Caracal caracal ( Schreber, 1776 )

Veals, Amanda M, Burnett, Alexandra D, Morandini, Marina, Drouilly, Marine & Koprowski, John L, 2020, Caracal caracal (Carnivora: Felidae), Mammalian Species 52 (993), pp. 71-85 : 72-76

Caracal caracal ( Schreber, 1776)

Caracal

Felis caracal Schreber, 1776 :pl. 110. Type locality “Vorgebirge der guten Hofnung,” restricted to “Table Mountain, near Cape Town, South Africa,” by J. A. Allen (1924:281).

Felis caracal Müller, 1776:30 . Type locality “Arabia” (but probably near Constantine, Algeria). Preoccupied by, but not, Felis caracal Schreber, 1776 ; see discussion in J. A. Allen (1924:281).

Felis caracal algiricus J. B. Fischer, 1829:210 . No type locality specified.

Felis caracal nubica J. B. Fischer, 1829:210 . No type locality specified, designated as “Meroe, Nubia [ Sudan],” by J. A. Allen (1924:281).

Felis caracal bengalensis J. B. Fischer, 1829:210 . No type locality specified; preoccupied by Felis bengalensis Kerr, 1792 .

Felis caracal var. algira J. A. Wagner (in M. Wagner), 1841: vol. 3, 76, atlas, Tab IV. Type locality “ Algeria.”

Caracal melanotis Gray, 1843:46 . Replacement name (unnecessary) for Felis caracal Schreber, 1776 ; see above.”

Lynx lucani Rochebrune, 1885:87 . Type locality “Landana,” northern Angola.

Caracal caracal: Matschie, 1892:114 . First use of current name combination.

Caracal berberorum Matschie,1892:114 . Type locality“Nordafrika. (Constantine …) [Constantine, Algeria]; ” corrected to “ Staouely, Algeria,” by Matschie 1912.

Felis (Caracal) berberorum spatzi Matschie, 1912:61 . Type locality “zwischen Feriana und Tebessa in den Aleppokiefernwaldungen zwischen Bir bou Haya und Ain bou Dries gesammelt hat [between Fériana and Tébessa, in the Aleppo pine forests between Bir Bou Haya and Ain Bou Dries],” Aleppo, Tunisia.

C [aracal]. berberorum medjerdae Matschie, 1912:62. Type locality “ Tunis,” Tunisia.

Felis (Caracal) nubicus corylinus Matschie, 1912:63 . Type locality “Tangiers,” Morocco.

Felis (Caracal) caracal schmitzi Matschie, 1912:64–65 . Type locality “Ain ed dschuheijir nordwestlich des Toten Meeres [Ain Dcherer, northwest of the Dead Sea, Palestine = Israel].”

Felis (Caracal) caracal aharonii Matschie, 1912:66 . Type locality “Einmündung des Chabur in den Euphrat [at the confluence of the Khabur and the Euphrates],” Syria.

Caracal caracal poecilotis Thomas and Hinton, 1921:3 . Type locality “Mt. Baguezan, Asben, 5,200 feet [=Mont Idoukal-n- Taghès (Mt. Bagzane), Asben, Air, Niger, 1,600 m].”

Caracal caracal coloniae Thomas, 1926:181 . Type locality “Deelfontein, central Cape Colony, South Africa.”

Lynx caracal roothi Roberts, 1926:247 . Type locality “Elandshoek, Drakensberg, Barberton district,” Transvaal, South Africa.

Lynx caracal limpopoensis Roberts, 1926:248 . Type locality “Nijellele River, north of Zoutpansberg and near the Limpopo River,” Northern Transvaal, South Africa.

Lynx caracal damarensis Roberts, 1926:248 . Type locality “Quickborn, Okahandja, S.W.A. [= Namibia].”

Felis (Caracal) caracal michaëlis Heptner, 1945:310 . Type locality Bokurdak, Turkmenistan.

[ Caracal caracal ] melanotix Wozencraft, 2005:533. Nomen nudum.

CONTEXT AND CONTENT. Context as for genus. The number of subspecies reported from different authors varies, mainly due to a lack of recent morphological and molecular studies that examine geographical variation in this species ( Kitchener et al. 2017). Wozencraft (2005) recognized eight subspecies of Caracal caracal based on Smithers (1971); algira, caracal , damarensis, limpopoensis, lucani , nubica, poecilotis, and schmitzi. Sunquist and Sunquist (2009) recognized C. c. michaelis ( Heptner 1945) as a ninth subspecies. The latest taxonomic revision of the Felidae proposes the possibility of as few as three subspecies, caracal , nubica, and schmitzi but suggests the need for further research to elucidate the geographical variation within C. caracal ( Kitchener et al. 2017) . The following are the eight subspecies presented by Wozencraft (2005):

C. c. algira ( Wagner, 1841). See above; berberorum Matschie, 1892 ; corylinus Matschie, 1912; medjerdae Matschie, 1912; and spatzi Matschie, 1912 are synonyms.

C. c. caracal ( Schreber, 1776) . See above; coloniae Thomas, 1926; melanotis Gray, 1843 ; and roothi ( Roberts, 1926) are synonyms.

C. c. damarensis ( Roberts, 1926). See above.

C. c. limpopoensis ( Roberts, 1926). See above.

C. c. lucani ( Rochebrune, 1885) . See above.

C. c. nubica (J. B. Fischer, 1829). See above.

C. c. poecilotis (Thomas and Hinton, 1921). See above.

C. c. schmitzi ( Matschie, 1912). See above; aharonii Matschie, 1912; bengalensis Fischer, 1829 ; and michaelis Heptner, 1945 are synonyms.

NOMENCLATURE NOTES. Though originally classified under the genus Felis ( Roberts 1951; Groves 1982; Meester et al. 1986; Heptner and Sludskii 1992), caracals have since been reclassified under a separate genus, Caracal . Early on Caracal caracal was aligned with the lynxes due to its prominent ear tufts and short tail, both characteristics shared with the Canadian lynx ( Lynx canadensis ), the bobcat ( Lynx rufus ), the Iberian lynx ( Lynx pardinus ), and the Eurasian lynx ( Lynx lynx ). However, C. caracal belongs to a much older lineage that diverged from the ancestral felid line more than 1 million years before the lynx evolved ( Johnson et al. 2006). C. caracal is most closely related to the African golden cat ( Janczewski et al. 1995; Johnson and O’Brien 1997; Johnson et al. 2006). A phylogenetic analysis indicated that the caracal lineage (serval, caracal, and African golden cat) diverged from other felids about 8.5 million years ago ( Janczewski et al. 1995; Pecon-Slattery and O’Brien 1998). The serval diverged about 5.6 million years ago, and the African golden cat and C. caracal bifurcated about 1.9 million years ago ( Johnson et al. 2006). Some authors continue to keep two different lineages for C. caracal and the African golden cat ( Profelis aurata — Rolland et al. 2015); however, a more recent classification groups the caracal and African golden cat in the genus Caracal ( C. caracal and C. aurata , respectively), and places the serval in the monotypic genus Leptailurus ( Leptailurus serval — Kitchener et al. 2017).

Buffon (1761) chose the name caracal in reference to its Turkic name “Karrah-kulak” or “Kara-coulac,” meaning cat with black ears. Other vernacular names for C. caracal are African lynx, caracal lynx, desert lynx, karakal, red cat, red lynx, and rooikat (Kenmuir and Williams 1975; Stuart 1982).

DIAGNOSIS

Caracal caracal is a slender, medium-sized cat characterized by a short tail and long ear tufts ( Hoath 2003; Fig. 1 ); no other African felid has distinct ear tufts. The body color is a uniform reddish brown in southern Africa ( Hufnagel 1972) but can be sable in Asia and the Arabic peninsula (Kingdon and Hoffmann 2013). These distinct physical characteristics help to distinguish C. caracal from the other six felids of southern Africa. C. caracal is sympatric with the lion ( Panthera leo ), leopard ( Panthera pardus ), cheetah ( Acinonyx jubatus ), serval, African wild cat ( Felis lybica ), and black-footed cat ( Felis nigripes ) in southern Africa (Caro and Stoner 2003). C. aurata is sympatric with C. caracal in portions of their respective geographic ranges, which can generate some confusion for identification. However, C. aurata selects for different habitat types than C. caracal and is found only in the humid equatorial rainforests ( Furstenburg 2010).

The key to distinguishing these two species is a combination of coat pattern and color, body size, shape of the ears, and the pattern, color, and carriage of the tail. C. caracal has triangular, black-backed ears with long (ca. 50 mm) black tufts on the tip. C. aurata has shorter ears (ca. 53 mm) that are solid black (Kingdon and Hoffmann 2013). The tail of C. caracal is sandy-red to golden-red throughout, and dangles nearly straight down when the cat is standing, not reaching the hocks. The tail of C. aurata is black-tipped, and when the animal is standing, the tip of the tail curves away from the body with the lowest part of the tail reaching the hocks (Kingdon and Hoffmann 2013).

GENERAL CHARACTERS

Classified as a medium-sized felid, head and body lengths for Caracal caracal are 61–106 cm with the largest individuals inhabiting South Africa and the smallest occurring in Israel and the Sahara Desert ( Smithers 1971; Stuart 1981; Nader 1984; Sunquist and Sunquist 2002; Furstenburg 2010). The hindquarters are set slightly higher than the forequarters ( Hufnagel 1972), with the height of the shoulders about 46 cm ( Hufnagel 1972). C. caracal has a relatively short (18–34 cm), robust tail compared to most cats, roughly one-third of the body length and it dangles straight down (Kingdon and Hoffmann 2013). Most individuals have furry paws; individuals from the Karakum Desert in Turkey have stiff bristle hair under the paws ( Furstenburg 2010). Both front and hind paws of adults have four toe cushions and are typically 50–60 mm long and 40–50 mm wide ( Furstenburg 2010). Weights vary between 5.8 and 22 kg across its geographic range. Adult males can weigh anywhere between 8 and 20 kg (average 12.7 kg), while females range between 6.2 and 15.9 kg (average 10.1 kg — Smithers 1971; Roberts 1977; Pringle and Pringle 1979; Smithers and Wilson 1979; Smithers 1983; Nader 1984; Weisbein and Mendelssohn 1990; Heptner and Sludskii 1992; Sunquist and Sunquist 2002; Sunquist and Sunquist 2009).

Caracal caracal body color is uniform golden or a rufous brown that explains the Afrikaans name of “rooikat” meaning “red cat”; the underbelly is cream with orange or brown spots, with the inner legs being pale (Kingdon and Hoffmann 2013). Intraspecific color variation is slight, with females generally lighter in color (Sunquist and Sunquist 2002). Animals from areas with low rainfall tend to be paler than those from higher rainfall regions (Kingdon and Hoffmann 2013). Melanistic individuals have been recorded in Kenya and Uganda ( Rosevear 1974). Guard hairs are 15–30 mm and seasonal variation can occur in coat thickness providing effective insulation from extreme temperatures ( Pocock 1939; Smithers 1983).

The head is rather angular and strikingly marked. The eyes are commonly brilliant green or yellow, and occasionally blue in color, contoured by a distinctive dark line, which runs from above the eye down along the tear line and down the nose. C. caracal exhibits a lowered upper eyelid and black tear streaks around the eyes, which are likely protective adaptations against the sun’s glare (Sunquist and Sunquist 2002). White fur covers the chin, throat, upper lips, and around the eyes. The eyes and lips are often lined with black ( Hufnagel 1972). The large, pointed ears measure up to 80 mm long including the tufts. These conspicuous ear tufts have been suggested to play a role in intraspecific communication (Nowell and Jackson 1996).

DISTRIBUTION

Caracal caracal has a very wide distribution, one of the most expansive of extant carnivores, ranging from the southern tip of the African continent, north to the Arabian Peninsula, the Middle East, and Turkey, eastwards to central India and northwards to Kazakhstan and Turkmenistan ( Fig. 2 ; Stuart 1984; Nowell and Jackson 1996; Sunquist and Sunquist 2002; Hoath 2003). It is widely distributed on the African continent, with the exception of the equatorial rainforests and the Sahara interior ( Ray et al. 2005). C. caracal is distributed across the drier regions of central Asia, and southwest Asia and India as well ( Avgan et al. 2016). It is present in the montane massifs of the Sahara Desert and its fringes, including the Adar des Iforas in northeastern Mali (Sidiyéne and Trainer 1990), Hoggar and Tassil mountains of southeastern Algeria and the Saharan Atlas ( De Smet 1989; Kowalski and Rzebik-Kowalska 1991), the Air of Niger (Dragesco-Joffé 1993), edges of the great sand areas of the Eastern Great Erg, and Tanzania ( Pettorelli et al. 2010). In Egypt, C. caracal is recorded only from the Eastern Desert and north and southwestern Sinai ( Hoath 2003). In Turkey, it is present in the southwest of the country, specifically in Datça and Bozburun peninsulas (Ilemin and Gürkan 2010); in Iran C. caracal is present in central and eastern areas of Iran ( Farhadinia et al. 2007; Ziaie 2008). The presence of C. caracal is equivocal in the savanna regions of the Congo basin ( Pettorelli et al. 2010), north of the Congo River, and in central Sinai ( Hoath 2003).

Sunquist and Sunquist (2009) reported distinct geographical areas for all nine of the subspecies described in the literature. C. c. caracal ( Schreber, 1776) is in eastern and southern Africa, C. c. algira ( Wagner, 1841) in northern Africa, C. c. damarensis ( Roberts, 1926) throughout Namibia, C. c. limpopoensis ( Roberts, 1926) north of the province Transvaal, C. c. lucani ( Rochebrune, 1885) in the grasslands of southeastern Gabon ( Coetzee 1977), C. c. michaelis ( Heptner, 1945) in desert regions of the Caspian Sea Region and east to Amu Darya River, C. c. nubica ( Fischer, 1829) eastern Cameroon and north to the Nubian Desert, C. c. poecilotis (Thomas and Hinton, 1921) in western Africa, and C. c. schmitzi ( Matschie, 1912) in Turkey, Palestine, Iran, and India.

Caracal caracal is still widely distributed throughout its historic range in Africa; however, there have been substantial losses along the peripheries. In northern and western Africa in particular, C. caracal has been locally extirpated from areas with high land conversion rates to livestock areas ( Ray et al. 2005). In Africa, it is estimated to have lost 37.7% of its historical distribution ( Ray et al. 2005), occurring less frequently and in patchily distributed pockets of drier habitat (Nowell and Jackson 1996). However, C. caracal is most abundant in South Africa and Namibia, where its range is expanding (Stuart and Wilson 1988; Nowell and Jackson 1996), possibly due to local extirpation of the black-backed jackal Canis mesomelas by farmers (Pringle and Pringle 1979; Stuart 1982; Nowell and Jackson 1996). In Asia, C. caracal remains patchily distributed and limited by harsh winters with the Himalayas acting as a natural barrier preventing expansion further north (Nowell and Jackson 1996; Furstenburg 2010)

FOSSIL RECORD

The subfamily Felinae comprise the common ancestor of all living cats and the Felidae family originated near the end of the Eocene ( Werdelin et al. 2010). Modern felids arose in Asia 10.8 million years ago before the progenitor of the caracal lineage arrived in Africa between 8.5 and 5.6 million years ago ( Johnson et al. 2006). The earliest record of the caracal lineage is from about 4 million years from eastern and southern Africa ( Werdelin et al. 2010).

Based on molecular ages of lineages, the specimens from the caracal lineage group into two distinct size classes, large, represented by the stem containing the C. caracal and C. aurata lineage, and small represented by the serval stem ( Werdelin et al. 2010). However, whether these fossils are conspecific with extant species in the caracal lineage remains unclear. Available materials, mostly isolated teeth and fragmented jaws prevent further clarification of this ( Werdelin et al. 2010). “ Felis ” issiodorensis, a species that is generally placed in the genus Lynx , may be placed in the Caracal genus ( Morales et al. 2003) based upon biometric analyses that show similarities to caracal specimens, a possibility deserves further study ( Werdelin et al. 2010).

FORM AND FUNCTION

Caracal caracal is part of the subfamily Felinae , the conical tooth cats, due to the presence of a rounded canine cross-section ( Werdelin et al. 2010) and relatively large, well-enameled teeth ( Smithers 1983). The second upper premolars are typically absent in C. caracal (Skinner and Chimimba 2006) and in a sample of 100 skulls, P2 was present in only eight (Skinner and Chimimba 2006). The dental formula of C. caracal is I 3/3, C 1/1, P 3/2, M 1/1, total 30 (Skinner and Chimimba 2006). The diastema on the upper jaw and small lower incisors allows for the upper canines to sink to full depth into prey (Pringle and Pringle 1979). C. caracal has relatively large canines; Christiansen and Adolfssen (2005) reported an upper canine length of 18.4 mm and lower canine length of 17.0 mm for an adult specimen while Pringle and Pringle (1979) reported upper canine lengths up to 20 mm. The skull and dentition of C. caracal may have evolved to handle relatively large prey in proportion to its body size (Pringle and Pringle 1979).

Among extant felids, C. caracal is characterized by relatively large morphological features with a rounded, blunt rostrum ( Fig. 3 ; Sicuru 2011). The sagittal crest is distinctive, providing extra surface area for jaw muscle attachment indicating a powerful bite force. Additional leverage for muscles involved in closing the jaw is provided by a thick mandible with a tall coronoid process ( Smithers 1983). Orbital sockets and auditory bullae are relatively large, indicating keen senses of vision and hearing ( Smithers 1983). From the posterior most part of occipital to anterior most extension of nasals, total skull length was 103.52 mm for a typical female specimen collected from Sudan. Furstenburg (2010) reported a total skull length of 150 mm, width of 113 mm, and bite width of 37 mm, which the author measured based on adapted methodology from Smithers (1983).

Muscle fibers of C. caracal are considered three times more powerful than those found in humans ( Neils 2018). Powerful hindquarters enable C. caracal to leap vertically ≥ 3 m in the air to capture avian prey ( Furstenburg 2010; Kohn and Noakes 2013). Relatively large paws with stiff guard hairs on the underside may be advantageous for moving thorough soft substrate such as sand (Heptner and Sludskii 1992) and provide protection from extreme midday heat in more arid environments ( Furstenburg 2010). The forepaw has an additional fifth digit, a sturdy dewclaw used for hunting ( Grobler 1981).

ONTOGENY AND REPRODUCTION

Kittens are born blind, opening their eyes at 6–10 days. Ears will lie flat on the head at birth and only become erect after 30 days when the tufts on ear tips are about 10 mm long ( Furstenburg 2010). Cubs will begin to hunt with the mother 4–6 months after weaning and their canines erupt (Sunquist and Sunquist 2009). Juveniles may leave the natal range by 9–10 months or may remain with the mother for about a year (Sunquist and Sunquist 2009).

Caracal caracal females reach sexual maturity between 7 and 12 months and males between 9 and 14 months (Bernard and Stuart 1987). However, males likely do not begin mating until they are between 18 and 20 months due to older, dominant males controlling higher quality territories ( Furstenburg 2010). Females likely do not begin mating until 14 months of age and can remain reproductive until 18 years of age ( Furstenburg 2010). Females are polyestrous and estrus cycles last anywhere from 1 to 6 days (Nowell and Jackson 1996). Estrus depends on female nutritional status, which may be why spermatogenesis in males is aseasonal with a reduction in reproductive activity in the winter (Bernard and Stuart 1987). Reproduction is timed with the phenology of prey species, peaking when vegetation, and therefore prey, production is highest. Thus, timing of reproduction varies by geographic location with usually only a single litter produced each year (Bernard and Stuart 1987). However, for captive C. caracal , births can occur throughout the year with a pronounced peak in the summer (Bernard and Stuart 1987). Copulation tends to be short and gestation period averages 78–81 days. Births are typically separated by 1 year (Bernard and Stuart 1987) but consecutive litters can be spaced approximately 10 months apart ( Furstenburg 2010). Litters average 2–4 kittens (Bernard and Stuart 1987) but up to six kittens may be born (Sunquist and Sunquist 2009).

There has been a greater awareness that many felid species are at risk of extinction, which has led to exploration of the possibility of using assisted reproductive technology for captive breeding programs ( Pope et al. 2006). Several studies have been conducted on the reproductive biology of captive C. caracal to gain insights for breeding programs. In fecal samples, progesterone levels increased in pregnant females and estradiol levels peaked near estrus ( Graham et al. 1993). Additionally, sexual behaviors were positively correlated with an increase in fecal estradiol while progesterone only rose after mating or during pregnancy ( Graham et al. 1995). Fecal PGFM (13,14-dihydro- 15-keto-prostaglandin F2α) levels increased 41 days after mating until parturition, then returned to basal levels postparturition ( Dehnhard et al. 2012). Many felids have difficulty producing offspring in captivity and with advancements of in vitro fertilization (IVF) systems for the domestic cat ( Felis catus ), preliminary studies have been conducted to test the viability of these systems for captive wild felids, including C. caracal ( Goodrowe et al. 1991) . Female C. caracal responded positively to pregnant mare’s serum gonadotropin (PMSG) treatments and mature oocytes could be collected using laparoscopic techniques ( Goodrowe et al. 1991). Two live kittens were produced from females that received auto-transferred day-4 or day-5 embryos ( Pope et al. 2006). C. caracal embryos were successfully cryopreserved before thawing and transferring to recipients on days 5 or 6. This had a 33% success rate for establishing pregnancies which led to a total of three kittens being born ( Pope et al. 2006).

Figures

Fig. 1.—Subadult Caracal caracal from Anysberg Nature Reserve, South Africa. Photograph by Marine Drouilly.

Fig. 2.—Geographic distribution of Caracal caracal. Map redrawn from IUCN Red List of Threatened Species.The type locality for each subspecies is indicated within the historic range of C. caracal: 1) C. c. caracal; 2) C. c. algira; 3) C. c. damarensis; 4) C. c. limpopoensis; 5) C. c. lucani; 6) C. c. nubica; 7) C. c. poecilotis; 8) C. c. schmitzi.

Fig. 3.—Dorsal, ventral, and lateral views of skull and lateral view of mandible of an adult female Caracal caracal (FMNH [Field Museum of Natural History] 105607) collected by H. H. Hoogstraal (collector # 13919) on 9 December 1972 at Africa, Sudan, Kassala, Ethiopian: Kassala, Abu Gamal. Total skull length is 103.52 mm. f8e611ef-14e0- 431f-acd8-8955c8521c97 Field Museum of Natural History (Zoology) Mammal Collection. https://collections-zoology.fieldmuseum.org/ catalogue/2604376.