Difference between revisions of "Hyrax" - New World Encyclopedia

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{{Taxobox
 
{{Taxobox
 
| name = Hyraxes
 
| name = Hyraxes
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| subdivision_ranks = [[Family (biology)|Families]]
 
| subdivision_ranks = [[Family (biology)|Families]]
 
| subdivision =
 
| subdivision =
* †Pliohyracidae
 
 
* '''Procaviidae'''
 
* '''Procaviidae'''
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* [[extinct|†]]Pliohyracidae
 
}}
 
}}
  
'''Hyrax''' is the common name for any of the small, largely herbivorous [[mammal]]s comprising the [[Order (biology)|order]] '''Hyracoidea''' and characterized by thickset bodies, short tails, and three-toed hind feet with rounded, hoof-like nails on two of the toes and a claw on the third. These mammals, which can be described as having a [[guinea pig]] or [[woodchuck]]-like body, and are similar in size to a large [[rabbit]], are also known as '''cories''' and '''dassies'''. The presence of specialized muscles in the soles of the feet, combined with the soft and elastic nature of the [[sweat-gland]] moistened soles, helps to create a suction-like friction with rocks and other substrate.  Hyraxes are often mistaken for [[rodent]]s, but are more closely related to [[elephant]]s. There is one extant family of hyraxes, Procaviidae, distributed only in [[Africa]] and the [[Middle East]].
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'''Hyrax''' is the common name for any of the largely herbivorous [[mammal]]s comprising the [[Order (biology)|order]] '''Hyracoidea''', whose modern members are characterized by thickset bodies, short tails, and three-toed hind feet with rounded, hoof-like nails on two of the toes and a claw on the third. These mammals, which can be described as having a [[guinea pig]] or [[woodchuck]]-like body, and are similar in size to a large [[rabbit]], are also known as '''cories''' and '''dassies'''. The presence of specialized muscles in the soles of the feet, combined with the soft and elastic nature of the [[sweat-gland]] moistened soles, helps to create a suction-like friction with rocks and other substrate, making them agile climbers.   
  
 +
There is one extant family of hyraxes, Procaviidae, distributed only in [[Africa]] and the [[Middle East]]. Four species are recognized: the [[rock hyrax]] (''Procavia capensis''), the [[yellow-spotted rock hyrax|yellow-spotted rock hyrax]] (''Heterohyrax brucei''), the [[western tree hyrax|western tree hyrax]] (''Dendrohyrax dorsalis'') and the [[southern tree hyrax|southern tree hyrax]] (''Dendrohyrax arboreus''). However, in evolutionary history hyraxes were more diverse, and with larger and smaller representatives, and for many millions of years they were the primary terrestrial herbivore in Africa. Hyraxes are often mistaken for [[rodent]]s, but are more closely related to [[elephant]]s.
  
values food chain and to humans
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Hyraxes provide diverse values for the [[ecosystem]] and for humans. As part of [[food chain]]s, they consume many tree, shrub, and grass plant parts (leaves, petioles, twigs, shoots, fruits, seeds) and occasionally insects, while being consumed by birds of prey ([[eagle]]s, [[owl]]s), large cats (lions, leopards), jackals, [[snake]]s, and [[mongoose]]s. Hyraxes form the almost exclusive diet of Verreaux eagles. Hyraxes also disperse the seeds of fruits they eat and play a role in nutrient cycling. For humans, hyraxes have been hunted for food and [[fur]] and their crystallized excrement has been used by Europeans and African tribes to treat various illness (epilepsy, hysteria, convulsions) and as vitamin supplements.
 
 
 
 
 
 
 
 
Four species are recognised; the [[rock hyrax]] (''Procavia capensis''), the [[Yellow-spotted Rock Hyrax|yellow-spotted rock hyrax]] (''Heterohyrax brucei''), the [[Western Tree Hyrax|western tree hyrax]] (''Dendrohyrax dorsalis'') and the [[Southern Tree Hyrax|southern tree hyrax]] (''Dendrohyrax arboreus'').
 
  
 
==Physical description==
 
==Physical description==
  
Hyraxes are well-furred, shot-tailed, rotund animals about the size of a large [[rabbit]]. Typically, they measure between 30 and 70 centimeters (12-28 inches) long and weigh between 2 and 5 kilograms (4-11 pounds).   
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Hyraxes are well-furred, short-tailed, rotund animals about the size of a large [[rabbit]]. Typically, they measure between 30 and 70 centimeters (12-28 inches) long and weigh between 2 and 5 kilograms (4-11 pounds).   
  
Hyraxes have 3-toed hind feet with two of the toes with rounded nails that resemble hooves, while the inner toe has a claw (Myers 2000). Forefeet generally have four or five toes. The soles of their fore- and hindfeet are soft, elastic, rubbery like pads, moistened by numerous specialized sweat glands, which increases friction and help the animal maintain its grip when quickly moving up steep, rocky surfaces. The soles of the feet also have specialized muscles in this regard, which helps the soles to work somewhat like suction cups. The plane of symmetry of the feet goes through the third digit (Myers 2000).
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[[File:hyraxfootcloseup.jpg|thumb|The characteristic foot pads]]
 
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Hyraxes have 3-toed hind feet with two of the toes with rounded nails that resemble hooves, while the inner toe has a claw. Forefeet have four or five toes. The soles of the fore- and hindfeet are soft, elastic, rubbery like pads, moistened by numerous specialized sweat glands, which increases friction and help the animal maintain its grip when quickly moving up steep, rocky surfaces. The soles of the feet also have specialized muscles in this regard, which helps the soles to work somewhat like suction cups. The plane of symmetry of the feet goes through the third digit (Myers 2000).
Hyraxes have skulls with a short rostrum and prominent postorbital processes. They have a single incisor on each side of the upper jaw and two on the lower. Like [[rodent]]s, the incisors grow continuously throughout life, and the presence of enamel on only one side (the anterior surface) creates a self-sharpening cutting edge. The molars have strongly developed lophs. Unlike most other browsing and grazing animals, they do not use the [[incisor]]s at the front of the jaw for slicing off leaves and grass, rather, they use the [[molar (tooth)|molar]] teeth at the side of the jaw. There is a small [[diastema (dentistry)|diastema]] between the incisors and the cheek teeth. The [[dentition|dental formula]] for hyraxes is {{DentalFormula|upper=1.0.4.3|lower=2.0.4.3}}.
 
 
 
 
 
 
 
 
 
They also have efficient [[kidney]]s, retaining water so that they can better survive in arid environments.
 
 
 
 
 
* Myers, P. 2000. [http://animaldiversity.ummz.umich.edu/accounts/Hyracoidea/ Hyracoidea]. ''Animal Diversity Web''. Retrieved January 30, 2014.
 
 
 
 
 
Hyraxes retain a number of primitive mammalian characteristics; in particular, they have poorly developed internal temperature regulation, which they compensate for by behavioural [[thermoregulation]] such as  huddling and by basking in the sun.
 
 
 
[[File:Hyraxchewing.ogv|thumb|right|A hyrax showing the characteristic chewing/grunting behaviour.  Note the incisor tusks.]]
 
Although not [[ruminant]]s, hyraxes have complex, multi-chambered stomachs that allow [[Symbiosis|symbiotic]] bacteria to break down tough plant materials; their overall ability to digest fibre is similar to that of the [[ungulates]].<ref>von Engelhardt et al. (1978) Production of methane in two non-ruminant herbivores. Comparative Biochemistry and Physiology Part A: Physiology 60 (3) 309-311</ref> Their mandibular motions (see video) have often been described as chewing [[cud]],<ref>{{cite journal |first=Hubert |last=Hendrichs |year=1966 |title=Vergleichende Untersuchung des Wiederkauverhaltens |trans_title=Comparative investigation of cud retainers |journal=Biologisches Zentralblatt |volume=84 |issue=6 |pages=671–751 |language=German |type=dissertation |oclc=251821046 |quote=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}}{{Verify source|Where did the quote come frome? The article is in German. Who translated?|date=February 2011}}</ref> although there is no evidence this behaviour is associated with the [[Regurgitation (digestion)|regurgitation]] of stomach contents<ref>JB Sale - JE Afr. Nat. Hist. Soc, 1966</ref> as in the [[even-toed ungulate]]s and some of the [[Macropodidae|macropod]]s.  This behaviour is referred to in a passage in the Bible ([http://www.biblegateway.com/passage/?search=Leviticus%2011:5&version=NIV Leviticus 11:5]) which describes hyraxes as chewing the cud.  Some authors believe these chewing motions are a form of [[agonistic behaviour]] when the animal feels threatened, rather than being related to ingestion or [[mastication]].<ref name="zootorah.com">http://www.zootorah.com/hyrax/hyrax.pdf</ref>
 
  
 +
Hyraxes have skulls with a short rostrum and prominent postorbital processes. They have a single incisor on each side of the upper jaw and two on the lower. Like [[rodent]]s, the incisors grow continuously throughout life, and the presence of enamel on only one side (the anterior surface) creates a self-sharpening cutting edge. The molars have strongly developed lophs. Unlike most other browsing and grazing animals, they do not use the [[incisor]]s at the front of the jaw for slicing off leaves and grass, rather, they use the [[molar (tooth)|molar]] teeth at the side of the jaw. There is a small [[diastema (dentistry)|diastema]] between the incisors and the cheek teeth. The [[dentition|dental formula]] for hyraxes is {{DentalFormula|upper=1.0.4.3|lower=2.0.4.3}} (Myers 2000).
  
 +
Although not [[ruminant]]s, hyraxes have complex, multi-chambered stomachs that allow [[Symbiosis|symbiotic]] bacteria to break down tough plant materials; their overall ability to digest fibers is similar to that of the [[ungulates]] (von Engelhardt et al. 1978). Their mandibular motions have often been described as chewing [[cud]] (Hendrichs 1966), although there is no evidence this behavior is associated with the [[Regurgitation (digestion)|regurgitation]] of stomach contents (Sale 1966b) as in the [[even-toed ungulate]]s and some of the [[Macropodidae|macropod]]s. This behavior is referred to in a passage in the Bible (Leviticus 11:5), which describes hyraxes as chewing the cud. Hyraxes also have efficient [[kidney]]s, retaining water so that they can better survive in arid environments.
  
 +
Hyraxes retain a number of primitive mammalian characteristics; in particular, they have poorly developed internal temperature regulation, which they compensate for by behavioral [[thermoregulation]] such as  huddling and by basking in the sun.
  
 
==Distribution and habitat==
 
==Distribution and habitat==
 +
[[image:Rock Hyrax area.png|right|thumb|200px|Rock hyrax (''Procavia capensis'') range]]
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Hyraxes inhabit rocky terrain, forests, and moist savannah across [[sub-Saharan Africa]] and the [[Middle East]]. The rock hyrax (''Procavia capensis'') occurs across [[sub-Saharan Africa]], with the exception of the [[Congo basin]] and [[Madagascar]]. The western tree hyrax (''Dendrohyrax dorsalis''), is found in West and Central Africa: [[Benin]], [[Cameroon]], [[Central African Republic]], [[Republic of the Congo]], [[Democratic Republic of the Congo]], [[Ivory Coast]], [[Equatorial Guinea]], [[Gabon]], [[Gambia]], [[Ghana]], [[Guinea]], [[Guinea-Bissau]], [[Liberia]], [[Nigeria]], [[Rwanda]], [[Senegal]], [[Sierra Leone]], [[Sudan]], [[Togo]], [[Uganda]], and possibly [[Niger]]. The southern tree hyrax (''Dendrohyrax arboreus'') is found in Angola, Democratic Republic of the Congo, Kenya, Mozambique, South Africa, Tanzania, and Zimbabwe. ''Heterohyrax brucei'', the yellow-spotted rock hyrax or bush hyrax, is found in Angola, Botswana, Burundi, Democratic Republic of the Congo, Southern Egypt, Eritrea, Ethiopia, Kenya, Malawi, Mozambique, Rwanda, Somalia, Northern South Africa, Sudan, Tanzania, Uganda, Zambia, and Zimbabwe.
  
Hyraxes inhabit rocky terrain across [[sub-Saharan Africa]] and the [[Middle East]].  
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Some species are arboreal (the western tree hyrax and southern tree hyrax) and some live on rock outcrops (rock hyrax and yellow-spotted rock hyrax). While southern tree hyraxes are arboreal, and tend to live in dens created in trees and feed primarily in trees, there are reports of them sometimes occupying rocks (Williams 2004).  
  
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==Behavior==
  
==Behavior==
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Hyraxes are agile climbers and quick, and active during the daylight hours. Arboreal hyraxes tend to be solitary while those dwelling in rock outcrops tend to live in colonies (Myers 2000). Feeding is often in short, concentrated periods during the day, limiting their exposure to predators.
  
 
Female hyraxes give birth to up to four young after a [[gestation]] period of between seven and eight months, depending on the species. The young are [[weaning|weaned]] at one to five months of age, and reach sexual maturity at 16 to 17 months.     
 
Female hyraxes give birth to up to four young after a [[gestation]] period of between seven and eight months, depending on the species. The young are [[weaning|weaned]] at one to five months of age, and reach sexual maturity at 16 to 17 months.     
  
Hyraxes live in small family groups, dominated by a single male who aggressively defends the territory from rivals. Where there is abundant living space, the male may dominate 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.<ref name=EoM>{{cite book |editor=Macdonald, D. |author=Hoeck, Hendrik |year=1984 |title=The Encyclopedia of Mammals |publisher=Facts on File |location=New York |pages=462–5 |isbn=0-87196-871-1}}</ref>
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Hyraxes living in small family groups are dominated by a single male, who aggressively defends the territory from rivals. Where there is abundant living space, the male may dominate 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 (Hoeck 1984).
 
 
  
 
==Historical accounts==
 
==Historical accounts==
 
[[File:Klipdas.jpg|thumb|right|Hyrax on [[Mount Kenya]]]]
 
[[File:Klipdas.jpg|thumb|right|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 language|Hebrew]] שָּׁפָן ''shaphan''),<ref>[http://www.htmlbible.com/sacrednamebiblecom/kjvstrongs/FRMSTRHEB82.htm#S8227 "Shaphan" in ''Strong's Concordance'']</ref> and therefore no name for them. There are references to hyraxes in the [[Old Testament]],<ref>[http://www.gnpcb.org/esv/search/?q=Lev+11%3A4-8%3B+Deut+14%3A7%3B+Ps+104%3A18%3B+Prov+30%3A26 Lev 11:4-8; Deut 14:7; Ps 104:18; Prov 30:26]</ref> particularly in Leviticus 11, where they are described as lacking a split hoof and therefore being not [[kosher]]. It also incorrectly claims that the hyrax chews its cud. Some of the modern translations refer to them as rock badgers. [[Shaphan]] was also the name of a scribe of King [[Josiah]].
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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 language|Hebrew]] שָּׁפָן ''shaphan''),and therefore no name for them. There are references to hyraxes in the [[Old Testament]] (Lev 11:4-8; Deut 14:7; Ps 104:18; Prov 30:26) and notably Leviticus 11 in which they are described as lacking a split hoof and therefore being not [[kosher]]. It also incorrectly claims that the hyrax chews its cud. Some of the modern translations refer to them as rock badgers.
  
[[Phoenicia]]n [[sailor]]s visiting the coast of [[Spain]] ''circa'' [[12th century B.C.E.|1100s B.C.E.]], mistaking the [[European rabbit]] for the rock hyrax from their native homeland, gave it the name ''i-shepan-ham''. A theory exists that an adaptation and/or corruption of this name, used by the [[Ancient Rome|Romans]], became [[Hispania]], leading to Spanish ''España'' and English ''Spain'', although this theory is somewhat controversial.<ref>[http://books.google.com/books?id=S1lDAAAAIAAJ&printsec=frontcover&dq=A+System+of+Ancient+and+Medi%C3%A6val+Geography+for+the+Use+of+Schools+and+Colleges Anthon, Charles. A System of Ancient and Mediæval Geography for the Use of Schools and Colleges pg.14]</ref>
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[[Phoenicia]]n [[sailor]]s visiting the coast of [[Spain]] ''circa'' [[12th century B.C.E.|1100s B.C.E.]], mistaking the [[European rabbit]] for the rock hyrax from their native homeland, gave it the name ''i-shepan-ham''. A theory exists that an adaptation and/or corruption of this name, used by the [[Ancient Rome|Romans]], became [[Hispania]], leading to Spanish ''España'' and English ''Spain'', although this theory is somewhat controversial.
  
==Evolution==
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==Classification==
[[File:Tree-hyrax.jpg|thumb|right|Tree hyrax in the [[Serengeti]], [[Tanzania]]]]
 
All modern hyraxes are members of the family '''Procaviidae''' (the only living family within the 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.<ref>{{Cite journal|author=Eugenie Barrow; Erik R. Seiffert; Elwyn L. Simons.|year=2010|title=A primitive hyracoid (Mammalia, Paenungulata) from the early Priabonian (Late Eocene) of Egypt|journal=Journal of Systematic Palaeontology|volume=8|issue=2|pages=213–244|doi=10.1080/14772010903450407|url=http://www.informaworld.com/smpp/content~db=all~content=a922470879~frm=titlelink}}</ref> For many millions of years, hyraxes were the primary terrestrial herbivore in Africa, just as [[odd-toed ungulate]]s were in North America. Through the middle to late [[Eocene]], there were many different species,<ref>{{Cite book
 
  | last = Prothero
 
  | first = Donald R.
 
  | title = After the Dinosaurs: The Age of Mammals
 
  | publisher = [[Indiana University Press]]
 
  | year = 2006
 
  | location = Bloomington, Indiana
 
  | pages = 132
 
  | isbn = 978-0-253-34733-6}}</ref> 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 [[bovid]]s, which were very efficient grazers and browsers, displaced the hyraxes out of the prime territory and into marginal niches. Nevertheless, the order remained widespread, diverse and successful 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 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 [[toenail]]s, 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 [[bone]]s.<ref>"Hyrax: The Little Brother of the Elephant", ''Wildlife on One'', BBC TV.</ref>
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===Extant species===
 +
Four extant species currently are recognized; the [[rock hyrax]] (''Procavia capensis''), the [[yellow-spotted rock hyrax|yellow-spotted rock hyrax]] (''Heterohyrax brucei''), the [[western tree hyrax|western tree hyrax]] (''Dendrohyrax dorsalis'') and the [[southern tree hyrax|southern tree hyrax]] (''Dendrohyrax arboreus'').  
  
Hyraxes are sometimes described as being the closest living relative to the elephant,<ref>{{Cite web|url=http://www.economist.com/science/displaystory.cfm?story_id=12926018|title=Hirax song is a menu for mating|accessdate=15 January 2009|work=The Economist|date=15 January 2009}}</ref> 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 (cladistics)|outgroup]] to the assemblage of elephants, sirenians, and extinct orders such as [[Embrithopoda]] and [[Desmostylia]].<ref name=Asher>{{cite journal | author = Asher, R.J., Novacek, M.J., Geisher, J.H.| year = 2003 | title = Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence | journal = J. Mamm. Evol. | volume = 10 | pages = 131–194 | doi = 10.1023/A:1025504124129}}</ref>
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More species used to be recognized, but in the 2000s, taxonomists [[lumpers and splitters|reduced]] the number of recognized species of hyrax. In 1995, there were eleven or more recognized species; in 2013, only four are recognized. The animals which are no longer recognized as species are regarded as subspecies of the recognized four. There are over 50 recognized subspecies and species, many of which are considered highly endangered (Shoshani 2005).
  
===Similarities with elephants and sirenia===
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* '''ORDER HYRACOIDEA'''
Hyraxes share several unusual characteristics with [[elephant]]s and [[sirenia]] (manatees and dugongs) which have resulted in them 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;<ref name="Carnaby2008">{{cite book|author=Trevor Carnaby|title=Beat about the Bush: Mammals|url=http://books.google.com/books?id=hjt83FfPShsC&pg=PA293|date=1 January 2008|publisher=Jacana Media|isbn=978-1-77009-240-2|page=293}}</ref><ref name="Sisson1914">{{cite book|author=Septimus Sisson|title=The anatomy of the domestic animals|url=http://books.google.com/books?id=UgszAQAAIAAJ&pg=PA577|year=1914|publisher=W.B. Saunders Company|page=577}}</ref> the same as elephants, manatees, and dugongs.<ref>{{cite book|author=Marshall Cavendish Corporation|title=Mammal Anatomy: An Illustrated Guide|url=http://books.google.com/books?id=mTPI_d9fyLAC&pg=PA63|date=1 September 2010|publisher=Marshall Cavendish|isbn=978-0-7614-7882-9|page=63}}</ref> 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; in most other mammals, tusks develop from the [[Canine tooth|canines]].  Hyraxes, like elephants, have flattened [[Nail (anatomy)|nails]] on the tips of their digits, rather than curved, elongated [[claws]] which are usually seen on mammals (see<ref>[http://ts4.mm.bing.net/th?id=H.4688013754893047&pid=15.1 Picture of hyrax feet]</ref>).
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** '''Family Procaviidae'''
 
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*** Genus ''[[Dendrohyrax]]''
Hyraxes have highly charged [[myoglobin]], which has been inferred to reflect an aquatic ancestry.<ref>{{Cite web|first=|title=One Protein Shows Elephants and Moles Had Aquatic Ancestors|url=http://phenomena.nationalgeographic.com/2013/06/13/one-protein-shows-elephants-and-moles-had-aquatic-ancestors/|doi=|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. —>{{inconsistent citations}}}}</ref>
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**** [[Southern tree hyrax]], ''Dendrohyrax arboreus''
 
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**** [[Western tree hyrax]], ''Dendrohyrax dorsalis''
==Classification==
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*** Genus ''[[Heterohyrax]]''
 +
**** [[Yellow-spotted rock hyrax]], ''Heterohyrax brucei''
 +
*** Genus ''[[Procavia]]''
 +
**** [[Rock hyrax]], ''Procavia capensis''
  
The [[extinct]] [[meridiungulata|meridiungulate]] [[Family (biology)|family]] '''[[Archaeohyracidae]]''', consisting of four [[genus|genera]] of [[Notoungulata|notoungulate]] [[mammal]]s known from the [[Paleocene]] through the [[Oligocene]] of [[South America]]<ref>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</ref> is a group unrelated to the true hyraxes.
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===Extinct species===
  
===List of extinct species===
 
 
*[[Pliohyracidae]]
 
*[[Pliohyracidae]]
 
**''[[Geniohyinae]]''
 
**''[[Geniohyinae]]''
Line 123: Line 103:
 
****''[[Procavia transvaalensis]]''
 
****''[[Procavia transvaalensis]]''
  
==Extant species==
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The [[extinct]] [[meridiungulata|meridiungulate]] [[Family (biology)|family]] '''[[Archaeohyracidae]]''', consisting of four [[genus|genera]] of [[Notoungulata|notoungulate]] [[mammal]]s known from the [[Paleocene]] through the [[Oligocene]] of [[South America]] (McKenna and Bell 1997), is a group unrelated to the true hyraxes.
In the 2000s, taxonomists [[lumpers and splitters|reduced]] the number of recognized species of hyrax. In 1995, there were eleven or more recognized species; in 2013, only four are recognized. The animals which are no longer recognised as species are regarded as subspecies of the recognized four. There are over 50 recognized subspecies and species, many of which are considered highly endangered.<ref name=MSW3>{{MSW3 Shoshani|pages=87–89|id=11400002}}</ref>
 
  
* '''ORDER HYRACOIDEA'''
+
==Evolution==
** '''Family Procaviidae'''
+
[[File:Tree-hyrax.jpg|thumb|right|Tree hyrax in the [[Serengeti]], [[Tanzania]]]]
*** Genus ''[[Dendrohyrax]]''
+
All modern hyraxes are members of the family '''Procaviidae''' (the only living family within the 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 (Barrow et al. 2010). For many millions of years, hyraxes were the primary terrestrial herbivore in Africa, just as [[odd-toed ungulate]]s were in North America. Through the middle to late [[Eocene]], there were many different species (Prothero 2006), 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 [[bovid]]s, which were very efficient grazers and browsers, displaced the hyraxes out of the prime territory and into marginal niches. Nevertheless, the order remained widespread, diverse, and successful as late as the end of the [[Pliocene]] (about two million years ago) with representatives throughout most of Africa, [[Europe]], and [[Asia]].
**** [[Southern tree hyrax]], ''Dendrohyrax arboreus''
+
 
**** [[Western tree hyrax]], ''Dendrohyrax dorsalis''
+
The descendants of the giant hyracoids 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 [[siren]]ians. Hyraxes have highly charged [[myoglobin]], which has been inferred to reflect an aquatic ancestry (Yong 2013).
*** Genus ''[[Heterohyrax]]''
+
 
**** [[Yellow-spotted rock hyrax]], ''Heterohyrax brucei''
+
===Similarities with elephants and sirenia===
*** Genus ''[[Procavia]]''
+
 
**** [[Rock hyrax]], ''Procavia capensis''
+
Hyraxes share several unusual characteristics with [[elephant]]s and [[sirenia]] (manatees and dugongs), which have resulted in them 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 (Carnaby 2008; Sisson 1914)); the same as elephants, manatees, and dugongs (Marshall Cavendish 2010). 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; in most other mammals, tusks develop from the [[Canine tooth|canines]].  Hyraxes, like elephants, have flattened [[Nail (anatomy)|nails]] on the tips of their digits, rather than curved, elongated [[claws]], which are usually seen on mammals.
 +
 
 +
The small modern hyraxes share other features with elephants, such as [[toenail]]s, 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 [[bone]]s (BBC 2000). [[DNA]] evidence also supports the hypothesis of a close connection between hyrax, elephants, and sirenians.
  
 +
Hyraxes are sometimes described as being the closest living relative to the elephant (Economist 2009), 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 (cladistics)|outgroup]] to the assemblage of elephants, sirenians, and extinct orders such as [[Embrithopoda]] and [[Desmostylia]] (Asher et al. 2003).
  
 
==References==
 
==References==
{{Reflist}}
+
* Asher, R. J., M. J. Novacek, and J. H.. Geisher. 2003. Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence. ''J. Mamm. Evol.'' 10: 131–194.
 +
 
 +
* Barrow, E., E. R. Seiffert, and  E. L. Simons. 2010. [http://www.informaworld.com/smpp/content~db=all~content=a922470879~frm=titlelink A primitive hyracoid (Mammalia, Paenungulata) from the early Priabonian (Late Eocene) of Egypt]. ''Journal of Systematic Palaeontology'' 8(2): 213–244. Received February 2, 2014.
 +
 
 +
* BBC TV. 2000. [http://www.ovguide.com/tv_episode/wildlife-on-one-season-29-episode-8-hyrax-little-brother-of-the-elephant-684361 Hyrax: The little brother of the elephant]. ''Wildlife on One'' Season 29, Episode 8, Air date Oct. 2, 2000. Retrieved February 2, 2014.
 +
 
 +
* Carnaby, R. 2008. ''Beat about the Bush: Mammals''. Jacana Media. IBSN 9781770092402.
 +
 
 +
* Economist. 2009. [http://www.economist.com/science/displaystory.cfm?story_id=12926018 The song of songs: A hyrax’s song gives a detailed account of his wares]. ''The Economist'' January 15, 2009. Retrieved February 2, 2104.
 +
 
 +
* Hendrichs, H. 1966. Vergleichende untersuchung des wiederkauverhaltens (transl. Comparative investigation of cud retainers. ''Biologisches Zentralblatt'' 84(6): 671–751.
 +
 
 +
* Hoeck, H. 1984. Hyraxes. Pages 462-5 in D. Macdonald, ''The Encyclopedia of Mammals''. New York: Facts on File.  ISBN 0871968711.
 +
 
 +
* Marshall Cavendish Corp. 2010. [http://books.google.com/books?id=mTPI_d9fyLAC&pg=PA63 ''Mammal Anatomy: An Illustrated Guide'']. Marshall Cavendish. ISBN 9780761478829. Retrieved February 2, 2014.
 +
 
 +
* McKenna, M. C., and S. K. Bell. 1997. ''Classification of Mammals Above the Species Level.'' New York: Columbia University Press. ISBN 0231110138.
 +
 
 +
* Myers, P. 2000. [http://animaldiversity.ummz.umich.edu/accounts/Hyracoidea/ Hyracoidea]. ''Animal Diversity Web''. Retrieved January 30, 2014.
 +
 
 +
* Prothero, D. R. 2006. ''After the Dinosaurs: The Age of Mammals. Bloomington, IN: Indiana University Press. ISBN 9780253347336.
 +
 
 +
* Sale, J. B. 1966a. The habitat of the rock hyrax. ''JE Afr. Nat. Hist. Soc.'' 25: 205-214.
 +
 
 +
* Sale, J. B. 1966b. Daily food consumption in the hyrax. ''JE Afr. Nat. Hist. Soc.'' 25: 215-224.
 +
 
 +
* Shoshani, J. 2005. Order Hyracoidea. In D. E. Wilson and D. M. Reeder, eds., ''Mammal Species of the World'' 3rd ed. Johns Hopkins University. ISBN 9780801882210.
 +
 
 +
* Sisson, S. 1914. ''The Anatomy of the Domestic Animals''. W.B. Saunders Company.
 +
 
 +
* von Engelhardt, W., S. Wolter, H. Lawrenz, and J. A. Hemsley. 1978. Production of methane in two non-ruminant herbivores. ''Comparative Biochemistry and Physiology. Part A: Physiology'' 60(3): 309-311.
 +
 
 +
* Williams, A. 2004. [http://animaldiversity.ummz.umich.edu/accounts/Dendrohyrax_arboreus/ Dendrohyrax arboreuseastern tree hyrax(Also: southern tree hyrax)]. ''Animal Diversity Web''. Retrieved January 30, 2014.
 +
 
 +
* Yong, E. 2013. [http://phenomena.nationalgeographic.com/2013/06/13/one-protein-shows-elephants-and-moles-had-aquatic-ancestors One protein shows elephants and moles had aquatic ancestors]. ''National Geographic'' June 13, 2013. Retrieved February 2, 2013.
  
{{Mammals}}
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{{Mammals|state=collapsed}}
{{Hyracoidea}}
 
  
  

Latest revision as of 21:54, 11 May 2014

Hyraxes
Fossil range: Eocene-Recent, 55.8–0 Ma
Yellow-spotted hyrax (Heterohyrax brucei)
Yellow-spotted hyrax (Heterohyrax brucei)
Scientific classification
Families
  • Procaviidae
  • Pliohyracidae

Hyrax is the common name for any of the largely herbivorous mammals comprising the order Hyracoidea, whose modern members are characterized by thickset bodies, short tails, and three-toed hind feet with rounded, hoof-like nails on two of the toes and a claw on the third. These mammals, which can be described as having a guinea pig or woodchuck-like body, and are similar in size to a large rabbit, are also known as cories and dassies. The presence of specialized muscles in the soles of the feet, combined with the soft and elastic nature of the sweat-gland moistened soles, helps to create a suction-like friction with rocks and other substrate, making them agile climbers.

There is one extant family of hyraxes, Procaviidae, distributed only in Africa and the Middle East. Four species are recognized: the rock hyrax (Procavia capensis), the yellow-spotted rock hyrax (Heterohyrax brucei), the western tree hyrax (Dendrohyrax dorsalis) and the southern tree hyrax (Dendrohyrax arboreus). However, in evolutionary history hyraxes were more diverse, and with larger and smaller representatives, and for many millions of years they were the primary terrestrial herbivore in Africa. Hyraxes are often mistaken for rodents, but are more closely related to elephants.

Hyraxes provide diverse values for the ecosystem and for humans. As part of food chains, they consume many tree, shrub, and grass plant parts (leaves, petioles, twigs, shoots, fruits, seeds) and occasionally insects, while being consumed by birds of prey (eagles, owls), large cats (lions, leopards), jackals, snakes, and mongooses. Hyraxes form the almost exclusive diet of Verreaux eagles. Hyraxes also disperse the seeds of fruits they eat and play a role in nutrient cycling. For humans, hyraxes have been hunted for food and fur and their crystallized excrement has been used by Europeans and African tribes to treat various illness (epilepsy, hysteria, convulsions) and as vitamin supplements.

Physical description

Hyraxes are well-furred, short-tailed, rotund animals about the size of a large rabbit. Typically, they measure between 30 and 70 centimeters (12-28 inches) long and weigh between 2 and 5 kilograms (4-11 pounds).

The characteristic foot pads

Hyraxes have 3-toed hind feet with two of the toes with rounded nails that resemble hooves, while the inner toe has a claw. Forefeet have four or five toes. The soles of the fore- and hindfeet are soft, elastic, rubbery like pads, moistened by numerous specialized sweat glands, which increases friction and help the animal maintain its grip when quickly moving up steep, rocky surfaces. The soles of the feet also have specialized muscles in this regard, which helps the soles to work somewhat like suction cups. The plane of symmetry of the feet goes through the third digit (Myers 2000).

Hyraxes have skulls with a short rostrum and prominent postorbital processes. They have a single incisor on each side of the upper jaw and two on the lower. Like rodents, the incisors grow continuously throughout life, and the presence of enamel on only one side (the anterior surface) creates a self-sharpening cutting edge. The molars have strongly developed lophs. 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. There is a small diastema between the incisors and the cheek teeth. The dental formula for hyraxes is (Myers 2000).

Although not ruminants, hyraxes have complex, multi-chambered stomachs that allow symbiotic bacteria to break down tough plant materials; their overall ability to digest fibers is similar to that of the ungulates (von Engelhardt et al. 1978). Their mandibular motions have often been described as chewing cud (Hendrichs 1966), although there is no evidence this behavior is associated with the regurgitation of stomach contents (Sale 1966b) as in the even-toed ungulates and some of the macropods. This behavior is referred to in a passage in the Bible (Leviticus 11:5), which describes hyraxes as chewing the cud. Hyraxes also have efficient kidneys, retaining water so that they can better survive in arid environments.

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

Distribution and habitat

Rock hyrax (Procavia capensis) range

Hyraxes inhabit rocky terrain, forests, and moist savannah across sub-Saharan Africa and the Middle East. The rock hyrax (Procavia capensis) occurs across sub-Saharan Africa, with the exception of the Congo basin and Madagascar. The western tree hyrax (Dendrohyrax dorsalis), is found in West and Central Africa: Benin, Cameroon, Central African Republic, Republic of the Congo, Democratic Republic of the Congo, Ivory Coast, Equatorial Guinea, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Liberia, Nigeria, Rwanda, Senegal, Sierra Leone, Sudan, Togo, Uganda, and possibly Niger. The southern tree hyrax (Dendrohyrax arboreus) is found in Angola, Democratic Republic of the Congo, Kenya, Mozambique, South Africa, Tanzania, and Zimbabwe. Heterohyrax brucei, the yellow-spotted rock hyrax or bush hyrax, is found in Angola, Botswana, Burundi, Democratic Republic of the Congo, Southern Egypt, Eritrea, Ethiopia, Kenya, Malawi, Mozambique, Rwanda, Somalia, Northern South Africa, Sudan, Tanzania, Uganda, Zambia, and Zimbabwe.

Some species are arboreal (the western tree hyrax and southern tree hyrax) and some live on rock outcrops (rock hyrax and yellow-spotted rock hyrax). While southern tree hyraxes are arboreal, and tend to live in dens created in trees and feed primarily in trees, there are reports of them sometimes occupying rocks (Williams 2004).

Behavior

Hyraxes are agile climbers and quick, and active during the daylight hours. Arboreal hyraxes tend to be solitary while those dwelling in rock outcrops tend to live in colonies (Myers 2000). Feeding is often in short, concentrated periods during the day, limiting their exposure to predators.

Female hyraxes give birth to up to four young after a gestation period of 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 living in small family groups are dominated by a single male, who aggressively defends the territory from rivals. Where there is abundant living space, the male may dominate 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 (Hoeck 1984).

Historical accounts

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 shaphan),and therefore no name for them. There are references to hyraxes in the Old Testament (Lev 11:4-8; Deut 14:7; Ps 104:18; Prov 30:26) and notably Leviticus 11 in which they are described as lacking a split hoof and therefore being not kosher. It also incorrectly claims that the hyrax chews its cud. Some of the modern translations refer to them as rock badgers.

Phoenician sailors visiting the coast of Spain circa 1100s B.C.E., mistaking the European rabbit for the rock hyrax from their native homeland, gave it the name i-shepan-ham. A theory exists that an adaptation and/or corruption of this name, used by the Romans, became Hispania, leading to Spanish España and English Spain, although this theory is somewhat controversial.

Classification

Extant species

Four extant species currently are recognized; the rock hyrax (Procavia capensis), the yellow-spotted rock hyrax (Heterohyrax brucei), the western tree hyrax (Dendrohyrax dorsalis) and the southern tree hyrax (Dendrohyrax arboreus).

More species used to be recognized, but in the 2000s, taxonomists reduced the number of recognized species of hyrax. In 1995, there were eleven or more recognized species; in 2013, only four are recognized. The animals which are no longer recognized as species are regarded as subspecies of the recognized four. There are over 50 recognized subspecies and species, many of which are considered highly endangered (Shoshani 2005).

  • ORDER HYRACOIDEA
    • Family Procaviidae
      • Genus Dendrohyrax
        • Southern tree hyrax, Dendrohyrax arboreus
        • Western tree hyrax, Dendrohyrax dorsalis
      • Genus Heterohyrax
        • Yellow-spotted rock hyrax, Heterohyrax brucei
      • Genus Procavia
        • Rock hyrax, Procavia capensis

Extinct species

  • Pliohyracidae
    • Geniohyinae
      • Seggeurius
      • Geniohyus
    • Saghatheriinae
      • Microhyrax
      • Meroehyrax
      • Selenohyrax
      • Bunohyrax
      • Pachyhyrax
      • Megalohyrax
      • Saghatherium
      • Thyrohyrax
    • Titanohyracinae
      • Antilohyrax
      • Titanohyrax
    • Pliohyracinae
      • Sogdohyrax
      • Kvabebihyrax
      • Prohyrax
      • Parapliohyrax
      • Pliohyrax
      • Postschizotherium
  • Procaviidae
    • Procaviinae
      • Gigantohyrax
      • Procavia (Rock hyrax)
        • Procavia antigua
        • Procavia transvaalensis

The extinct meridiungulate family Archaeohyracidae, consisting of four genera of notoungulate mammals known from the Paleocene through the Oligocene of South America (McKenna and Bell 1997), is a group unrelated to the true hyraxes.

Evolution

Tree hyrax in the Serengeti, Tanzania

All modern hyraxes are members of the family Procaviidae (the only living family within the 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 (Barrow et al. 2010). For many millions of years, hyraxes were the primary terrestrial herbivore in Africa, just as odd-toed ungulates were in North America. Through the middle to late Eocene, there were many different species (Prothero 2006), 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 out of the prime territory and into marginal niches. Nevertheless, the order remained widespread, diverse, and successful 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 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. Hyraxes have highly charged myoglobin, which has been inferred to reflect an aquatic ancestry (Yong 2013).

Similarities with elephants and sirenia

Hyraxes share several unusual characteristics with elephants and sirenia (manatees and dugongs), which have resulted in them 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 (Carnaby 2008; Sisson 1914)); the same as elephants, manatees, and dugongs (Marshall Cavendish 2010). 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; in most other mammals, 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.

The small modern hyraxes share other 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 (BBC 2000). DNA evidence also supports the hypothesis of a close connection between hyrax, elephants, and sirenians.

Hyraxes are sometimes described as being the closest living relative to the elephant (Economist 2009), 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 extinct orders such as Embrithopoda and Desmostylia (Asher et al. 2003).

References
ISBN links support NWE through referral fees

  • Asher, R. J., M. J. Novacek, and J. H.. Geisher. 2003. Relationships of endemic African mammals and their fossil relatives based on morphological and molecular evidence. J. Mamm. Evol. 10: 131–194.
  • Carnaby, R. 2008. Beat about the Bush: Mammals. Jacana Media. IBSN 9781770092402.
  • Hendrichs, H. 1966. Vergleichende untersuchung des wiederkauverhaltens (transl. Comparative investigation of cud retainers. Biologisches Zentralblatt 84(6): 671–751.
  • Hoeck, H. 1984. Hyraxes. Pages 462-5 in D. Macdonald, The Encyclopedia of Mammals. New York: Facts on File. ISBN 0871968711.
  • McKenna, M. C., and S. K. Bell. 1997. Classification of Mammals Above the Species Level. New York: Columbia University Press. ISBN 0231110138.
  • Myers, P. 2000. Hyracoidea. Animal Diversity Web. Retrieved January 30, 2014.
  • Prothero, D. R. 2006. After the Dinosaurs: The Age of Mammals. Bloomington, IN: Indiana University Press. ISBN 9780253347336.
  • Sale, J. B. 1966a. The habitat of the rock hyrax. JE Afr. Nat. Hist. Soc. 25: 205-214.
  • Sale, J. B. 1966b. Daily food consumption in the hyrax. JE Afr. Nat. Hist. Soc. 25: 215-224.
  • Shoshani, J. 2005. Order Hyracoidea. In D. E. Wilson and D. M. Reeder, eds., Mammal Species of the World 3rd ed. Johns Hopkins University. ISBN 9780801882210.
  • Sisson, S. 1914. The Anatomy of the Domestic Animals. W.B. Saunders Company.
  • von Engelhardt, W., S. Wolter, H. Lawrenz, and J. A. Hemsley. 1978. Production of methane in two non-ruminant herbivores. Comparative Biochemistry and Physiology. Part A: Physiology 60(3): 309-311.
Mammals
Monotremata (platypus, echidnas)

Marsupialia: | Paucituberculata (shrew opossums) | Didelphimorphia (opossums) | Microbiotheria | Notoryctemorphia (marsupial moles) | Dasyuromorphia (quolls and dunnarts) | Peramelemorphia (bilbies, bandicoots) | Diprotodontia (kangaroos and relatives)

Placentalia: Cingulata (armadillos) | Pilosa (anteaters, sloths) | Afrosoricida (tenrecs, golden moles) | Macroscelidea (elephant shrews) | Tubulidentata (aardvark) | Hyracoidea (hyraxes) | Proboscidea (elephants) | Sirenia (dugongs, manatees) | Soricomorpha (shrews, moles) | Erinaceomorpha (hedgehogs and relatives) Chiroptera (bats) | Pholidota (pangolins)| Carnivora | Perissodactyla (odd-toed ungulates) | Artiodactyla (even-toed ungulates) | Cetacea (whales, dolphins) | Rodentia (rodents) | Lagomorpha (rabbits and relatives) | Scandentia (treeshrews) | Dermoptera (colugos) | Primates |

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