Procavia capensis, Pallas, 1766

Don E. Wilson & Russell A. Mittermeier, 2011, Procaviidae, Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona: Lynx Edicions, pp. 41-47 : 43-44

1.

Rock Hyrax

Procavia capensis

French: Daman des rochers / German: Klippschliefer / Spanish: Daman de rocas

Other common names: Klipdassie; Abyssinian Hyrax (habessinica), Cape Hyrax ( capensis ), Johnston Hyrax (johnstoni), Kaokoveld Hyrax (welwitschii), Western Hyrax (ruficeps)

Taxonomy. Cavia capensis Pallas, 1766 ,

South Africa, Western Cape Prov., Cape of Good Hope.

Recent authors have recognized only a single species, P. capensis . However, some authorities consider P. capensis to be restricted to southern Africa and treat the following as distinct species: P. welwitschii, south-western Angola and Namibia; P. ruficeps, North and West Africa, and Sudan; P. johnston, south-western Tanzania, Malawi, Mozambique, and Zimbabwe; and P. habessinica, Egypt, Sudan, Israel, and Arabian Peninsula. Recent studies on geographic variation in mtDNA in South Africa indicate that at least two species exist in what has conventionally been regarded as P. capensis . Further genetic studies over the whole range will most likely divide Procavia into several species. Chromosome number is 2n = 54. The X chromosomeis the largest, with a submedian centromere; the Y chromosome is a very small acrocentric. No fewer than seventeen subspecies have been recognized and more than 65 synonyms are listed. The validity of many of these is doubtful, and some may actually represent distinct species.

Subspecies and Distribution.

P.c.capensisPallas,1766—SouthAfrica,includingLesothoandSwaziland.

P.c.bamendaeBrauer,1913—CameroonandCentralAfricanRepublic.

P.c.capillosaBrauer,1917—SEthiopia.

P.c.erlanger:Neumann,1901—SSomalia.

P.c.habessinicaHemprich&Ehrenberg,1832—Egypt,NSudan,Israel,SaudiArabia,andYemen.

P.c.jacksoniThomas,1900—EKenya.

P.c.jayakariThomas,1892—Oman.

P.c.johnston:Thomas,1894—SWTanzania,Malawi,Mozambique,andZimbabwe.

P.c.kerstingiMatschie,1899—TogoandBenin.

P.c.mackinder:Thomas,1900—WKenya.

P.c.matschietNeumann,1900—DRCongoandTanzania.

P.c.pallidaThomas,1891—NSomalia.

P.c.ruficepsHemprich&Ehrenberg,1832—NandWAfrica.

P.c.scioanaGiglioli,1888—NEthiopia.

P.c.sharicaThomas&Wroughton,1907—Chad.

P.c.syriacaSchreber,1784—Syria,Lebanon,Jordan,andIsrael.

P. c. welwitschii Gray, 1868 — SW Angola and Namibia. The distribution information for this species is still incomplete; the Rock Hyrax is also present in Eritrea, Niger, Nigeria, C & S Sudan, Uganda, Rwanda, Burundi, Zambia and E Botswana, but the subspecific identity of these populations still requires confirmation.

Descriptive notes. Head-body 39-58 cm; weight 1.8-5. 4 kg. Males and females are approximately the same size. The average adult size varies greatly across Africa, and seems to be closely linked to average annual precipitation, which in turn affects the availability of food; size increases up to a mean annual rainfall of 700 mm and decreases thereafter. On the other hand,size variation in the skull of Rock Hyraxes from different regionsis positively correlated with temperature, indicating that this species conforms to Bergmann’s rule. There is extensive variation in coat color, which varies widely throughout their range from a yellowish-buff to a dark brown; this variation has been associated with mean annual rainfall patterns. The pelage is dense, up to 25 mm long, and has a grizzled appearance due to banding of the hairs (dark at the base, with a paler band of varying width and a black tip). Underfuris short, soft, and thick. The underparts are paler in color than the upper, and the hair is slightly longer and lacks the banding. Long black vibrissae (tactile hairs) 60-70 mm in length (though longer on the face) are widely distributed over their bodies, probably for orientation in dark fissures and holes. Rock Hyraxes have a dorsal gland, surrounded by a creamy, yellowcolored (typical of ruficeps) or brown to black (typical of capensis ) margin of hairs that can be fanned when the animal is excited; this dorsal margin is not conspicuous in the Rock Hyrax of southern Africa. Females have three pairs of mammae, one pair pectoral and two pairs inguinal. Testes are permanently abdominal and the uterus is duplex. The penis is short, simply built with a slightly elliptical cross section, the diameter increasing slightly toward the tip. Mean distance between anus and penis is 35 mm. Characteristic features of the skull of the Rock Hyrax include: widely spaced and anteriorly situated eye sockets; well-developed interparietal; small tympanic bullae; the premaxillae form a tubercle between the incisive foramina; the coronoid process is small and recurved; and the hyoid bone is unusually scoop-shaped in structure. The zygomatic arches are broad and heavily built, indicative of a powerful set of masseter muscles that operate the lowerjaw. In contrast to the typically hyracoid dentition, dental formula is typically 11/2 C0/0 P 4/3 M 3/3 (x2)= 32, although specimens from northern parts of Africa often have the first lower premolar present (as in Heterohyrax brucei ). The length of the upper molar toothrow M'~ is much greater than that of the premolar toothrow P'**. The two upper, ever-growing incisor teeth, one on each side, are separated by about the width of one tooth. The upperincisors are tusk-like, ridged or triangular in cross-section in males, but rounded in females.

Habitat. Rock Hyraxes occupy a wide range of habitats, from arid deserts to rainforest, and from sea level to the alpine zone of Mount Kenya at 4000 m. However, as their name implies, Rock Hyraxes are dependent on the presence of rocky outcrops (kopjes), mountain cliffs, or loose boulders that provide suitable refuge in the form of crevices and crannies in which to shelter. The nature of the refuge environment differs substantially across Africa, but rock outcrops appear to be favored due to the extensive networks of crevices and fissures, access to safe foraging areas, and good vantage points. Overall the refuge environment provides stability compared with the surrounding habitats, where conditions are more extreme. However, Rock Hyraxes also have been found in erosion gullies (e.g. in the Karoo, a habitat they have colonized recently), in culverts under roads, holes in stone walls, and even in the holes of other species such as Aardvark (Orycteropus afer) and Meerkat (Suricata suricatta). These refuges seem to be common in areas where rocky habitats are overpopulated; Rock Hyraxes may traverse considerable distances between areas of suitable rocky habitat. In several parts of Africa (e.g. the Serengeti National Park in Tanzania, Matobo National Park in Zimbabwe, and the northern parts of South Africa), Rock Hyraxes and Bush Hyraxes ( Heterohyrax brucei ) occur together and live in close associations on rocky habitats. Rock Hyraxes are the most important prey in the diet of Verreaux’s eagle (Aquila verreauxit). Because males are forced to disperse when mature, oneto two-year-old males are particularly at risk of predation. Juveniles constituted 11-33% of Procavia remains in Verreaux's eagle nests in the Western Cape, and 18% of remains in the Matobo National Park, Zimbabwe. Rock Hyraxes form a main component of the diet of crowned hawk-eagles (Stephanoaetus coronatus). Other predators include martial (Polemaetus bellicosus) and tawny eagles (Aquila rapax), Leopards (Panthera pardus) particularly on Mount Kenya, Lions (Panthera leo), Caracals (Caracal caracal), jackals (Canis spp), Spotted Hyenas (Crocuta crocuta), and snakes. There is an extensive literature on the variety of external parasites—ticks, biting and sucking lice, mites, and fleas— that have been collected from Rock Hyraxes. In South Africa only ten of some 10,000 ticks recovered belonged to species that could infest domestic livestock. Dust-bathing probably helps keep parasite burdens relatively low. Rock Hyraxes harbor a number of internal parasites, including nematodes and cestodes, which could play a role in hyrax mortality in some areas. In Mountain Zebra National Park, South Africa, female Rock Hyraxes had the highest densities ofticks and biting lice in the summer. The difference between the sexes was related to the Rock Hyrax’s social structure. Females are more social than males making them vulnerable to increased infestation rates. Females also tend to be in worse physiological condition during the summer. No significant seasonal differences were noted in endoparasite densities. In the Serengeti, the sarcoptic mite, which causes mange, is an important cause of mortality for Rock Hyraxes, and females with symptoms of mange have been seen on Mount Kenya. In Kenya and Ethiopia, Rock Hyraxes might be an important reservoir for the parasitic diseases Leishmaniasis and Trypanosomiasis, as they have been reported to carry this parasite. Zoos have documented herpes virus infections in captive Rock Hyraxes.

Food and Feeding. The Rock Hyrax diet includes a variety of grasses, forbs, and shrubs, and they favor new shoots, buds, fruits, and berries. In the Serengeti, Rock Hyraxes were observed feeding on 79 plant species. The animals have a high seasonal adaptability. In the wet season they showed a preference for grasses (78%), but in the dry season when grasses became parched and poor in quality they browsed extensively (57%), more or less in proportion to the foliage density of each vegetation class. They were observed to feed on 24 grass species, including Panicum maximum, Pennisetum mezianum, and Themeda triandra; and on dicotyledonous plants such as Cordial ovalis, Maerua trphylla, Hoslundia opposita, Iboza sp., Hibiscus lunarifolius, Ficus ingens, Solanum incanum, Grewia fallax, and Acacia tortilis. In southern Africa, Rock Hyraxes exhibited similar seasonal preferences. In Namibia, a study of diet composition from two areas found that grasses were eaten in significant quantities only at the end of the hot-dry season and at the beginning of the wet season. Some 35 grass species were consumed, including Anthephora pubescens, Aristida congesta, Cynodon dactylon, Enneapogon scaber, E. brachystachyus, Eragrostis trichophora, Stipagrostis hirtigluma, and S. uniplumis. The important dicotyledonous plants in the diet were Acacia mellifera, and Ziziphus mucronata. Although they fed on a wide variety of plant species, ten species constituted more than 80% oftheir dietary biomass: Acacia karroo, Olea europaea, Felicia filifolia, Grewia occidentalis, Cussonia paniculata, Maytenus heterophylla, Pentzia spp., Clematis brachiata, Lycium oxycarpum, and Diospyros lycoides. Leaves of trees and shrubs formed the major portion of the diet; grazing largely depended on the seasonal availability of grasses. Examination of C":C'* ratios of carbonate and collagen fractions of bone and microwear patterns of the molariform teeth confirm that Rock Hyraxes switch between grazing and browsing at different times of the year. Rock Hyraxes in Kenya were observed feeding on a poisonous plant, Phytolacca dodecandra, and they are known to eat toxic Lobelia spp. However, there are a number of plant species that they avoid, such as Anthoxanthum nivale, Sedum ruwenzoriense, and Carduus keniensis. Most feeding occurs in groups. Rock Hyraxes often assume a fan-like orientation, which may serve to avoid conflict or to spot predators. Often a few individuals act as sentinels, as feeding away from their refuges makes the group vulnerable to attack by predators. Group feeding is intensive, usually only lasting about 20 minutes. Rock Hyraxes usually do not spend more than two hours per day feeding. Their disproportionally large jaws may be an adaptation that enables intensive feeding. In the Karoo, South Africa, the average daily distance travelled during group feeding ranged from 169 m to 572 m, but Rock Hyraxes seldom feed more than 15-20 m away from shelter. Feeding in groups, with sentinels,lets the hyraxes feed farther away from their crevices. Casual feeding by single individuals occurs more sporadically and at any time of the day, usually only a short distance from the protection of crevices.

Breeding. There is a single breeding season per year, and for both males and females, this represents a very short period of sexual activity. In South Africa male sexual activity is from February to May with a peak in April; testes mass increases dramatically during this time, with a more than tenfold difference between active and quiescent males. Females have a mean estrous cycle length of 13 days, but can cycle several times over a seven-week period. Gestation is 212-240 days, which is exceptionally long for such a small mammal. It may represent a primitive characteristic, given the much larger body sizes of some of the ancestral hyrax species, or it may reflect the adaptive advantages of producing precocial young. Within a family group, the pregnant femalesall give birth within a period of about three weeks;this birth synchrony appears to be a mechanism to avert predation by predators such as eagles. Some authors have suggested that parturition may be linked to rainfall or photoperiod. For example, in most areas in South Africa the breeding season is in late summer, with a peak in April, but there is a shift from earlier conception in the south-west (January—May) to later in the north-east (May-July). This results in a shift in births from the end of August in the south-west to the end of March in the north-east. In the Serengeti, births were recorded from March to May; births in Israel occurred in April, supporting photoperiod as one of the proximate causes. Timing of the breeding season reflects the adaptation to more favorable temperatures for the newborn young. In arid areas of north-western South Africa, conception is from September to November and parturition from May to July. The number of young per female varies between one and four (mean 2-4) for the Serengeti; in southern Africa, a female with six embryos was collected. Mean litter sizes recorded include: 3-3 in the Western Cape; 2-7 in the Eastern Cape; and 2: 37 in Zimbabwe. Some authors suggested an increase in littersize with latitude. Others demonstrated a relationship between litter size and female age. First breeders have only 1-2 young. Females between two and eight years produce the largest litters, followed by a decline in older females. Nutritional conditions probably affect both litter size and age at first breeding. The young are fully developed at birth, fully haired and with eyes open. They are capable of agile movement within a few hours and can ingest solid food within a few weeks. Birth weight varies to some degree depending on the number in the litter (mean 195 g). The average litter weight (581 g) is high relative to female body weight. Suckling young assume a strict teat order. Weaning occurs at 1-5 months. Both sexes reach sexual maturity at about 16-17 months. However, there is a report that some females breed at five months and males reach sexual maturity at 28-29 months in southern Africa. These findings suggest a difference between physiological and behavioral sexual maturity in males; the dominant male effectively prevents mating by sexually mature young males. The sex ratio is equal at birth and until two years of age, after which females sometimes tend to outnumber males. A high level ofjuvenile mortality appears to be an important factor controlling Rock Hyraxes population dynamics. At sexual maturity, females usually join the adult female group. Males disperse before they reach 30 months. Adult femaleslive significantly longer than adult males and may reach an age of over ten years; in captivity, animals have lived more than 14 years. Juvenile male dispersal appears to play a major role in the female-biased adult sex ratio. However, preliminary results of microsatellite DNA analysis of Rock Hyrax in the Serengeti show no sex bias and reveal extremely low levels of genetic variation within and between neighboring colonies. In South Africa, patterns of geographic variation of maternally inherited mtDNA suggest that there are high levels of historical connectedness among localities. Additional molecular markers are needed to reconcile direct and indirect estimates of dispersal and gene flow. Although occupancy of some habitat islandsis stable over time, most regions are characterized by fluctuations in Rock Hyrax numbers and may experience local extinctions and recolonizations. This is seen in colonies occupying kopijes in the Serengeti, where low levels of allelic diversity and heterozygosity at eight locations suggest metapopulation dynamics and a population bottleneck. The Rock Hyrax’s dynamic population structure may be a consequence of the species’ unpredictable environment.

Activity patterns. Rock Hyraxes are predominantly diurnal, though they occasionally are active on moonlit nights. It is not unusual to find them feeding at any time of the day, although there are peaks in mid-morning and mid-afternoon during warm periods. In winter, peak periods are later, or may become one extended period. More than 90% of the day is spent resting. Heaping, where several individuals are stacked on top of each other, is observed inside crevices and also outside during very cold conditions. Young can often be observed heaping with their mother. Although heaping can be observed inside crevices, huddling behavior is more common when groups are resting, especially early in the morning when the hyraxes first emerge from their crevices to sun themselves. During warmer times of day, solitary resting is seen. Hyrax social behavior is directly linked to daily temperature fluctuations and predation pressure. Different behavior patterns are expressed under varying environmental conditions. In southern Africa, in summer, Rock Hyraxes use rock crevices to avoid high temperatures. In the early morning, temperatures fall within the animals’ thermoneutral zones, which enables them to forage without the need to heat up. In winter, basking and huddling become essential for their survival.

Movements, Home range and Social organization. Rock Hyraxes are gregarious,living in cohesive and stable family groups or colonies numbering as many as 80 individuals and consisting of 3-7 related adult females, one adult territorial male, dispersing males, subadult females, and juveniles of both sexes. Their numbers vary depending on the size of the kopje. Smaller kopjes or rocky outcrops support only a single colony, but larger kopjes may support several family groups, each occupying a traditional range. There are four classes of mature male: territorial, peripheral, and early and late dispersers. Territorial males are the most dominant, and repel all intruding males from an area largely encompassing the females’ core area. Their aggressive behavior towards other adult males escalates particularly in the mating season, when they monopolize all receptive females. On small kopjes, peripheral males are unable to settle, but on large kopjes they can occupy areas on the periphery ofthe territorial males’ territories. Males are solitary, and the highest ranking among them takes over a female group whenever a territorial male disappears; some studies have reported regular replacement of the dominant male. The females’ home ranges are not defended and may overlap. Rarely, an adult female from outside a group will be incorporated into the family group. The majority ofjuvenile males—the early dispersers—leave their birth sites at 16-24 months old, soon after reaching sexual maturity. The late dispersers leave a year later, but before they are 30 months old. Individual Rock Hyraxes have been observed to disperse over a distance of at least 2 km, although gene flow over distances greater than 10 km is unlikely. The farther a dispersing animal has to travel across open country, where there is little cover and few hiding places, the greater its chances of dying, either through predation or as a result ofits inability to cope with temperature stress. Observers in southern Africa reported shorter dispersal distances, indicating that the rocky habitat and the intervening vegetation between habitat islands strongly influence dispersal distance. Some of the resting time is spent self-grooming, using the lower incisors and the curved claw on the second digit. Grooming and dust bathing help rid Rock Hyraxes of ectoparasites. Rock Hyraxes urinate and defecate in latrines; over time, calcium carbonate in their urine crystallizes, forming deposits that whiten the cliff faces below latrines. Although Rock Hyraxes are gregarious, low levels of intraspecific aggression play an important role in maintaining colonial life. Visual communications include flaring the hairs surrounding the dorsal gland and appeasement behavior. Pilo-erection of the dorsal spot can either signal alarm (if the hairs are erected at a 45° angle) or threat (90°). These clear signals and stereotypical appeasement behavior limit serious aggressive encounters between individuals; agonistic behavior is mostly observed between males during the breeding season. Olfactory communication functions during reproduction and to establish mother—infant bonds. In southern Africa 21 vocal and four non-vocal sounds were recorded. The multitude of grunts, growls, snarls, spits, snorts, and squeals are used in a variety of contexts, but most commonly in showing aggression, appeasement, or defensive retreat. The alarm call in the form of a sharp bark is characteristic and differs from that of the Bush Hyrax and tree hyraxes. The alarm calls used by the sentinels, especially during group feeding, appear to be specific to the particular threat. A repetitious bark or song appears to function in transmitting territorial and sexual signals. Observations in Israel have shown that these songs provide accurate information regarding body weight, size and condition, social status, and hormonal state of the caller. Resident males and some bachelor males sing complex songs in individually distinct voices throughout most of the year, and also countersing with neighboring males. Singing males differ from the general adult male population in that their cortisol levels are higher than those of silent males. In singers, cortisol levels are associated with social rank, with dominants showing the highest levels. Singers are also on average older and more dominant, and they copulate more often than non-singers, suggesting that singing males may have higher reproductive success. Males are more vocal during the breeding season and also use dorsal gland secretions for signalling. These displays may be composed of components that are important in male-male competition, as well as mate choice. The dominantterritorial males monitor urine deposits routinely in search of receptive females. Most copulations are between the territorial males and adult females; peripheral males most often mate with subadult females. However, females mate with more than one territorial male and with peripheral males. Receptive females approach the dominant male and signal their readiness to mate by flaring their dorsal spot hairs, sniffing the male’s anogenital region, and presenting their hindquarters. Males initiate mating through a mating call, weaving head movements, and dorsal spot flaring. Copulations are brief. The young are born inside the rock crevices. Juveniles form nursery groups that often engage in social play; both juveniles and subadults have a much larger area of activity compared with the territorial and peripheral adults. In the Rock Hyrax, a strong correlation was found in females, but not in males, between androgens and cortisol. In most mammal species, the male shows this association. Female and male Rock Hyraxes have similar testosterone levels, and a significant relationship between social status and testosterone levels was observed. These levels differed only between lower-ranking males and females, with lower-ranking females showing higher levels. Dominant females had a significantly lower testosterone level than subordinate females. No association was detected between litter size and a female’s rank, testosterone, or cortisol levels.

Status and Conservation. Currently classified as Least Concern on The IUCN Red List. Although this species is subject to some localized hunting, it is widely distributed on the African continent, is present in a number of protected areas across its range, and is generally not believed to be at any risk of extinction in the wild. However, more and better information on its status and distribution is desperately needed. Rock Hyrax populations in South Africa’s Cape Province were once listed as vermin due to their high numbers and grazing impact. However, about ten years ago populations in the KwaZulu-Natal province became locally extinct. Two recent reintroduction attempts of captive and wild groups in a reserve failed.

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Figures

1. Rock Hyrax (Procavia capensis), 2. Bush Hyrax (Heterohyrax brucei), 3. Western Tree Hyrax (Dendrohyrax dorsalis), 4. Southern Tree Hyrax (Dendrohyrax arboreus), 5. Eastern Tree Hyrax (Dendrohyrax validus)

Subspecies and Distribution. P.c.capensisPallas,1766—SouthAfrica,includingLesothoandSwaziland. P.c.bamendaeBrauer,1913—CameroonandCentralAfricanRepublic. P.c.capillosaBrauer,1917—SEthiopia. P.c.erlanger:Neumann,1901—SSomalia. P.c.habessinicaHemprich&Ehrenberg,1832—Egypt,NSudan,Israel,SaudiArabia,andYemen. P.c.jacksoniThomas,1900—EKenya. P.c.jayakariThomas,1892—Oman. P.c.johnston:Thomas,1894—SWTanzania,Malawi,Mozambique,andZimbabwe. P.c.kerstingiMatschie,1899—TogoandBenin. P.c.mackinder:Thomas,1900—WKenya. P.c.matschietNeumann,1900—DRCongoandTanzania. P.c.pallidaThomas,1891—NSomalia. P.c.ruficepsHemprich&Ehrenberg,1832—NandWAfrica. P.c.scioanaGiglioli,1888—NEthiopia. P.c.sharicaThomas&Wroughton,1907—Chad. P.c.syriacaSchreber,1784—Syria,Lebanon,Jordan,andIsrael. P. c. welwitschii Gray, 1868 — SW Angola and Namibia . The distribution information for this species is still incomplete; the Rock Hyrax is also present in Eritrea , Niger , Nigeria , C & S Sudan , Uganda , Rwanda , Burundi , Zambia and E Botswana , but the subspecific identity of these populations still requires confirmation.