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Hyrax

Hyraxes (order Hyracoidea) are small to medium-sized, herbivorous mammals in the family Procaviidae, native primarily to sub-Saharan Africa and parts of the Middle East, with five extant species divided among three genera: the rock hyrax (Procavia capensis), the tree hyraxes (genus Dendrohyrax, including D. arboreus, D. validus, and D. dorsalis), and the bush hyrax (Heterohyrax brucei).^[1] Despite their rodent- or lagomorph-like appearance, hyraxes are ungulates with hoof-like nails on their toes and are the closest living relatives to elephants (Proboscidea) and sirenians (manatees and dugongs) within the placental mammal superorder Afrotheria.^[2]^[3] These mammals typically weigh 2–5 kg (4–11 lb) and measure 30–56 cm (12–22 in) in length, featuring rounded heads, short tails, and specialized feet with moist, rubbery pads that enable strong adhesion to rocks and trees.^[4] Rock hyraxes inhabit rocky outcrops and cliffs in arid and semi-arid regions, where they form diurnal colonies of 5–80 individuals centered around a dominant male, while tree hyraxes are mostly nocturnal and solitary or live in small family groups in forested areas, and bush hyraxes occupy bushy, rocky savannas.^[4]^[5] All species are primarily folivorous, browsing on leaves, fruits, and grasses, with some tolerance for toxic plants due to specialized gut bacteria.^[4]^[6] Hyrax societies exhibit complex social structures, including communal latrines, vocal communication (such as loud alarm calls in rock hyraxes), and alloparenting behaviors, with reproduction generally seasonal and litters of 1–4 precocial young after gestations of 6–8 months.^[4]^[7] Predators include leopards, eagles, snakes, and caracals, prompting hyraxes to rely on sentinel systems and rapid escapes into crevices.^[8] Conservation statuses vary: the rock and bush hyraxes are classified as Least Concern by the IUCN due to their wide distributions and stable populations, while the eastern tree hyrax is Near Threatened due to habitat loss and population declines, and the other tree hyrax species are Least Concern but decreasing in some areas.^[9]

Taxonomy and evolution

Classification

Hyraxes are classified in the kingdom Animalia, phylum Chordata, class Mammalia, superorder Afrotheria, order Hyracoidea, and family Procaviidae, the sole extant family in the order.^[10]^[11] The name of the order Hyracoidea derives from the Greek hyrakios, meaning "shrew-mouse," reflecting early perceptions of their small, rodent-like appearance.^[11] Within the family Procaviidae, there are three extant genera: Procavia (rock hyraxes), Heterohyrax (bush hyraxes), and Dendrohyrax (tree hyraxes).^[11]^[9] Current taxonomy recognizes five extant species: Procavia capensis (Cape rock hyrax), Heterohyrax brucei (yellow-spotted bush hyrax), Dendrohyrax arboreus (southern tree hyrax), Dendrohyrax dorsalis (western tree hyrax), and Dendrohyrax validus (eastern tree hyrax). Taxonomic revisions in the early 21st century reduced the number of recognized species from around 11 to these five, reclassifying many as subspecies; over 50 subspecies are described across the genera.^[12]^[13] Hyraxes form a close evolutionary relationship with elephants and sirenians in the clade Paenungulata.^[10]

Phylogenetic relationships

Hyraxes (order Hyracoidea) were historically misclassified by morphologists as rodents or ungulates due to superficial resemblances such as their small size, rodent-like appearance, and herbivorous dentition, a view prevalent from the late 19th century until molecular analyses in the late 20th century.^[14] This traditional placement emphasized convergent adaptations for terrestrial herbivory rather than deep evolutionary affinities, leading to their grouping with groups like the "condylarths" or other early ungulate-like mammals. Cladistic analyses incorporating both morphological and molecular data later corrected this, revealing hyraxes as part of a distinct African radiation.^[15] Molecular phylogenetic studies from the 1990s onward firmly established hyraxes within the superorder Afrotheria, a clade encompassing six orders: Hyracoidea (hyraxes), Proboscidea (elephants), Sirenia (sirenians), Tubulidentata (aardvarks), Macroscelidea (elephant shrews or sengis), and Afrosoricida (tenrecs and golden moles).^[16] This monophyly is supported by extensive genomic evidence, including nuclear and mitochondrial DNA sequences, retroposon insertions, and chromosomal data, which demonstrate shared ancestry originating in Africa during the Cretaceous.^[17] Within Afrotheria, hyraxes form the order Hyracoidea in the subclade Paenungulata alongside Proboscidea and Sirenia, with Afrosoricida and Macroscelidea as more distant relatives; this topology has been consistently recovered across multiple datasets, resolving earlier uncertainties in interordinal relationships.^[15] Hyraxes share several derived morphological traits with proboscideans and sirenians that support their paenungulate affinity, including testicondy (internal testes retained in the abdominal cavity), high-crowned lophodont molars adapted for grinding vegetation, and hoof-like nails on the digits rather than claws.^[15]^[18] These synapomorphies, combined with molecular markers, indicate a common ancestor for Paenungulata that diverged around 63-65 million years ago in the early Paleocene, shortly after the Cretaceous-Paleogene boundary, marking a rapid radiation within Afrotheria.^[19] This timeline aligns with fossil-calibrated molecular clocks, emphasizing the role of Africa's isolation in fostering these unique evolutionary linkages.^[16]

Fossil record

The fossil record of hyraxes (Hyracoidea) originates in the late Eocene, approximately 37 million years ago, in Africa, where early forms such as primitive hyracoids from the Priabonian stage of Egypt exhibited small, rodent-like morphologies with generalized dental features suited to browsing.^[20] These initial representatives, including genera like Microhyrax from slightly earlier Eocene deposits in North Africa, weighed around 3 kg and displayed arboreal adaptations such as a mobile midtarsal joint in the tarsals.^[14] This emergence aligns with the broader afrotherian radiation following the Cretaceous-Paleogene extinction event, as evidenced by the diverse Paleogene mammal assemblages in North African sites, marking the diversification of endemic African placentals into herbivorous niches.^[21] Hyrax diversity peaked during the Oligocene and Miocene epochs, encompassing over 30 genera across Africa and into Eurasia, reflecting a wide range of body sizes and ecological roles from small browsers to large herbivores.^[22] Notable examples include Titanohyrax from the Eocene-Oligocene boundary, which achieved elephant-like proportions with estimated body masses up to 1,300 kg, based on cranial and postcranial remains from North African localities. Similarly, Megalohyrax eocaenus from the early Oligocene Fayum Depression in Egypt reached pig- to tapir-sized dimensions, with a skull length of 391 mm indicating a body mass of several hundred kilograms and lophodont molars adapted for grinding vegetation.^[23] Key fossil sites contributing to this record include the Fayum Depression in Egypt, renowned for its rich late Eocene to early Oligocene vertebrate assemblages that document the early phases of afrotherian evolution, and the East African Rift, such as the Rukwa Rift Basin in Tanzania, where late Oligocene forms like Rukwalorax jinokitana reveal sub-equatorial diversification.^[24]^[25] By the late Miocene, hyrax lineages began a marked decline, attributed to intensified competition from emerging bovid artiodactyls, which were more efficient grazers and browsers, coupled with climatic shifts toward drier environments that favored open savannas over forested habitats.^[26] Fossil evidence from East African sites shows a reduction in hyrax abundance and variety during this period, with large-bodied forms like those in the Pliohyracidae becoming rare.^[22] Most ancient hyracoid lineages had gone extinct by the Pliocene, leaving only the modern procaviids as survivors, a pattern underscored by sparse late Miocene records in regions like Greece and the absence of diverse forms thereafter.^[22]

Physical description

Morphology

Hyraxes are small, rotund mammals with a stocky build that superficially resembles rodents such as marmots or groundhogs, featuring short tails, rounded ears, and a compact body form.^[27] They typically measure 30 to 70 cm in body length and weigh between 2 and 5 kg, with variations across species; for instance, the rock hyrax (Procavia capensis) averages 30 to 50 cm long and 3.5 to 5 kg.^[28]^[29] Their fur is coarse and sparsely distributed, ranging in color from yellowish-brown to gray, which provides camouflage in rocky environments.^[27] Some species, including the rock hyrax, possess a dorsal glandular spot—a bare patch on the midline of the back covered by longer, often darker hairs—that secretes an oily substance for scent marking.^[29]^[28] Hyrax dentition includes two upper incisors and four lower ones that grow continuously throughout life, similar to rodent teeth, though the upper pair in males develops into prominent tusk-like structures.^[27] The total number of teeth is 32 to 34, with a dental formula of 1/2, 0/0, 4/4, 3/3; the molars are high-crowned and equipped with transverse enamel ridges suited for grinding vegetation.^[27]^[30] This incisor morphology shares evolutionary parallels with that of elephants, underscoring their afrotherian affinities.^[27] Skeletally, hyraxes have short limbs adapted for a plantigrade stance, with the forefeet bearing four toes and the hind feet three, all tipped with broad nails rather than hooves.^[27] They retain clavicles, a feature uncommon among many ungulate-like mammals, contributing to their agile climbing ability.^[27] In the wild, hyraxes have a lifespan of up to 12 years.^[4]

Adaptations

Hyraxes exhibit several specialized physiological and anatomical adaptations that facilitate their survival in rocky and arboreal environments across sub-Saharan Africa and the Middle East. Due to their small body size, typically ranging from 2 to 5 kg, hyraxes possess limited internal thermoregulatory capacity, relying heavily on behavioral mechanisms to maintain body temperature. They bask in the sun during cooler periods to elevate their core temperature and huddle in groups during cold nights to conserve heat, compensating for their inability to precisely regulate internal heat production. Body temperature in rock hyraxes (Procavia capensis) fluctuates by up to 4.5°C over a 24-hour cycle, reflecting this dependence on external cues rather than metabolic adjustments.^[31]^[32] The digestive system of hyraxes is adapted for processing fibrous vegetation through a combination of foregut and hindgut fermentation, enabling efficient nutrient extraction from low-quality forage. The stomach features a sacculation that serves as the primary foregut fermentation chamber, where microbial activity initiates the breakdown of plant material, producing volatile fatty acids similar to those in ruminants, though the structure is not fully multichambered. This is complemented by a large caecum in the hindgut, which acts as a secondary fermentation site, hosting microbes that degrade cellulose and hemicellulose in fibrous plants; during dry seasons when food quality declines, the caecum can enlarge to up to 4.6% of body mass to enhance this process. These dual fermentation sites allow hyraxes to derive significant energy from tough, lignified diets without extensive chewing.^[33] Locomotion in hyraxes is facilitated by specialized foot structures that provide strong adhesion and grip on vertical rock faces and tree trunks. The soles of their feet are covered in rubbery, moist pads composed of soft epidermal tissue, maintained wet by eccrine sweat glands that secrete a viscous fluid to enhance friction and prevent slipping on smooth surfaces. These pads incorporate unique hexagonal epidermal patterns and microscopic ridges that increase surface contact and tensile adhesion, akin to mechanisms in geckos but reliant on moisture rather than setae. Hind feet typically feature three digits, with the two medial digits bearing hoof-like nails for weight-bearing stability, while the lateral digits and a central claw on the middle digit aid in gripping and grooming; front feet have four digits with similar adaptations. These features enable agile climbing despite the absence of a tail for balance.^[34]^[35] Sensory adaptations in hyraxes emphasize vigilance against predators in open habitats. They possess excellent eyesight, with eyes positioned for a wide field of view and an iris that protrudes slightly over the pupil to reduce glare from overhead light. Hearing is also acute, supporting the use of complex vocalizations for long-distance communication; male rock hyrax songs include low-frequency elements that propagate effectively over rocky terrain to convey territorial and social information.^[36]^[37]

Distribution and habitat

Geographic range

Hyraxes are native to sub-Saharan Africa and the Arabian Peninsula, with no established populations outside these regions. The rock hyrax (Procavia capensis) exhibits the broadest distribution among the species, spanning from Senegal and Mauritania in West Africa, through northern countries including Algeria, Libya, and Egypt, southward across the continent to South Africa, and extending into the Middle East from Syria and Jordan to Yemen and Oman.^[38]^[28] Tree hyraxes of the genus Dendrohyrax are restricted to forested equatorial Africa. The western tree hyrax (D. dorsalis) ranges from Guinea and Liberia across West and Central Africa to Uganda and Tanzania, while the southern tree hyrax (D. arboreus) occupies eastern and southern regions from the Democratic Republic of the Congo through Kenya, Tanzania, and Zambia to Mozambique, Angola, and South Africa; the eastern tree hyrax (D. validus) is distributed patchily in a narrow band of lowland and montane forests in Kenya and Tanzania.^[39] The Benin tree hyrax (D. interfluvialis) is found in wet and dry forests between the Niger and Volta Rivers in Ghana, Togo, Benin, and Nigeria.^[40]^[41]^[42] The bush hyrax (Heterohyrax brucei) is primarily found in eastern and central Africa, from Sudan, Eritrea, and Ethiopia southward to Angola, Zambia, Malawi, and South Africa, with extensions to the Sinai Peninsula and Djibouti.^[43]^[28] During the Pleistocene, hyrax distributions were more extensive, encompassing broader areas of North Africa, but subsequent aridification linked to glacial cycles led to habitat fragmentation and local extirpations in some regions.^[44]

Habitat preferences

Hyraxes exhibit distinct habitat preferences shaped by their need for shelter, foraging opportunities, and predator avoidance, with species specializing in rocky, arboreal, or mixed environments across Africa and the Middle East.^[45] Rock hyraxes (Procavia capensis) primarily inhabit rocky outcrops, kopjes, cliffs, and boulder screes, utilizing cavities and crevices within igneous, metamorphic, or sedimentary rock formations for shelter and evasion from predators like leopards.^[46] They avoid open plains, favoring sheltered entrances to holes that protect against wind and larger intruders, with microhabitats often featuring disused burrows of other animals or even human-made structures like road culverts.^[46] These populations thrive in proximity to vegetation for foraging, spanning altitudinal ranges from sea level to over 4,200 meters in areas like the Ethiopian highlands and Mount Kenya.^[46] While not strictly dependent on free-standing water sources, rock hyraxes rely on moisture from nearby vegetation, enabling survival in arid to semi-arid zones.^[45] Tree hyraxes (Dendrohyrax spp.), in contrast, favor tropical and montane forests with dense, multilayered canopies and tall trees exceeding 50 meters, where woody climbers and lianas facilitate arboreal movement, feeding, and daytime refuge in hollows or branches.^[47] Viable populations require large forest fragments over 90 hectares to support their shy, nocturnal lifestyle, though smaller patches near larger forests may sustain limited groups; they retreat to caves or rocky outcrops in degraded areas.^[47]^[12] Altitudinal distribution includes moist montane elevations up to 2,200 meters in the Taita Hills of Kenya and higher in ranges like the Ruwenzori Mountains, with preferences for intact natural forests providing foliage and protection.^[47]^[12] Like other hyraxes, they derive hydration from plant material, avoiding direct dependence on surface water.^[45] Bush hyraxes (Heterohyrax brucei), also known as yellow-spotted hyraxes, occupy woodland savannas and bushy areas, blending semi-arboreal habits with terrestrial shelter in rocky kopjes, krantzes, and boulder piles that offer crevices for hiding.^[48] They climb low trees and vertical trunks to browse, preferring habitats with moderate temperatures (17–25°C) and humidity (32–40%) near woody vegetation like Acacia and Combretum species.^[48] This species ranges from sea level to 3,800 meters in East Africa, sometimes utilizing fig tree holes near rivers for additional refuge, while obtaining all necessary water from their diet of leaves, buds, and fruits.^[48]

Behavior and ecology

Diet and feeding

Hyraxes are primarily folivorous herbivores, with diets consisting mainly of leaves, grasses, fruits, and bark from a variety of plant species.^[49] They exhibit opportunistic feeding, browsing on woody plants and grazing on available vegetation, and can tolerate certain toxic plants that are poisonous to other mammals, such as those in the Solanaceae and Euphorbiaceae families.^[4] Rock hyraxes (Procavia capensis) in arid regions particularly favor succulents and lichens growing on rocks, which provide moisture and nutrients in water-scarce environments.^[50] Foraging typically occurs during daylight hours, with peaks shortly after dawn and before dusk to minimize exposure to predators and heat.^[51] Individuals spend approximately 3-4 hours per day feeding, selectively browsing to maximize nutrient intake while avoiding high-toxin concentrations in plants.^[8] Hyraxes possess specialized dentition adapted for processing fibrous plant material, with high-crowned molars featuring transverse ridges that shear tough vegetation during mastication.^[52] Unlike rodents, they crop vegetation primarily using the sides of the premolars and molars by turning their heads sideways, rather than relying on incisors for initial clipping.^[53] These dental features, combined with hindgut fermentation, support their herbivorous lifestyle, though detailed digestive processes are addressed in adaptations.^[54] Dietary composition varies seasonally, with a shift toward seeds, fruits, and bark during dry periods when fresh leaves are scarce, allowing hyraxes to exploit available resources efficiently. They derive most of their water from food sources, rarely drinking free water even when available, which is crucial for survival in arid habitats.^[55] Hyraxes exhibit varied social structures depending on the species, with rock hyraxes (Procavia capensis) forming larger, more complex groups compared to the generally solitary tree hyraxes (Dendrohyrax spp.). Rock hyraxes live in stable colonies typically consisting of 9 to 32 individuals, including a dominant territorial male, 3 to 17 adult and subadult females that form the core female-bonded unit, and juveniles and subadults of both sexes, often with one or more peripheral males loosely associated.^[56] These groups display a matriarchal structure among females, who maintain low levels of aggression and egalitarian social networks, while males exhibit linear dominance hierarchies.^[56]^[57] In contrast, tree hyraxes are primarily solitary or live in small family units of 2 to 3 individuals, such as a mother with subadult young, though occasional groups of up to 9 have been observed in specific habitats.^[47]^[12] Communication among hyraxes serves to coordinate group activities, maintain bonds, and alert others to potential dangers. Rock hyraxes employ a variety of vocalizations, including alarm calls such as squeals, barks, chucks, and snorts, which convey information about predator proximity and type.^[58] They also use scent marking via the prominent dorsal gland, a specialized sebaceous gland on the back that secretes odorous substances for territorial demarcation and social signaling, often combined with visual displays like gland erection.^[58]^[59] A sentinel system enhances group vigilance, where individual rock hyraxes, often from elevated positions, monitor for threats during foraging and emit alarm calls to warn the group, allowing rapid retreat to rocky refuges.^[60]^[61] Tree hyraxes rely heavily on nocturnal vocalizations for communication, including alarm calls like snorts, hacs, and wheezes with frequencies ranging from 220 to 15,000 Hz, though scent marking details are less documented beyond communal middens.^[12] Social hierarchies in rock hyraxes are maintained through territorial defense by dominant males, who monopolize access to females and core areas, displacing subadults and peripheral males.^[56] Females, as the stable social core, engage in affiliative behaviors like allogrooming to strengthen bonds and reduce tension, with grooming networks predicting social support and longevity in egalitarian groups.^[62]^[57] Leadership roles are context-dependent and sex-specific: younger males often lead daily movements under moderate risk, while less central individuals may initiate responses during threats, and females share leadership more equally.^[63] Activity patterns align with habitat and predation pressures, influencing social interactions. Rock hyraxes are diurnal, with groups foraging and basking together during the day before retreating to shelters at night, though some active nighttime proximity maintains bonds.^[64] Juveniles engage in play behaviors, such as chasing, mock mating, fur nipping, and climbing, observed frequently among subadults to develop social skills.^[56] Tree hyraxes, adapted to arboreal life, are nocturnal, with peak activity from 23:00 to 04:00, limiting overt social displays but enabling vocal coordination in small units.^[12]

Reproduction and life cycle

Hyraxes exhibit breeding seasons that typically occur one or two times per year, often synchronized with rainfall patterns to ensure food availability for offspring, though this varies by species and location; for instance, rock hyraxes (Procavia capensis) in arid regions may breed in spring following winter rains.^[8] Mating systems are polygynous, with dominant males aggressively chasing and herding receptive females within social groups to secure multiple mates during the season.^[4] Gestation periods last approximately seven to eight months across hyrax species, an unusually long duration for their body size, resulting in litters of one to four precocial young that are born fully furred with open eyes and capable of limited mobility shortly after birth.^[4] Tree hyraxes (Dendrohyrax spp.) generally produce smaller litters of one to two offspring, while rock hyraxes can have up to four, with an average of two to three.^[7]^[65] Young hyraxes are weaned between one and three months of age, after which they begin foraging independently, though they may remain in the natal group for protection.^[59] Sexual maturity is reached at around 16 to 17 months for females and slightly later, up to 24 months, for males, with full physical growth attained by about three years.^[4]^[66] In the wild, hyraxes have a maximum lifespan of up to 12 years, with averages around 5-9 years influenced by high juvenile mortality rates—often exceeding 50% due to predation by eagles, leopards, and snakes; in captivity, individuals can survive up to 14-17 years with reduced threats.^[67]^[45]^[68] Social group dynamics, including communal vigilance, aid in mitigating some predation risks during early development.^[59]

Conservation

Status and threats

The conservation status of hyrax species varies, with most classified as Least Concern by the IUCN Red List due to their wide distribution and adaptability. For instance, the rock hyrax (Procavia capensis), the most widespread species, is assessed as Least Concern with a stable population trend, reflecting its abundance in rocky habitats across sub-Saharan Africa and parts of the Middle East.^[69] In contrast, certain tree hyrax species face greater risks; the eastern tree hyrax (Dendrohyrax validus) is categorized as Near Threatened, with a decreasing population driven by ongoing habitat fragmentation and exploitation.^[39] Overall, hyrax populations remain stable on a continental scale but are increasingly fragmented, as isolated habitat patches limit gene flow and increase vulnerability to local extirpations. The primary anthropogenic threats to hyraxes include habitat loss and degradation from agricultural expansion and deforestation, which have reduced available rocky outcrops and forested areas essential for shelter and foraging, particularly in eastern and western Africa. Hunting for bushmeat and skins further exacerbates pressures, leading to localized population reductions, especially in areas with high human-wildlife overlap.^[69] ^[39] Natural predation by species such as leopards, martial and Verreaux's eagles, and various snakes also influences population dynamics, though these are balanced in undisturbed ecosystems.^[59] Common hyrax species, like the rock hyrax, are estimated to number in the millions across their range, supported by densities of 20–100 individuals per km² in suitable habitats such as Mount Kenya.^[69] However, localized declines have been reported in East Africa, including Ethiopia's Zegie Peninsula and Tanzania's forests, where habitat loss and hunting have led to reported population declines, with 55% of local residents noting decreases in some surveyed areas.^[70] These declines highlight the need for targeted monitoring in fragmented regions.

Protection efforts

Hyrax populations benefit from inclusion in several key protected areas across their range, such as Serengeti National Park in Tanzania, where studies indicate higher densities near human-inhabited sites due to reduced predation, and Kruger National Park in South Africa, which supports ongoing wildlife monitoring and habitat preservation.^[71]^[72] In Ethiopia, the Bale Mountains National Park safeguards endemic subspecies like the rock hyrax Procavia capensis capillosa, contributing to regional conservation through habitat protection and population assessments.^[73] These sites collectively encompass significant portions of suitable hyrax habitats, aiding in the maintenance of stable populations for least concern species. Legally, hyraxes are not included in the CITES appendices, though Sudan proposed listing the rock hyrax (Procavia capensis) under Appendix III in 2019 to regulate international trade, reflecting national protections already in place there.^[74] In South Africa, regulations vary by province; for instance, in Limpopo, rock hyraxes are classified as protected wild animals requiring permits for hunting, while in KwaZulu-Natal, no permit is needed during open seasons.^[75]^[76] Israel's rock hyrax population holds protected status under wildlife laws, though temporary derogations for culling have been authorized since 2018 to manage leishmaniasis transmission.^[77] The IUCN Species Survival Commission (SSC) Afrotheria Specialist Group leads research and monitoring efforts, developing standardized protocols for population tracking across afrotherian species, including hyraxes, with initiatives funded since 2016 to conduct range-wide surveys and genetic analyses.^[78]^[79] Camera trap surveys, increasingly employed in the 2010s for mammal inventories in African protected areas, have captured hyrax occurrences alongside other species, providing data on distribution and relative abundance in regions like the Dja Biosphere Reserve in Cameroon.^[80] Reintroduction efforts for hyraxes remain limited and mixed in success. In South Africa, two attempts to reintroduce rock hyraxes into a KwaZulu-Natal reserve failed due to predation and habitat challenges.^[81] In Namibia, releases of rehabilitated rock hyraxes from sanctuaries, such as those by Naankuse Wildlife Sanctuary, support population supplementation in communal conservancies, where ecotourism generates community benefits and incentivizes habitat conservation.^[82]^[83]

Cultural and human significance

Historical references

In the Hebrew Bible, the hyrax is identified as the "shaphan," an animal deemed unclean in Leviticus 11:5 and Deuteronomy 14:7 because it chews the cud yet lacks cloven hooves.^[84] This description reflects an observation of the hyrax's feeding behavior, which mimics rumination through reingestion of partially digested food, though it is not a true ruminant.^[85] In various African cultures, hyraxes hold symbolic significance in folklore. For instance, in some regions of southern Africa, rock hyraxes are regarded as symbols of fertility and rain due to their plump bodies and ability to thrive in arid environments. Additionally, hyraceum—a sticky mass of dung and urine produced by hyraxes—has been used in traditional South African folk medicine to treat ailments such as epilepsy and convulsions, and in perfumery as a musk substitute.^[86] Ancient depictions of hyraxes appear in San rock art in South Africa, dating back approximately 4,000 years, where they are portrayed in hunting scenes by these hunter-gatherer communities.^[87] A notable example is found at Modderpoort in the Free State, featuring one of South Africa's few unequivocal representations of the rock hyrax (Procavia capensis).^[88] In ancient Egypt, hyraxes are referenced in textual and artistic records, including hieroglyphs that denote the animal, with evidence indicating their role as a supplementary food source alongside vegetation like acacia leaves and pods.^[89] The term "hyrax" originates from the ancient Hebrew shaphan, meaning "the hidden one," alluding to the animal's elusive, rock-dwelling nature, a usage shared in Phoenician nomenclature. A longstanding but disputed etymological theory links the name "Spain" (from Latin Hispania) to the Phoenician phrase I-Shpania or I-Shaphan, interpreted as "land of hyraxes" or "coast of hyraxes," stemming from Phoenician explorers' misidentification of abundant Iberian rabbits as hyraxes.^[90]

Modern interactions

Rock hyraxes have been maintained in zoos and as occasional pets since at least the mid-20th century, with early records of captive breeding emerging in the 1930s in European and African institutions.^[91] However, keeping them presents significant challenges, including nutritional deficiencies that lead to metabolic bone disease from imbalanced calcium-phosphorus ratios in captive diets, as well as iron overload potentially exacerbated by high-iron feeds and genetic factors.^[92]^[93] Stress from confinement and handling further complicates care, often resulting in elevated cortisol levels and reduced welfare, as observed in reintroduction attempts where captives exhibited disease susceptibility. Breeding programs, such as those at Israel's Tisch Family Zoological Park (Jerusalem Biblical Zoo), have successfully propagated rock hyraxes while integrating research on their social behaviors, contributing to both ex situ conservation and public education.^[94] In ecotourism, hyraxes attract visitors to natural reserves and urban-adjacent sites, enhancing local economies through wildlife viewing opportunities. In South Africa, where rock hyraxes are common in protected areas, ecotourism centered on observing these animals alongside other species generates substantial revenue, supporting rural jobs and habitat preservation, though exact figures vary by region and year. Guided hikes on Table Mountain National Park, for instance, frequently highlight dassies (rock hyraxes) basking on rocks or foraging, providing educational encounters that draw thousands of hikers annually and promote biodiversity awareness.^[95]^[96] Hyraxes serve as valuable models in scientific research, particularly for elucidating afrotherian evolution, as molecular and fossil evidence positions them as close relatives to elephants and sirenians within this ancient placental mammal clade.^[10]^[17] Non-invasive studies of their vocalizations, conducted via audio recordings in wild populations, reveal complex song structures used in mating and territorial displays, offering insights into mammalian communication without disturbing natural behaviors.^[97]^[98] Hyraxes have gained prominence in modern media, with tree hyrax "singing" calls inspiring viral social media trends in 2024, where their eerie, melodic vocalizations were memed as "awawa" sounds on platforms like TikTok and Instagram, amassing millions of views and highlighting their quirky appeal. They also feature in documentaries, such as BBC Earth's segments in the "Africa" series and standalone clips narrated by David Attenborough, which showcase their survival strategies and evolutionary ties to elephants, educating global audiences on lesser-known wildlife.^[99]^[100]

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Molecular phylogeneticists have identified an Afrotheria clade, which includes several taxa as different as tenrecs (Tenrecidae), golden moles (Chrysochloridae) ...
[15]
A new estimate of afrotherian phylogeny based on simultaneous ...
Nov 13, 2007 · The placental mammalian clade Afrotheria is now supported by diverse forms of genomic data, but interordinal relationships within, and ...
[16]
Afrotheria: Plate tectonics meets genomics - PNAS
Afrotheria is one of the most remarkable hypotheses in mammal evolution. It suggests that one-third of the orders of placental mammals form an ancient group ...
[17]
A Retroposon Analysis of Afrotherian Phylogeny - Oxford Academic
Abstract. Recent comprehensive studies of DNA sequences support the monophyly of Afrotheria, comprising elephants, sirenians (dugongs and manatees), hyraxe.
[18]
https://www.sciencedirect.com/science/article/pii/S0022519398908219
[19]
Placental mammal diversification and the Cretaceous–Tertiary ...
Among interordinal splits, only divergences within Paenungulata occurred after the K/T boundary, and only when constrained with a maximum of 65 mya. Within ...
[20]
(PDF) A primitive hyracoid (Mammalia, Paenungulata) from the early ...
Aug 7, 2025 · and Geniohyus and is absent in Antilohyrax, Saghatherium,. Selenohyrax, Thyrohyrax and Titanohyrax.
[21]
[PDF] The oldest and youngest records of afrosoricid placentals from the ...
2010. The oldest and youngest records of afrosoricid placentals from the Fayum Depression of northern. Egypt. Acta Palaeontologica Polonica 55 (4): 599–616.
[22]
The Fossil Record of Hyraxes (Mammalia: Hyracoidea - ResearchGate
They appeared at the Eocene but later, during the Late Eocene–Oligocene, diversified with several taxa. Their number decreased at the beginning of the Miocene ...
[23]
Skull of Megalohyrax eocaenus (Hyracoidea, Mammalia) from the ...
Aug 10, 2025 · The reference point for Paleogene hyracoid diversity has long been the Eocene and Oligocene localities of the Fayum Depression in Egypt ...Missing: peak | Show results with:peak
[24]
New Age Determinations for the Eocene-Oligocene Boundary ...
The Fayum Depression of northern Egypt contains the richest collection of fossil vertebrates known from the Afro-Arabian Paleogene.
[25]
(PDF) A Hyracoid from the Late Oligocene Red Sandstone Group of ...
Aug 10, 2025 · Late Eocene to early Oligocene sites of the Fayum in Egypt are particularly famous for their rich hyracoid fossil record illustrating the ...
[26]
Revision of the Early Miocene Hyracoidea (Mammalia) of East Africa
Despite differences in cranial and dental morphology from Megalohyrax and Bunohyrax, Whitworth [18] classified two species in these genera that are recorded ...
[27]
Hyracoidea (hyraxes) | INFORMATION - Animal Diversity Web
There is a single family of hyraxes, Procaviidae, and seven living species. Hyraxes are found in sub-Saharan Africa and Middle East. Technical characters.Missing: list taxonomy<|control11|><|separator|>
[28]
Hyraxes - IUCN Afrotheria Specialist Group
Size Variation in the Rock Hyrax (Procavia capensis) and Late Quartenary climatic change in South Africa. Quaternary Research 46, 193-207. Olds, N. and J.Missing: skeletal | Show results with:skeletal
[29]
Rock Hyrax - San Diego Zoo Animals & Plants
The rock hyrax is covered in short brown fur, with a lighter underbelly. There are extra-long hairs that stick out around the body called guard hairs to help ...About · Habitat And Diet · Family Life
[30]
Hyrocoidea (Hyraxes) - Veterian Key
Aug 27, 2016 · The deciduous dental formula for the hyrax is: incisors (I) 2/2, canines (C) 1/1, premolars (P) 4/4, for a total of 28 teeth. The permanent ...
[31]
Rock hyrax (Procavia capensis) longevity, ageing, and life history
In the wild, animals have been reported to live for 8.5 years [0434]. One captive specimen lived for 14.8 years [0671]. Life history traits (averages).
[32]
[PDF] SEASONAL VARIATION IN THE THERMAL BIOLOGY OF THE ...
Our results show that rock hyrax seasonally and daily altered both their physiological and behavioural thermoregulation to control body temperature efficiently.
[33]
regulation of body temperature in the rock hyrax, heterohyrax brucei
REGULATION OF BODY TEMPERATURE IN THE ROCK HYRAX,. HETEROHYRAX BRUCEI. GEORGE A. BARTHOLOMEW AND MICHAEL RAINY. ABSTRACT.-The effects of ambient temperature on ...
[34]
[PDF] THE DIGESTION IN THE HYRAX PROCAVIA CAPENSIS
JIGESTIVE SYSTEM. The digestive system of the hyrax is schematically shown in Fig. 2. The oesophagus is a muscular tube 10 cm long. The stomach is 12 cm long.
[35]
Adaptive evolution of secretory cell lines in vertebrate skin
well as adhesion to tissue substrates (KocH et al. 1991). When the hagfish ... glands of the foot pads of the rock hyrax (Procavia capensis, Hyracoidea ...
[36]
Fingerprint ridges allow primates to regulate grip - PNAS
Nov 30, 2020 · Sweating improves grip as demonstrated, for example, by measuring the sliding resistance of the footpads of rats, tenrecs, rock hyrax, and ...
[37]
Review of sensory modalities of sirenians and the other extant ...
Aug 23, 2021 · ... hyraxes is the largest in absolute size amongst Paenungulata (Ekdale, 2013). Elephants and sirenians share a low position of their lateral ...
[38]
Male hyraxes increase song complexity and duration in the ...
Aug 25, 2014 · Long-range vocalizations were recorded from a distance of 10–50 m with a Sennheiser ME 67 shotgun microphone (frequency response 50–20000 Hz ± ...
[39]
Procavia capensis (rock hyrax) - Animal Diversity Web
These hyraxes are heavily built for their size, weighing as much as 4.3 kg. Males (4 kg) are slightly larger than females (3.6 kg) and have blunter features ...Missing: anatomy dentition skeletal
[40]
Procavia capensis • Rock Hyrax - ASM Mammal Diversity Database
South Africa, Western Cape Prov., Cape of Good Hope. Biogeographic realm. Palearctic · Afrotropic. Country distribution. Syria · Lebanon · Israel · Palestine · ...
[41]
Dendrohyrax dorsalis • Western Tree Hyrax
Description of a new species of Hyrax from Fernando Po. Proceedings of the ... IUCN Red List status. Least Concern. Species Permalink. https://www ...<|separator|>
[42]
Dendrohyrax arboreus • Southern Tree Hyrax
IUCN Red List status. Least Concern. Species Permalink. https://www ... Dendrohyrax arboreus arboreus. Shortridge, 1934. Family. Procaviidae. Root name. arboreus ...
[43]
Heterohyrax brucei • Bush Hyrax - ASM Mammal Diversity Database
IUCN Red List status. Least Concern. Species Permalink. https://www.mammaldiversity.org/taxon/1000527. Country distribution map. The map below provides a ...
[44]
[PDF] Multilocus phylogeography of rock hyrax (Procavia capensis) from ...
Repeated climatic fluctuations and aridification caused by Pleistocene global glacial cycles have caused periodic habitat fragmentation, resulting in isolation, ...
[45]
Hyrax | African Wildlife Foundation
Life span. Up to 12 years in the wild; average 8.5 years. Habitat. Dry savanna to dense rainforest to rocky outcrops. Diet. Herbivorous. Gestation. 7 to 8 ...
[46]
[PDF] The Habitat of the Rock Hyrax - Biodiversity Heritage Library
Rock hyrax live in cavities within rocky outcrops, often in boulder heaps at the foot of cliffs, and in volcanic areas. They are found in the Rift Valley and ...Missing: preferences | Show results with:preferences
[47]
Habitat preferences, estimated abundance and behavior of tree ...
Apr 15, 2022 · Calling frequency of tree hyraxes provides useful data that can be used for estimating relative population abundance28. Generally, acoustic ...
[48]
(PDF) Heterohyrax brucei - ResearchGate
Aug 9, 2025 · Hyrax population size is determined by kopje size and number of crevices from which to hide from predators (Hoeck, 1989).
[49]
Procavia - an overview | ScienceDirect Topics
Procavia refers to a genus of small herbivorous mammals known as hyraxes, with the rock hyrax (Procavia capensis) being a representative species found in ...Missing: fur skeletal
[50]
(PDF) The Biology of Hyraxes - ResearchGate
Jan 2, 2016 · The dentition of the hyrax is unusual as only the second incisors are retained in the upper jaw. These continually growing teeth are triangular ...Missing: testicondy | Show results with:testicondy
[51]
Patch use and site selection by a rock hyrax (Procavia capensis) in a ...
... foraging and resource renewal by plant growth. beginning shortly after dawn and again for 1 h before dusk As rock hyraxes are suspected to be the reservoir ...
[52]
Mammalian dental function and wear: A review - ScienceDirect.com
Wet season rock hyrax teeth had more microwear striations than did those of bush hyraxes, or those of rock hyraxes collected during the dry season when less ...
[53]
(PDF) The hyrax - a most mysterious mammal - ResearchGate
Hyraxes are small herbivorous mammals which occupy a wide range of habitats from coastal rocks, alpine moorland, and dry savannah to dense rainforest in Africa ...Missing: classification source
[54]
Teeth and the gastrointestinal tract in mammals: when 1 + 1 = 3 - NIH
Harvesting of microbes occurs either via coprophagy after separation from indigestible material by a separation mechanism in the hindgut, or from a forestomach ...Missing: recycling | Show results with:recycling
[55]
Stable isotope analyses of rock hyrax faecal pellets, hyraceum and ...
Rock hyraxes are generally independent of water (i.e. drought-tolerant), but when available, they will drink freely (Skinner and Chimimba, 2005). In South ...
[56]
Social behaviour and spatial relationships of the rock hyrax
Hyraxes associate to form colonies consisting of a dominant territorial male, several adult females and sub- adults and juveniles of both sexes; ...
[57]
Variance in Centrality within Rock Hyrax Social Networks Predicts ...
Jul 27, 2011 · Several recent studies have demonstrated that the social integration of adult females is positively associated with infant survival, and female ...
[58]
[PDF] Complex call in male rock hyrax (Procavia capensis)
Dec 24, 2008 · microphone (frequency response 50–20,000 Hz ±2.5 dB) powered by a Sennheiser K6 module, covered with a. Sennheiser MZW70-1 blimp windscreen ...<|control11|><|separator|>
[59]
Using path analysis to explore vigilance behavior in the rock hyrax ...
Feb 16, 2011 · During these group foraging bouts, occasionally 1 hyrax will act as a sentinel, watching for predators from a lookout (Kotler et al. 1999). ...
[60]
High-resolution tracking of hyrax social interactions highlights ...
Dec 15, 2022 · This pattern likely helps hyraxes to maintain long-term social bonds while adapting to short-term changes in their physical environment.
[61]
[PDF] Strong social relationships are associated with decreased longevity ...
Jan 17, 2018 · We focused on affiliative interactions [27,28] that included cohesive behaviours such as allogrooming, greeting, foraging together, playing.
[62]
Sex-associated and context-dependent leadership in the rock hyrax
Apr 15, 2022 · The findings demonstrated that male and female rock hyraxes take on different leadership positions, depending on the context.
[63]
High-resolution tracking of hyrax social interactions highlights ...
Dec 15, 2022 · We investigated the nighttime sociality of a wild population of rock hyraxes (Procavia capensis) and its relation to daytime social structure.Missing: taxonomy | Show results with:taxonomy
[64]
Rock Hyrax - Center for Perinatal Discovery
There are three hyrax genera: Procavia, Heterohyrax, and Dendrohyrax. Varied opinions have been expressed regarding their validity. The three genera, however, ...
[65]
Sex-associated and context-dependent leadership in the rock hyrax
Apr 15, 2022 · Native males reach maturity at the age of 17–24 months, and soon after gradually leave their natal group or are occasionally forced to ...
[66]
Rock Hyrax Facts and Information | United Parks & Resorts
rock hyrax, hyrax, dassie; Kingdom: Animalia; Phylum: Chordata; Class: Mammalia ... Status: IUCN: Not listed. CITES: Not listed. USFWS: Not listed. Fun Facts.<|separator|>
[67]
The IUCN Red List of Threatened Species
### Summary for Procavia capensis
[68]
The IUCN Red List of Threatened Species
### Summary for Dendrohyrax validus
[69]
Population size, habitat association, and local residents' attitude ...
We conducted this study to determine the population size and examine local residents' attitudes toward the rock hyrax in the Zegie Peninsula, Ethiopia.
[70]
(PDF) Do hyraxes benefit from human presence in Serengeti?
This study focuses on the factors that affect hyrax population size, including factors such as human presence, habitat type, kopje size and kopje height. We ...
[71]
Protecting Kruger's Legacy: Conservation Efforts | Africa Moja Tours
Oct 27, 2024 · The research conducted in Kruger informs not only local strategies but also global conservation efforts. Studies from the park help scientists ...
[72]
Population size, habitat association, and local residents' attitude ...
Jun 30, 2025 · Five extant species of hyraxes are recognized: the rock hyrax (Procavia capensis) and the yellow-spotted rock hyrax (Heterohyrax brucei), which ...
[73]
[PDF] Head quarter of police forces - WILDLIFE CONSERVATION - CITES
3/ import the Rock hyrax (Procavia capensis) in the appendix III of the CITES agreement, as it includes in the National Law of Sudan in the appendix III and ...
[74]
Is Rock Hyrax a legal game animal? | AfricaHunting.com
Apr 3, 2013 · In Limpopo South Africa just across the river they are a protected wild animal and need a permit to hunt one . The first thing you should look ...SOUTH AFRICA Hunting InformationBarrel length for handguns in South Africa?More results from www.africahunting.com
[75]
[PDF] HUNTING AND PERMITS IN KZN. WHAT YOU NEED TO KNOW
Bush pig, Rock Hyrax (Dassie),. Chacma Baboon, Vervet Monkey,. BB Jackal, Black and Pied Crow. • No Closed Season. • No Permit or License required. • Hunting ...
[76]
Hyraxes May Lose Protected Status in Israel to Prevent Spread of ...
Aug 26, 2018 · Rock hyraxes in Israel may lose protected status for three years in an attempt to curb the spread of leishmaniasis, a parasitic infection they host that causes ...
[77]
[PDF] IUCN SSC Afrotheria Specialist Group
Research activities: (1) develop five stan- dardised monitoring protocols for each group of Afrotherians to track trends over time and produce more data for ...
[78]
None
### Hyrax Conservation Efforts Summary
[79]
[PDF] Camera-Trap Survey for Larger Terrestrial Wildlife in the Dja ... - ZSL
This report summarises camera-trap results from the first systematic camera-trap survey conducted in the Northern Sector of the Dja Biosphere Reserve (DBR).
[80]
Two unsuccessful reintroduction attempts of rock hyraxes (Procavia ...
Two unsuccessful reintroduction attempts of rock hyraxes (Procavia capensis) into a reserve in the KwaZulu-Natal Province, South Africa.Missing: Israel Namibia ecotourism
[81]
Rock hyrax release. Some of our sanctuary dassies not only recently ...
May 2, 2020 · Rock hyrax release. Some of our sanctuary dassies not only recently received their health checks, but also took their steps to freedom.
[82]
Inspiring responsible travel to Namibia's communal conservancies
Aug 8, 2020 · The economic impact of joint venture tourism and the value it adds to keeping wildlife on the land cannot be underestimated, particularly in ...<|separator|>
[83]
https://conservationnamibia.com/articles/cnam2020-conservation-tourism.php
[84]
Coney - Meaning & Verses | Bible Encyclopedia
Neither the hyrax nor the hare is a ruminant, as seems to be implied in Leviticus 11:5 and Deuteronomy 14:7, but their manner of chewing their food may readily ...
[85]
San Rock Art of the Drakensberg | Nature - PBS
Mar 2, 2011 · They are the San, hunter-gatherers who are believed to have lived in this area for 4,000 years, as far back as the Stone Age. Though the San ...Missing: hyrax | Show results with:hyrax
[86]
San Rock Art in Free State
This site is truly unique in that it has South Africa's only unequivocal example of a dassie or rock hyrax (Procaviacapensis). Modderpoort. Bushman rock ...
[87]
The Mammals of Ancient Egypt on JSTOR
Hyraxes are diurnal and nocturnal, but feed mainly at night. Acacia leaves and pods are s taple food,... Save.
[88]
Rabbits, fish and mice, but no rock hyrax
Nov 9, 2014 · Apparently the Phoenicians named the Iberian peninsula I-Shaphan (Hispania ) meaning "coast of hyraxes ", a misidentification on the part of ...Missing: Shafen | Show results with:Shafen
[89]
Rock hyrax - Comparative Placentation
Feb 4, 2008 · The name "hyrax" originates from the Greek "hurax", for mouse. Despite their similarity to rodents, they form a distinct, separate Order of ...Missing: etymology | Show results with:etymology
[90]
[PDF] Rock Hyrax: Diet Recommendations and Nutritional Pathology
The normal body temperatures of rock hyrax only fluctuate plus or minus (+/) 2º C. (3.6º F) and fluctuations greater or less than 2º C (3.6º F) result in cold ...<|control11|><|separator|>
[91]
Hyrax - an overview | ScienceDirect Topics
Modern hyraxes are classified in a single family with three genera and four species. Limited to Africa, fossil hyraxes are also found in the Middle East and ...<|control11|><|separator|>
[92]
The hyrax that sings best will have the healthiest pups - ISRAEL21c
Oct 19, 2022 · Study of the mating song rhythm and structure of rock badgers suggests that virtuosos are chosen because they'll father the most surviving ...
[93]
Private Hiking Tour of Table Mountain - Cape Town - Viator
Rating 5.0 (36) A nature guide will accompany you on both routes, pointing out Table Mountain's flora and fauna—including adorable rock hyrax—en route. Private guided hike up ...
[94]
What Animals and Plants Live on Table Mountain? - AWOL Tours
Jul 28, 2025 · In this guide, we'll explore the animals and plants you might encounter on Table Mountain, from rare frogs and rock hyraxes to proteas and ...
[95]
Social status and cortisol levels in singing rock hyraxes
In this study, we examine whether singers, which are more conspicuous by the act of singing, have higher cortisol (i.e. basal stress; C) levels than non-singers ...
[96]
Acoustic Communication in the Rock Hyrax, Procavia capensis
We propose a method for non-invasive estimation of occupancy of the burrows ... These low-frequency vocalizations, also found in other non-related ...
[97]
Meet the Hyrax: The Adorable New Animal Taking Over Social Media
Nov 13, 2024 · The hyrax is a rock-dwelling animal with an unforgettable scream that's taken over social media with funny memes.
[98]
Meet the Endangered Hyrax | Little Brother of the Elephant | BBC Earth
Jul 3, 2009 · Sir David Attenborough narrates this BBC footage of the endangered hyrax species. Visit http://www.bbcearth.com for all the latest animal ...