The range map of Procaviidae, the only living family within Hyracoidea
Hyraxes (from
ancient Greekὕραξ (húrax)
"
shrew-mouse"), also called
dassies,[1][2]
are small, stout, herbivorous
mammals in the
orderHyracoidea. Hyraxes are well-furred, rotund animals with short tails.[3] Modern hyraxes are typically between 30 and 70 cm (12 and 28 in) long and weigh between 2 and 5 kg (4 and 11 lb). They are superficially similar to
marmots, or over-large
pikas, but are paradoxically much more closely related to
elephants and
sea cows. Hyraxes have a life span from 9 to 14 years. Both types of 'rock' hyrax (
P. capensis and
H. brucei) live on rock outcrops, including cliffs in Ethiopia[4]
and isolated granite outcrops called
koppies in southern Africa;[5]
With one exception, all hyraxes are limited to
Africa; the exception is the
rock hyrax (P. capensis) which is also found in adjacent parts of the
Middle East.
Hyraxes were a much more diverse group in the past encompassing species considerably larger than modern hyraxes. The largest known extinct hyrax, Titanohyrax ultimus has been estimated to weigh 600–1,300 kilograms (1,300–2,900 lb), comparable to a rhinoceros.[6]
Characteristics
Hyraxes retain or have redeveloped a number of primitive mammalian characteristics; in particular, they have poorly developed
internal temperature regulation,[7]
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 those of 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.
A hyrax showing its characteristic chewing, grunting behavior, and incisor tusks
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.[8]
Their mandibular motions are similar to chewing
cud,[9][a]
but the hyrax is physically incapable of
regurgitation[10][11]
as in the
even-toed ungulates and some of the
macropods. This behaviour is referred to in a passage in the Bible which describes hyraxes as "chewing the cud".[12]
This chewing behaviour may be a form of
agonistic behaviour when the animal feels threatened.[13]
The hyrax does not construct dens, as most
rodents and rodent-like mammals do, but over the course of its lifetime rather seeks shelter in existing holes of great variety in size and configuration.[14]
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; four toes are on each front foot and three are on each back foot.[15]
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 of 7–8 months, depending on the species. The young are
weaned at 1–5 months of age, and reach sexual maturity at 16–17 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 its own range. The remaining males live solitary lives, often on the periphery of areas controlled by larger males, and mate only with younger females.[16]
Hyraxes have highly charged
myoglobin, which has been inferred to reflect an aquatic ancestry.[17]
Similarities with Proboscidea and Sirenia
Hyraxes share several unusual characteristics with mammalian orders
Proboscidea (elephants and their extinct relatives) and
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,[18][19]
as do those of elephants, manatees, and dugongs.[20]
Female hyraxes have a pair of
teats near their armpits (
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.[21][22]
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 the curved, elongated
claws usually seen on mammals.[23]
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.[24] For many millions of years, hyraxes, proboscideans, and other afrotherian
mammals 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.[25] The smallest of these were the size of a mouse but others were much larger than any extant relatives. Titanohyrax could reach 600 kg (1,300 lb) or even as much as over 1,300 kg (2,900 lb).[26]Megalohyrax from the upper Eocene-lower
Oligocene was as huge as a tapir.[27][28]
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 with other similar mammals, and the shape of some of their
bones.[29]
Hyraxes are sometimes described as being the closest living relative of the elephant,[30] 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.[31]
A phylogeny of hyracoids known from the early Eocene through the middle Oligocene epoch.[35]
^Modern day hyrax species (
Procaviidae) may have evolved from smaller members of one of the
Saghatheriinae.
^The relationship of
hyracoids and
perissodactyls is controversial, and not supported by molecular data.
^The position of the
phenacodontids has been long unresolved; some authorities suspect that they are basal members of the
perissodactyls, but that placement is contested.[33][34]
In the 2000s, taxonomists
reduced the number of recognized species of hyraxes. In 1995, they recognized 11 species or more, but as of 2013, only four were recognized, with the others all considered as subspecies of one of the recognized four. Over 50 subspecies and species are described, many of which are considered highly endangered.[38] The most recently identified species is Dendrohyrax interfluvialis, which is a tree hyrax living between the Volta and Niger rivers but makes a unique barking call that is distinct from the shrieking vocalizations of hyraxes inhabiting other regions of the African forest zone.[39]
The following cladogram shows the relationship between the extant genera:[40]
References are made to hyraxes in the
Hebrew Bible (
Leviticus 11:5;
Deuteronomy 14:7;
Psalm 104:18;
Proverbs 30:26). In Leviticus they are described as lacking a split hoof and therefore not being
kosher. It also describes the hyrax as chewing its cud, reflecting its observable ruminant-like mandible motions; the Hebrew phrase in question (מַעֲלֵה גֵרָה) means "bringing up cud". Some of the modern translations refer to them as rock hyraxes.[45][46]
... hyraxes are creatures of little power, yet they make their home in the crags; ...
The words "rabbit", "hare", "coney", or "daman" appear as terms for the hyrax in some English translations of the Bible.[47][48]
Early English translators had no knowledge of the hyrax, so no name for them, though "badger" or "rock-badger" has also been used more recently in new translations, especially in "common language" translations such as the
Common English Bible (2011).[49]
"Spain"
One of the proposed etymologies for "
Spain" is that it may be a derivation of the Phoenician I-Shpania, meaning "island of hyraxes", "land of hyraxes", but the Phoenecian-speaking
Carthaginians are believed to have used this name to refer to rabbits, animals with which they were unfamiliar.[50]
Roman coins struck in the region from the reign of
Hadrian show a female figure with a rabbit at her feet,[51]
and
Strabo called it the "land of the 'rabbits' ".[52]
The Phoenician shpania is cognate to the modern Hebrew shafan.[53][b]
Footnotes
^
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.[9][verification needed]
^Hispania, the name that the
Romans gave to the peninsular, derives from the
Phoeniciani-spn-ya, where the prefix i would translate as "coast", "island", or "land", ya as "region" and spn[,] in Hebrew saphan, as "rabbits" (in reality, hyraxes). The Romans, therefore, gave Hispania the meaning of "land abundant in rabbits", a use adopted by
Cicero,
Caesar,
Pliny the Elder and, in particular,
Catulo, who referred to Hispania as the cuniculus peninsula.[53]
^
"Hyracoidea". Grzimek's Animal Life Encyclopedia. Vol. 15 Mammals (online ed.). Gale Publishing.
^"Dassie, n.". Dictionary of South African English (web ed.). Dictionary Unit for South African English. 2018. Retrieved 25 February 2019.
^
Wilson, Don E.; Mittermeier, Russell A. (eds.). Handbook of the Mammals of the World. Vol. 2. Barcelona, ES: Lynx Edicions. p. 29.
ISBN978-84-96553-77-4.
^
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): 309–311.
doi:
10.1016/0300-9629(78)90254-2.
^
ab
Hendrichs, Hubert (1966). "Vergleichende Untersuchung des Wiederkauverhaltens" [Comparative investigation of cud retainers]. Biologisches Zentralblatt (dissertation) (in German). 84 (6): 671–751.
OCLC251821046.
^
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): 649–653.
doi:
10.1016/0300-9629(94)90205-4.
PMID8529006.
^"Leviticus 11:5". Bible Gateway. Zondervan. Retrieved 8 April 2016.
^
Slifkin, Natan (11 March 2004).
"Chapter Six: Shafan the hyrax"(PDF). The Camel, the Hare, and the Hyrax. Archived from
the original(PDF) on 16 June 2012. Retrieved 25 April 2012.
^Barrow, Eugenie; Seiffert, Erik R.; Simons, Elwyn L. (2010). "A primitive hyracoid (Mammalia, Paenungulata) from the early Priabonian (Late Eocene) of Egypt". Journal of Systematic Palaeontology. 8 (2): 213–244.
Bibcode:
2010JSPal...8..213B.
doi:
10.1080/14772010903450407.
S2CID84398730.
^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". Journal of Mammalian Evolution. 10: 131–194.
doi:
10.1023/A:1025504124129.
S2CID39296485.
^
McKenna, Malcolm C.; Bell, Susan K. (1997). Classification of Mammals above the Species Level. New York, NY: Columbia University Press.
ISBN0-231-11013-8.
^
Gheerbrant, E.; Donming, D.; Tassy, P. (2005).
"Paenungulata (Sirenia, Proboscidea, Hyracoidea, and relatives)". In Rose, Kenneth D.; Archibald, J. David (eds.). The Rise of Placental Mammals: Origins and relationships of the major extant clades. Baltimore, MD: Johns Hopkins University Press. pp. 84–105.
ISBN978-0-8018-8022-3 – via Google books.
^
Tabuce, R.; Seiffert, E.R.; Gheerbrant, E.; Alloing-Séguier, L.; von Koenigswald, W. (2017). "Tooth enamel microstructure of living and extinct hyracoids reveals unique enamel types among mammals". Journal of Mammalian Evolution. 24 (1): 91–110.
doi:
10.1007/s10914-015-9317-6.
S2CID36591482.
^
Wood, John George (1877). Wood's Bible Animals. London, UK: J.W. Lyon.
OCLC976950183. A description of the habits, structure, and uses of every living creature mentioned in the Scriptures, from the ape to the coral; and explaining all those passages in the Hebrew Bibles and the Christian Old Testament in which reference is made to beast, bird, reptile, fish, or insect. Illustrated with over one hundred new designs
^
Wood, John George (2014) [1888].
Story of the Bible Animals (scanned ed.). Philadelphia, PA: Charles Foster's Publications. p.
367–371.
OCLC979571526. Retrieved 12 June 2024 – via Project Gutenberg. A description of the habits and uses of every living creature mentioned in the scriptures, with explanation of passages in the Old and New Testament in which reference is made to them
^
Buel, James W. (1889). The Living World. St. Louis, MO: Holloway & Co.
doi:
10.5962/bhl.title.163548. A complete natural history of the world's creatures, fishes, reptiles, insects, birds and mammals
^
Elwell, Walter A.; Comfort, Philip Wesley (2008). Tyndale Bible dictionary. Tyndale House Publishers.
ISBN978-1-4143-1945-2.
OCLC232301052.