Hyrax

Hyraxes
Temporal range: Eocene-Recent, 55.8–0 Ma
Yellow-spotted rock hyrax (Heterohyrax brucei)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Class: Mammalia
Clade: Atlantogenata
Superorder: Afrotheria
Clade: Paenungulata
Order: Hyracoidea
Huxley, 1869
Families

Hyraxes (from the Greek ὕραξ, hurax, "shrewmouse"), also called dassies,[1] are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between 30 and 70 cm (12 and 28 in) long and weigh between 2 and 5 kg (4.4 and 11 lb). They are superficially similar to pikas or rodents (especially marmots), but are more closely related to elephants and manatees.

Four extant species are recognised; the rock hyrax (Procavia capensis), the yellow-spotted rock hyrax (Heterohyrax brucei), the western tree hyrax (Dendrohyrax dorsalis) and the southern tree hyrax (D. arboreus). Their distribution is limited to Africa and the Middle East.

Characteristics

Hyraxes retain a number of primitive mammalian characteristics; in particular, they have poorly developed internal temperature regulation, for which they compensate by behavioural thermoregulation, such as huddling together and basking in the sun.

Unlike most other browsing and grazing animals, they do not use the incisors at the front of the jaw for slicing off leaves and grass; rather, they use the molar teeth at the side of the jaw. The two upper incisors are large and tusk-like, and grow continuously through life, similar to rodents. The four lower incisors are deeply grooved 'comb teeth'. A diastema occurs between the incisors and the cheek teeth. The dental formula for hyraxes is 1.0.4.32.0.4.3.

Although not ruminants, hyraxes have complex, multichambered stomachs that allow symbiotic bacteria to break down tough plant materials, but their overall ability to digest fibre is lower than that of the ungulates.[2] Their mandibular motions (see video) are deceptively similar to chewing cud,[3] but the hyrax is physically incapable of regurgitation[4][5] as in the even-toed ungulates and the merycism of some of the macropods. This behaviour is referred to in a passage in the Bible which describes hyraxes as chewing the cud.[6] This chewing behaviour may be a form of agonistic behaviour when the animal feels threatened.[7]

Hyraxes inhabit rocky terrain across sub-Saharan Africa and the Middle East. Their feet have rubbery pads with numerous sweat glands, which may help the animal maintain its grip when quickly moving up steep, rocky surfaces. Hyraxes have stumpy toes with hoof-like nails; there are four toes on each front foot and three on each back foot.[8] They also have efficient kidneys, retaining water so that they can better survive in arid environments.

Female hyraxes give birth to up to four young after a gestation period between seven and eight months, depending on the species. The young are weaned at one to five months of age, and reach sexual maturity at 16 to 17 months.

Hyraxes live in small family groups, with a single male that aggressively defends the territory from rivals. Where living space is abundant, the male may have sole access to multiple groups of females, each with their own range. The remaining males live solitary lives, often on the periphery of areas controlled by larger males, and mate only with younger females.[9]

Hyraxes have highly charged myoglobin, which has been inferred to reflect an aquatic ancestry.[10]

Similarities with elephants and Sirenia

Hyraxes share several unusual characteristics with elephants and the Sirenia (manatees and dugongs), which have resulted in their all being placed in the taxon Paenungulata. Male hyraxes lack a scrotum and their testicles remain tucked up in their abdominal cavity next to the kidneys,[11][12] the same as elephants, manatees, and dugongs.[13] Female hyraxes have a pair of teats near their arm pits (axilla), as well as four teats in their groin (inguinal area); elephants have a pair of teats near their axillae, and dugongs and manatees have a pair of teats, one located close to each of the front flippers. The tusks of hyraxes develop from the incisor teeth as do the tusks of elephants; most mammalian tusks develop from the canines. Hyraxes, like elephants, have flattened nails on the tips of their digits, rather than curved, elongated claws which are usually seen on mammals.[14]

Historical accounts

Young hyrax on Mount Kenya

The words "rabbit", "hare", or "coney" appear as terms for the hyrax in some English translations of the Bible. Early English translators had no knowledge of the hyrax (Hebrew שָּׁפָן shafan),[15] and therefore no name for them. References are made to hyraxes in the Old Testament,[16] particularly in Leviticus 11, where they are described as lacking a split hoof and therefore not being kosher. It also describes the hyrax as chewing its cud; this is technically inaccurate as the hyrax does not regurgitate its food to chew it again. The Hebrew phrase in question (מַעֲלֵה גֵרָה) means "bringing up cud". Some of the modern translations refer to them as rock badgers.

Evolution

Pachyhyrax championi, a large fossil hyrax from the Miocene of Rusinga, Kenya (Natural History Museum collection)

All modern hyraxes are members of the family Procaviidae (the only living family within Hyracoidea) and are found only in Africa and the Middle East. In the past, however, hyraxes were more diverse, and widespread. The order first appears in the fossil record at a site in the Middle East in the form of Dimaitherium, 37 million years ago.[17] For many millions of years, hyraxes were the primary terrestrial herbivores in Africa, just as odd-toed ungulates were in North America.

Through the middle to late Eocene, many different species existed,[18] the largest of them weighing the same as a small horse and the smallest the size of a mouse. During the Miocene, however, competition from the newly developed bovids, which were very efficient grazers and browsers, displaced the hyraxes into marginal niches. Nevertheless, the order remained widespread and diverse as late as the end of the Pliocene (about two million years ago) with representatives throughout most of Africa, Europe, and Asia.

The descendants of the giant 'hyracoids' (common ancestors to the hyraxes, elephants, and sirenians) evolved in different ways. Some became smaller, and evolved to become the modern hyrax family. Others appear to have taken to the water (perhaps like the modern capybara), ultimately giving rise to the elephant family and perhaps also the sirenians. DNA evidence supports this hypothesis, and the small modern hyraxes share numerous features with elephants, such as toenails, excellent hearing, sensitive pads on their feet, small tusks, good memory, higher brain functions compared to other similar mammals, and the shape of some of their bones.[19]

Hyraxes are sometimes described as being the closest living relative of the elephant,[20] although whether this is so is disputed. Recent morphological- and molecular-based classifications reveal the sirenians to be the closest living relatives of elephants. While hyraxes are closely related, they form a taxonomic outgroup to the assemblage of elephants, sirenians, and the extinct orders Embrithopoda and Desmostylia.[21]

The extinct meridiungulate family Archaeohyracidae, consisting of four genera of notoungulate mammals known from the Paleocene through the Oligocene of South America,[22] is a group unrelated to the true hyraxes.

List of genera

Phylogeny of early hyracoids


Hyracoidea


Seggeurius


Microhyrax

Saghatheriinae


Bunohyrax


Pachyhyrax


Thyrohyrax




Selenohyrax

Saghatherium


Titanohyrax

Antilohyrax

Megalohyrax






Geniohyiinae

Geniohyus

Proboscidea

Perissodactyla[lower-alpha 1]

Phenacodontidae



A phylogeny of hyracoids known from the early Eocene through the middle Oligocene epoch. Modern day hyraxes (Procaviidae) may have evolved from small members that are here considered Saghatheriinae.[23]
  1. ↑ The relationship of hyracoids and Perissodactlya is controversial, and not supported by molecular data.

Extant species

In the 2000s, taxonomists reduced the number of recognized species of hyrax. In 1995, 11 or more species were recognized; in 2013, only four are recognized, with the others now each considered as a subspecies of one of the recognized four. Over 50 subspecies and species are described, many of which are considered highly endangered.[24]

See also

References

  1. ↑ "Hyracoidea" in Grzimek's Animal Life Encyclopedia, Vol. 15: Mammals. Gale Publishing. Online version accessed April 2014.
  2. ↑ von Engelhardt, W; Wolter, S; Lawrenz, H; Hemsley, J.A (1978). "Production of methane in two non-ruminant herbivores". Comparative Biochemistry and Physiology A. 60 (3): 309–11. doi:10.1016/0300-9629(78)90254-2.
  3. ↑ Hendrichs, Hubert (1966). "Vergleichende Untersuchung des Wiederkauverhaltens" [Comparative investigation of cud retainers]. Biologisches Zentralblatt (dissertation) (in German). 84 (6): 671–751. OCLC 251821046. All artiodactyl families and about 80% of the spp. were investigated. Chewing regurgitated fodder is an idle pastime as well as an instinct associated with appetite. Characteristic movements were analyzed for undisturbed samples of animals maintained on preserves. Group specific differences are reported in form, rhythm, frequency and side of chewing motion. The ungulate type is characterized as a specialization. The operation is described for the first time for the order Hyracoidea. On the basis of 12 spp. of the marsupial subfamily Macropodinae rumination is inferred for the whole category. Advantages of the process are debated
  4. ↑ Björnhag G, Becker G, Buchholz C, von Engelhardt W (1994). "The gastrointestinal tract of the rock hyrax (Procavia habessinica). 1. Morphology and motility patterns of the tract". Comparative Biochemistry and Physiology A. 109 (3): 649–53. PMID 8529006. doi:10.1016/0300-9629(94)90205-4.
  5. ↑ Sale, J. B. (1966). DAILY FOOD CONSUMPTION AND MODE OF INGESTION IN THE HYRAX. JE Afr.Nat.Hist.Soc., Vol XXV(No. 3), Page 219. Retrieved from http://www.biodiversitylibrary.org/content/part/EANHS/XXV_No.3_112__215_1966_Sale.pdf
  6. ↑ "Leviticus 11:5". Bible Gateway. Zondervan. Retrieved 2016-04-08.
  7. ↑ "Archived copy" (PDF). Archived from the original (PDF) on 16 June 2012. Retrieved 25 April 2012.
  8. ↑ http://www.awf.org/wildlife-conservation/hyrax%5B%5D
  9. ↑ Hoeck, Hendrik (1984). Macdonald, D., ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 462–5. ISBN 0-87196-871-1.
  10. ↑ "One Protein Shows Elephants and Moles Had Aquatic Ancestors".
  11. ↑ Trevor Carnaby (1 January 2008). Beat about the Bush: Mammals. Jacana Media. p. 293. ISBN 978-1-77009-240-2.
  12. ↑ Septimus Sisson (1914). The anatomy of the domestic animals. W.B. Saunders Company. p. 577.
  13. ↑ Marshall Cavendish Corporation (1 September 2010). Mammal Anatomy: An Illustrated Guide. Marshall Cavendish. p. 63. ISBN 978-0-7614-7882-9.
  14. ↑ Picture of hyrax feet
  15. ↑ "Shaphan" in Strong's Concordance
  16. ↑ Lev 11:4-8; Deut 14:7; Ps 104:18; Prov 30:26
  17. ↑ Eugenie Barrow; Erik R. Seiffert; Elwyn L. Simons. (2010). "A primitive hyracoid (Mammalia, Paenungulata) from the early Priabonian (Late Eocene) of Egypt". Journal of Systematic Palaeontology. 8 (2): 213–244. doi:10.1080/14772010903450407.
  18. ↑ Prothero, Donald R. (2006). After the Dinosaurs: The Age of Mammals. Bloomington, Indiana: Indiana University Press. p. 132. ISBN 978-0-253-34733-6.
  19. ↑ "Hyrax: The Little Brother of the Elephant", Wildlife on One, BBC TV.
  20. ↑ "Hirax song is a menu for mating". The Economist. 15 January 2009. Retrieved 15 January 2009.
  21. ↑ Asher, R.J., Novacek, M.J., Geisher, J.H. (2003). "Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence". J. Mamm. Evol. 10: 131–194. doi:10.1023/A:1025504124129.
  22. ↑ McKenna, Malcolm C., and Bell, Susan K. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, 631 pp. ISBN 0-231-11013-8
  23. ↑ Gheerbrant, E; D. Donming; P. Tassy (2005). "Paenungulata (Sirenia, Proboscidea, Hyracoidea, and Relatives)". In Kenneth D. Rose, J. David Archibald (eds.). The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades. Baltimore: Johns Hopkins University Press. pp. 84–105. ISBN 080188022X.
  24. ↑ Shoshani, J. (2005). Wilson, D.E.; Reeder, D.M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 87–89. ISBN 978-0-8018-8221-0. OCLC 62265494.

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