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Giraffidae

From Wikipedia, the free encyclopedia
Family of mammals belonging to even-toed ungulates

Giraffidae
Temporal range:20–0 MaEarlyMiocene – recent
Anokapi inBristol Zoo, England
Masai giraffe (G. tippelskirchi) at theMikumi National Park,Tanzania
Scientific classificationEdit this classification
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Artiodactyla
Superfamily:Giraffoidea
Family:Giraffidae
Gray, 1821
Type genus
Giraffa
Linnaeus, 1758
Subfamilies

TheGiraffidae are afamily ofruminantartiodactyl mammals that share a recent common ancestor withdeer andbovids. This family, once a diverse group spread throughoutEurasia and Africa, presently comprises only two extant genera, thegiraffe (between one and eight, usually four, species ofGiraffa, depending on taxonomic interpretation) and theokapi (the only known species ofOkapia). Both are confined tosub-Saharan Africa: the giraffe to the opensavannas, and the okapi to the denserainforest of theCongo. The two genera look very different on first sight, but share a number of common features, including a long, dark-coloured tongue, lobed canine teeth, and horns covered in skin, calledossicones.

Taxonomy

[edit]
ImageGenusLiving species
Okapia
Giraffa

Evolutionary background

[edit]
Shansitherium andPalaeotragus microdon, two giraffids from the Miocene of Asia

The giraffids areruminants of the cladePecora. Other extant pecorans are the familiesAntilocapridae (pronghorns),Cervidae (deer),Moschidae (musk deer), andBovidae (cattle,goats and sheep,wildebeests and allies, andantelopes). The exact interrelationships among the pecorans have been debated, mainly focusing on the placement of Giraffidae, but a recent large-scale ruminant genome sequencing study suggests Antilocapridae are the sister taxon to Giraffidae, as shown in thecladogram below.[1]

Ruminantia

The ancestors of pronghorn diverged from the giraffids in theEarly Miocene.[1] This was in part of a relatively late mammal diversification following a climate change that transformedsubtropicalwoodlands into opensavannahgrasslands.

The fossil record of giraffids and their stem-relatives is quite intensive, with fossil of these taxa includeGelocidae,Palaeomerycidae,Prolibytheridae, andClimacoceratidae.[2][3] It is thought that the palaeomerycids, prolibytherids, climacoceratids and the giraffids all form a clade of pecorans known asGiraffomorpha.[2][4] The relationship between the climacoceratids and giraffids is supported by the presence of a bilobed canine,[2] and have been postulated into two hypotheses. One is the climacoceratids were the ancestors of thesivatheres, as both groups were large, deer-likegiraffoids with branching antler-likeossicones, while an extinct basal group of giraffoids,canthumerycines, evolved into the ancestors of Giraffidae.[3] Another more commonly supported hypothesis is climacoceratids were merely the sister clade to giraffids, with sivatheres being either basal giraffids[2] or descended from a lineage that also includes the okapi.[5] While the current range of giraffids today is inAfrica, the fossil record of the group has shown this family was once widespread throughout ofEurasia.[2][3][5]

Below is the phylogenetic relationships of giraffomorphs after Solounias (2007),[2] Sánchez et al. (2015)[4] and Ríos et al. (2017):[5]

Giraffomorpha

Classification

[edit]
Skeletal illustration ofHelladotherium, now extinct
Skeletal mount ofPalaeotragus on display at theTianjin Natural History Museum.
Skeletal mount ofShansitherium tafeli on display at theBeijing Museum of Natural History.

Below is the total taxonomy of valid extant and fossil taxa (as well as junior synonyms which are listed in the brackets).

Family GiraffidaeJ.E.Gray, 1821

  • Basal extinct giraffids
    • CsakvarotheriumKretzoi, 1930
      • Csakvarotherium hungaricumKretzoi, 1930
    • InjanatheriumHeintz, Brunet & Sen, 1981
      • Injanatherium arabicumMorales, Soria & Thomas, 1987
      • Injanatherium hazimiHeintz, Brunet & Sen, 1981
    • PropalaeomeryxLydekker, 1883 [ProgiraffaPilgrim, 1908]
      • Propalaeomeryx sivalensisLydekker, 1883 [Progiraffa exiguaPilgrim, 1908]
    • ShansitheriumKillgus, 1922 [Schansitherium[sic]]
      • Shansitherium quadricornis(Bohlin, 1926) [Palaeotragus quadricornisBohlin, 1926]
      • Shansitherium tafeliKillgus, 1922
    • UmbrotheriumAbbazzi, Delfino, Gallai, Trebini & Rook, 2008
      • Umbrotherium azzaroliiAbbazzi, Delfino, Gallai, Trebini & Rook, 2008
  • Subfamily †CanthumerycinaeHamilton, 1978
    • GeorgiomeryxParaskevaidis, 1940
      • Georgiomeryx georgalasiParaskevaidis, 1940
    • CanthumeryxHamilton 1973 [ZarafaHamilton, 1973]
      • Canthumeryx sirtensisHamilton 1973 [Zarafa zelteniHamilton, 1973]
  • Subfamily †ProgiraffinaePilgrim, 1911
    • PalaeogiraffaBonis & Bouvrain, 2003
      • Palaeogiraffa macedoniae(Geraads, 1989) [Decennatherium macedoniaeGeraads, 1989]
      • Palaeogiraffa majorBonis & Bouvrain, 2003
      • Palaeogiraffa pamiri(Ozansoy, 1965) [Samotherium pamiriOzansoy, 1965]
  • Subfamily †GiraffokerycinaeSolounias, 2007
    • GiraffokeryxPilgrim, 1910
      • Giraffokeryx anatoliensisGeraads & Aslan, 2003
      • Giraffokeryx primaevus(Churcher, 1970) [Palaeotragus primaevusChurcher, 1970;Samotherium africanumChurcher, 1970 andAmotherium africanum[sic]]
      • Giraffokeryx punjabiensisPilgrim, 1910
  • SubfamilyGiraffinaeJ.E.Gray, 1821
    • TribeGiraffiniJ.E.Gray, 1821
    • Tribe †BohlininiSolounias, 2007
      • HonanotheriumBohlin, 1927
        • Honanotherium bernoriSolounias & Danowitz, 2016
        • Honanotherium schlosseri(Pilgrim, 1911) [Giraffa schlosseriPilgrim, 1911]
      • QilinWang et al., 2025
        • Qilin tungurensis(Colbert, 1936) [Palaeotragus tungurensisColbert, 1936]
      • BohliniaMatthew, 1929
        • Bohlinia adoumiLikius, Vignaud & Brunet, 2007
        • Bohlinia attica(Gaudry & Lartet, 1856) [Giraffa attica(Gaudry & Lartet, 1856) andOrasius attica(Gaudry & Lartet, 1856)]
        • Bohlinia nikitiaeKostopoulos, Koliadimou & Koufos, 1996
    • TribePalaeotraginiPilgrim, 1910
      • Subtribe †PalaeotraginaPilgrim, 1910
        • MitilanotheriumSamson & Radulesco, 1966 [MacedonitheriumSickenberg, 1967;SogdianotheriumSharapov, 1974]
          • Mitilanotherium inexpectatumSamson & Radulesco, 1966[Macedonitherium martiniiSickenberg, 1967;Mitilanotherium inexpectatumSamson & Radulesco, 1966;Mitilanotherium kuruksaense(Sharapov, 1974);Mitilanotherium martinii(Sickenberg, 1967);Palaeotragus inexspectatus(Samson & Radulesco, 1966);Palaeotragus priasovicusGodina & Bajgusheva, 1985 andSogdianotherium kuruksaenseSharapov, 1974]
        • PalaeotragusGaudry, 1861 [AchtiariaBorissiak, 1914]
          • Palaeotragus coelophrys(Rodler & Weithofer, 1890) [Alcicephalus coelophrysRodler & Weithofer, 1890]
          • Palaeotragus germainiArambourg, 1959
          • Palaeotragus lavocantiHeintz, 1976
          • Palaeotragus robinsoniCrusafont-Pairó, 1979
          • Palaeotragus roueniiGaudry, 1861 [Palaeotragus microdonKoken, 1885]
        • PraepalaeotragusGodina, Vislobokova & Abdrachmanova, 1993
          • Praepalaeotragus actaensisGodina, Vislobokova & Abdrachmanova, 1993
        • SamotheriumForsyth Major, 1888 [AlcicephalusRodler & Weithofer, 1890;ChersenotheriumAlexajew, 1916 andAmotherium[sic]]
          • Samotherium boissieriForsyth Major, 1888
          • Samotherium eminens(Alexajew, 1916) [Chersenotherium eminensAlexajew, 1916]
          • Samotherium majorBohlin, 1926
          • Samotherium neumayri(Rodler & Weithofer, 1890) [Alcicephalus neumayriRodler & Weithofer, 1890]
          • Samotherium sinense(Schlosser, 1903) [Alcicephalus sinenseSchlosser, 1903]
      • SubtribeOkapiinaBohlin, 1926
  • †SubfamilySivatheriinaeBonaparte, 1850
    • BirgerbohliniaCrusafont Pairó, 1952
      • Birgerbohlinia schaubiCrusafont Pairó, 1952
    • BramatheriumFalconer, 1845 [HydaspitheriumLydekker, 1876]
      • Bramatherium giganteusKhan & Sarwar, 2002
      • Bramatherium grande(Lydekker, 1878) [Hydaspitherium grandeLydekker, 1878]
      • Bramatherium magnum(Pilgrim, 1910) [Hydaspitherium magnumPilgrim, 1910]
      • Bramatherium megacephalum(Lydekker, 1876) [Hydaspitherium megacephalumLydekker, 1876]
      • Bramatherium perimenseFalconer, 1845
      • Bramatherium progressusKhan, Sarwar & Khan, 1993
      • Bramatherium suchoviGodina, 1977
    • DecennatheriumCrusafont Pairó, 1952
      • Decennatherium rexRíos, Sánchez & Morales, 2017
      • Decennatherium pachecoiCrusafont Pairó, 1952
    • HelladotheriumGaudry, 1860
      • Helladotherium duvernoyi(Gaudry & Lartet, 1856) [Camelopardalis duvernoyiGaudry & Lartet, 1856]
    • SivatheriumFalconer & Cautley, 1836 [GriquatheriumHaughton, 1922;IndratheriumPilgrim, 1910;LibytheriumPomel, 1892 andOrangiatheriumvan Hoepen, 1932]
      • Sivatherium giganteumFalconer & Cautley, 1836
      • Sivatherium hendeyiHarris, 1976
      • Sivatherium maurusium(Pomel, 1892) [Libytherium maurusiumPomel, 1892;Griquatherium cingulatumHaughton, 1922;Helladotherium olduvaienseHopwood, 1934;Sivatherium olduvaiense(Hopwood, 1934);Libytherium olduvaienseHopwood, 1934 andOrangiatherium vanrhynivan Hoepen, 1932]
    • VishnutheriumLydekker, 1876
      • Vishnutherium iravadicumLydekker 1876

Characteristics

[edit]
Two giraffes

The giraffe stands 5–6 m (16–20 ft) tall, with males taller than females. The giraffe and the okapi have characteristic long necks and long legs. Ossicones are present on males and females in the giraffe, but only on males in the okapi.[6]

Giraffids share many common features with other ruminants. They have cloven hooves andcannon bones, much like bovids, and a complex, four-chambered stomach. They have no upperincisors or uppercanines, replacing them with a tough, horny pad. An especially longdiastema is seen between the front and cheek teeth. The latter areselenodont, adapted for grinding up tough plant matter.[7] Like most other ruminants, thedental formula for giraffids is0.0.3.33.1.3.3. Giraffids have prehensile tongues (specially adapted for grasping).[8]

The extant giraffids, the forest-dwelling okapi and the savannah-living giraffe, have several features in common, including a pair of skin-covered horns, called ossicones, up to 15 cm (5.9 in) long (absent in female okapis); a long, black, prehensile tongue; lobed canine teeth; patterned coats acting as camouflage; and a back sloping towards the rear. The okapi's neck is long compared to most ruminants, but not nearly so long as the giraffe's. Male giraffes are the tallest of all mammals: their horns reach 5.5 m (18 ft) above the ground and their shoulder 3.3 m (11 ft), whereas the okapi has a shoulder height of 1.7 m (5 ft 7 in).[9]

Distribution

[edit]

The two extant genera are now confined tosub-Saharan Africa. The okapi is restricted to a small range in the northern rainforest of theDemocratic Republic of Congo. Although the range of the giraffe is considerably larger, it once covered an area twice the present size – all parts of Africa that could offer an arid and dry landscape furnished with trees.[9]

Behavior

[edit]

The social structure and behavior is markedly different in okapis and giraffes, but although little is known of the okapi's behavior in the wild, a few things are known to be present in both species:[9]

  • They have an ambling gait similar to camels, with their weight supported alternately by their left and right legs, while their necks maintain balance. Giraffes can run up to 60 km/h (37 mph) this way and are documented to have covered 1,500 km (930 mi) in theSahel during the dry season.
  • The dominance hierarchy, which has been well-documented among giraffes, has also been seen among captive okapis. An adult giraffe head can weigh 30 kg (66 lb), and if necessary, male giraffes establish a hierarchy among themselves by swinging their heads at each other, horns first, a behavior known as "necking". A subordinate okapi signals submission by placing its head and neck on the ground.

Giraffes are sociable, whereas okapis live mainly solitary lives. Giraffes temporarily form herds of up to 20 individuals; these herds can be mixed or uniform groups of males and females, young and adults. Okapis are normally seen in mother-offspring pairs, although they occasionally gather around a prime food source. Giraffe are not territorial, but have ranges that can dramatically vary between – 5 and 654 km2 (1.9 and 252.5 sq mi) – depending on food availability, whereas okapis have individual ranges about 2.5–5 km2 (0.97–1.93 sq mi) in size.

  • Giraffes and okapis are normally silent, but both have a range of vocalizations, including coughing, snorting, moaning, hissing, and whistling. Giraffes have been suggested to be able to communicate usinginfrasonic sounds like elephants and blue whales.

Notes

[edit]
  1. ^A grade of giraffids.
  2. ^A paraphyletic grade of palaeotragines ancestral to Sivatheriinae.

References

[edit]
  1. ^abChen, L.; Qiu, Q.; Jiang, Y.; Wang, K. (2019)."Large-scale ruminant genome sequencing provides insights into their evolution and distinct traits".Science.364 (6446) eaav6202.Bibcode:2019Sci...364.6202C.doi:10.1126/science.aav6202.PMID 31221828.
  2. ^abcdefSolounias, N. (2007). "Family Giraffidae". In Prothero, D.R.; Foss, S.E. (eds.).The Evolution of Artiodactyls. The Johns Hopkins University Press. pp. 257–277.ISBN 978-0-8018-8735-2.
  3. ^abcSkinner, J.; Mitchell, G. (2011). "Family Giraffidae (Giraffe and Okapi)". In Wilson, D.E.; Mittermeier, R.A. (eds.).Handbook of the Mammals of the World – Volume II. Barcelona: Lynx Ediciones. pp. 788–802.ISBN 978-84-96553-77-4.
  4. ^abSánchez, Israel M.; Cantalapiedra, Juan L.; Ríos, María; Quiralte, Victoria; Morales, Jorge (2015)."Systematics and Evolution of the Miocene Three-Horned Palaeomerycid Ruminants (Mammalia, Cetartiodactyla)".PLOS ONE.10 (12) e0143034.Bibcode:2015PLoSO..1043034S.doi:10.1371/journal.pone.0143034.PMC 4668073.PMID 26630174.
  5. ^abcRíos, M.; Sánchez, I.M.; Morales, J. (2017)."A new giraffid (Mammalia, Ruminantia, Pecora) from the late Miocene of Spain, and the evolution of the sivathere-samothere lineage".PLOS ONE.12 (11) e0185378.Bibcode:2017PLoSO..1285378R.doi:10.1371/journal.pone.0185378.PMC 5665556.PMID 29091914.
  6. ^Dagg, A. I. (1971)."Giraffa camelopardalis"(PDF).Mammalian Species (5):1–8.doi:10.2307/3503830.JSTOR 3503830. Archived fromthe original(PDF) on 2017-04-21. Retrieved2015-04-17.
  7. ^Pellew, Robin (1984). MacDonald, D. (ed.).The Encyclopedia of Mammals. New York: Facts on File. pp. 534–541.ISBN 978-0-87196-871-5.
  8. ^Kingdon, Jonathan (2013).Mammals of Africa (1st ed.). London: A. & C. Black. pp. 95–115.ISBN 978-1-4081-2251-8.
  9. ^abcGrzimek, Bernhard (2003). Hutchins, Michael; Kleiman, Devra G; Geist, Valerius; et al. (eds.).Grzimek's Animal Life Encyclopedia, Vol 15, Mammals IV (2nd ed.). Farmington Hills, MI: Gale Group.ISBN 978-0-7876-5362-0.
ExtantArtiodactyla species
SuborderRuminantia
Antilocapridae
Antilocapra
Giraffidae
Okapia
Giraffa
Moschidae
Moschus
Tragulidae
Hyemoschus
Moschiola
Tragulus
Cervidae
Large family listed below
Bovidae
Large family listed below
FamilyCervidae
Cervinae
Muntiacus
Elaphodus
Dama
Axis
Rucervus
Elaphurus
Rusa
Cervus
Capreolinae
Alces
Hydropotes
Capreolus
Rangifer
Hippocamelus
Mazama
Ozotoceros
Blastocerus
Pudu
Pudella
Odocoileus
Subulo
FamilyBovidae
Hippotraginae
Hippotragus
Oryx
Addax
Reduncinae
Kobus
Redunca
Aepycerotinae
Aepyceros
Peleinae
Pelea
Alcelaphinae
Beatragus
Damaliscus
Alcelaphus
Connochaetes
Pantholopinae
Pantholops
Caprinae
Large subfamily listed below
Bovinae
Large subfamily listed below
Antilopinae
Large subfamily listed below
FamilyBovidae (subfamilyCaprinae)
Ammotragus
Arabitragus
Budorcas
Capra
Capricornis
Hemitragus
Naemorhedus
Oreamnos
Ovibos
Nilgiritragus
Ovis
Pseudois
Rupicapra
FamilyBovidae (subfamilyBovinae)
Boselaphini
Tetracerus
Boselaphus
Bovini
Bubalus
Bos
Pseudoryx
Syncerus
Tragelaphini
Tragelaphus
(includingkudus)
Taurotragus
FamilyBovidae (subfamilyAntilopinae)
Antilopini
Ammodorcas
Antidorcas
Antilope
Eudorcas
Gazella
Litocranius
Nanger
Procapra
Saigini
Saiga
Neotragini
Dorcatragus
Madoqua
Neotragus
Nesotragus
Oreotragus
Ourebia
Raphicerus
Cephalophini
Cephalophus
Philantomba
Sylvicapra
SuborderSuina
Suidae
Babyrousa
Hylochoerus
Phacochoerus
Porcula
Potamochoerus
Sus
Tayassuidae
Tayassu
Catagonus
Dicotyles
SuborderTylopoda
Camelidae
Lama
Camelus
SuborderWhippomorpha
Hippopotamidae
Hippopotamus
Choeropsis
Cetacea
Giraffidae
National
Other
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