week-old young whose mother had died; thus, continued suckling might not be a necessity ( Nowak 1991).

Females produce litters every 12–14 months, but this interval can be reduced if a litter is unsuccessful ( Frame et al. 1979). Average litter size is 10 young (range = 6–16 young— Nowak 1991). Lycaon pictus copulates seasonally at latitudes from 7– 25°S generally producing a single litter per year within a pack, coinciding with peak prey abundance and cooler temperatures ( Malcolm 1979; Reich 1981; McNutt et al. 2019). Reproduction is aseasonal at latitudes <2° ( McNutt et al. 2019). In Kruger National Park (KNP), young were born in April–September, peaking in late May and early June ( Reich 1981). Inbreeding has occurred in KNP ( Reich 1978). In Selous Game Reserve, peak birthing occurred in July during the driest part of the year when ungulates congregated at water sources ( Creel and Creel 2002). On the Serengeti Plains, peak birth rates occurred during the rainy season in March–June when ungulates underwent annual migration ( Schaller 1972; Frame et al. 1979). Births occurred on average on 3 and 4 June corresponding to conception dates of 23 and 24 March in Botswana and Zimbabwe, respectively ( McNutt et al. 2019). In northern Botswana, 96.3% of litters were born in May–July ( McNutt 1996a). Intervals between births are 12–14 months if young survive but can be as short as 6 months if they do not ( Frame et al. 1979; Malcolm 1979). Captive populations averaged 11.7 months between litters ( Van Heerden and Kuhn 1985).

In some packs of L. pictus , sex ratio of male and female offspring at birth is skewed toward males ( Frame et al. 1979), but it is equal in others ( Creel et al. 1995; Somers et al. 2008). Although many authors cite a male-biased sex ratio at all ages ( Fuller et al. 1992), there is extensive regional variation, and it is unclear if a skewed sex ratio is a distribution-wide characteristic or an artifact of certain populations. In northern Botswana, sex biases were most prominent in young females in smaller packs rather than older females in larger packs ( McNutt and Silk 2008).

Figures

Fig. 1.—Adult Lycaon pictus in Pilanesberg Game Reserve, North West Province, South Africa. Photograph courtesy of Edward Peach (https:// www.kruger-2-kalahari.com/photographing-african-wild-dogs.html)

Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view of mandible of an adult female Lycaon pictus (National Museum of Natural History, Smithsonian Institution # 368442) from Mabeleapudi, Botswana, Africa.

Fig. 3.—Distribution of Lycaon pictus including eight categories: resident, possible occurrence, recoverable areas, transient, connecting areas, unknown, extirpated, and areas outside its distribution. Modified by K. L. Nicholson and used with permission of R. Groom (Range Wide Conservation Program for Cheetah and African Wild Dogs; http://www.cheetahandwilddog.org, accessed 23 June 2021).

Fig. 4.—Adult Lycaon pictus moving with intent during a hunting session in typical grassland habitat near Third Bridge Campsite, Moremi Game Reserve, Botswana, June 2014; note distinct coat pattern, mostly white tail, and lean body condition suggesting it had not recently fed. Photograph courtesy of Dave Gale used with permission.