Giraffidae

Taxonomy

Evolutionary background

The giraffids are ruminants of the clade Pecora. Other extant pecorans are the families Antilocapridae (pronghorns), Cervidae (deer), Moschidae (musk deer), and Bovidae (cattle, goats and sheep, wildebeests and allies, and antelopes). 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 the cladogram below.

The ancestors of pronghorn diverged from the giraffids in the Early Miocene. This was in part of a relatively late mammal diversification following a climate change that transformed subtropical woodlands into open savannah grasslands.

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

By the Pliocene, giraffids had been displaced from many Afro-Eurasian fossil assemblages by cervids and bovids, whose faster life history, contrasting with the long gestation periods of giraffids, may have provided an advantage at exploiting a greater diversity of niches than giraffids, particularly in the context of the cooling climate of the latter part of the Neogene, during which both cervids and bovids experienced rapid rates of evolution.

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

Classification

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

Family Giraffidae J.E.Gray, 1821

• Basal extinct giraffids
  • †Csakvarotherium Kretzoi, 1930
    • †Csakvarotherium hungaricum Kretzoi, 1930
  • †Injanatherium Heintz, Brunet & Sen, 1981
    • †Injanatherium arabicum Morales, Soria & Thomas, 1987
    • †Injanatherium hazimi Heintz, Brunet & Sen, 1981
  • †Propalaeomeryx Lydekker, 1883 [Progiraffa Pilgrim, 1908]
    • †Propalaeomeryx sivalensis Lydekker, 1883 [Progiraffa exigua Pilgrim, 1908]
  • †Shansitherium Killgus, 1922 [Schansitherium [sic]]
    • †Shansitherium quadricornis (Bohlin, 1926) [Palaeotragus quadricornis Bohlin, 1926]
    • †Shansitherium tafeli Killgus, 1922
  • †Umbrotherium Abbazzi, Delfino, Gallai, Trebini & Rook, 2008
    • †Umbrotherium azzarolii Abbazzi, Delfino, Gallai, Trebini & Rook, 2008
• Subfamily †Canthumerycinae Hamilton, 1978
  • †Georgiomeryx Paraskevaidis, 1940
    • †Georgiomeryx georgalasi Paraskevaidis, 1940
  • †Canthumeryx Hamilton 1973 [Zarafa Hamilton, 1973]
    • †Canthumeryx sirtensis Hamilton 1973 [Zarafa zelteni Hamilton, 1973]
• Subfamily †Progiraffinae Pilgrim, 1911
  • †Palaeogiraffa Bonis & Bouvrain, 2003
    • †Palaeogiraffa macedoniae (Geraads, 1989) [Decennatherium macedoniae Geraads, 1989]
    • †Palaeogiraffa major Bonis & Bouvrain, 2003
    • †Palaeogiraffa pamiri (Ozansoy, 1965) [Samotherium pamiri Ozansoy, 1965]
• Subfamily †Giraffokerycinae Solounias, 2007
  • †Giraffokeryx Pilgrim, 1910
    • †Giraffokeryx anatoliensis Geraads & Aslan, 2003
    • †Giraffokeryx primaevus (Churcher, 1970) [Palaeotragus primaevus Churcher, 1970; Samotherium africanum Churcher, 1970 and Amotherium africanum [sic]]
    • †Giraffokeryx punjabiensis Pilgrim, 1910
• Subfamily Giraffinae J.E.Gray, 1821
  • Tribe Giraffini J.E.Gray, 1821
    • †Orea Solounias & Ríos, 2025
      • †Orea leptia Solounias & Ríos, 2025
    • Giraffa Brisson, 1762 [Camelopardalis von Schreber, 1784 and Orasius Oken, 1816]
      • Giraffa giraffa (von Schreber, 1784)
        • Giraffa giraffa angolensis Lydekker, 1903 – Angolan giraffe
        • Giraffa giraffa giraffa (von Schreber, 1784) – South African giraffe
      • Giraffa tippelskirchii Matschie, 1898
        • Giraffa tippelskirchii thornicrofti Lydekker, 1911 – Thornicroft's giraffe
        • Giraffa tippelskirchii tippelskirchii Matschie, 1898 – Masai giraffe
      • Giraffa reticulata de Winton, 1899 – Reticulated giraffe
      • Giraffa camelopardalis (Linnaeus, 1758) – Northern giraffe
        • Giraffa camelopardalis peralta Thomas, 1898 – West African giraffe
        • Giraffa camelopardalis antiquorum Jardine & Swainson, 1835 – Kordofan giraffe
        • Giraffa camelopardalis camelopardalis (Linnaeus, 1758) – Nubian giraffe [Giraffa camelopardalis rothschildi Lydekker, 1903 – Rothschild's giraffe]
      • †Giraffa jumae Leakey, 1967
      • †Giraffa priscilla Pilgrim, 1911
      • †Giraffa punjabiensis Pilgrim, 1911
      • †Giraffa pygmaea Harris, 1976
      • †Giraffa sivalensis (Falconer & Cautley, 1843) [Camelopardalis sivalensis Falconer & Cautley, 1843 and Camelopardalis affinis Falconer & Cautley, 1843]
      • †Giraffa stillei (Dietrich, 1942) [Okapia stillei Dietrich, 1942 and Giraffa gracilis Arambourg, 1947]
  • Tribe †Bohlinini Solounias, 2007
    • †Honanotherium Bohlin, 1927
      • †Honanotherium bernori Solounias & Danowitz, 2016
      • †Honanotherium schlosseri (Pilgrim, 1911) [Giraffa schlosseri Pilgrim, 1911]
    • †Qilin Wang et al., 2025
      • †Qilin tungurensis (Colbert, 1936) [Palaeotragus tungurensis Colbert, 1936]
    • †Bohlinia Matthew, 1929
      • †Bohlinia adoumi Likius, Vignaud & Brunet, 2007
      • †Bohlinia attica (Gaudry & Lartet, 1856) [Giraffa attica (Gaudry & Lartet, 1856) and Orasius attica (Gaudry & Lartet, 1856)]
      • †Bohlinia nikitiae Kostopoulos, Koliadimou & Koufos, 1996
  • Tribe Palaeotragini Pilgrim, 1910
    • Subtribe †Palaeotragina Pilgrim, 1910
      • †Mitilanotherium Samson & Radulesco, 1966 [Macedonitherium Sickenberg, 1967; Sogdianotherium Sharapov, 1974]
        • †Mitilanotherium inexpectatum Samson & Radulesco, 1966[Macedonitherium martinii Sickenberg, 1967; Mitilanotherium inexpectatum Samson & Radulesco, 1966; Mitilanotherium kuruksaense (Sharapov, 1974); Mitilanotherium martinii (Sickenberg, 1967); Palaeotragus inexspectatus (Samson & Radulesco, 1966); Palaeotragus priasovicus Godina & Bajgusheva, 1985 and Sogdianotherium kuruksaense Sharapov, 1974]
      • †Palaeotragus Gaudry, 1861 [Achtiaria Borissiak, 1914]
        • †Palaeotragus coelophrys (Rodler & Weithofer, 1890) [Alcicephalus coelophrys Rodler & Weithofer, 1890]
        • †Palaeotragus germaini Arambourg, 1959
        • †Palaeotragus lavocanti Heintz, 1976
        • †Palaeotragus robinsoni Crusafont-Pairó, 1979
        • †Palaeotragus rouenii Gaudry, 1861 [Palaeotragus microdon Koken, 1885]
      • †Praepalaeotragus Godina, Vislobokova & Abdrachmanova, 1993
        • †Praepalaeotragus actaensis Godina, Vislobokova & Abdrachmanova, 1993
      • †Samotherium Forsyth Major, 1888 [Alcicephalus Rodler & Weithofer, 1890; Chersenotherium Alexajew, 1916 and Amotherium [sic]]
        • †Samotherium boissieri Forsyth Major, 1888
        • †Samotherium eminens (Alexajew, 1916) [Chersenotherium eminens Alexajew, 1916]
        • †Samotherium major Bohlin, 1926
        • †Samotherium neumayri (Rodler & Weithofer, 1890) [Alcicephalus neumayri Rodler & Weithofer, 1890]
        • †Samotherium sinense (Schlosser, 1903) [Alcicephalus sinense Schlosser, 1903]
    • Subtribe Okapiina Bohlin, 1926
      • †Afrikanokeryx Harris, Solounias & Geraads, 2010
        • †Afrikanokeryx leakeyi Harris, Solounias & Geraads, 2010
      • Okapia Lankester, 1901
        • Okapia johnstoni (P. L. Sclater, 1901) – Okapi
• †Subfamily Sivatheriinae Bonaparte, 1850
  • †Birgerbohlinia Crusafont Pairó, 1952
    • †Birgerbohlinia schaubi Crusafont Pairó, 1952
  • †Bramatherium Falconer, 1845 [Hydaspitherium Lydekker, 1876]
    • †Bramatherium giganteus Khan & Sarwar, 2002
    • †Bramatherium grande (Lydekker, 1878) [Hydaspitherium grande Lydekker, 1878]
    • †Bramatherium magnum (Pilgrim, 1910) [Hydaspitherium magnum Pilgrim, 1910]
    • †Bramatherium megacephalum (Lydekker, 1876) [Hydaspitherium megacephalum Lydekker, 1876]
    • †Bramatherium perimense Falconer, 1845
    • †Bramatherium progressus Khan, Sarwar & Khan, 1993
    • †Bramatherium suchovi Godina, 1977
  • †Decennatherium Crusafont Pairó, 1952
    • †Decennatherium rex Ríos, Sánchez & Morales, 2017
    • †Decennatherium pachecoi Crusafont Pairó, 1952
  • †Helladotherium Gaudry, 1860
    • †Helladotherium duvernoyi (Gaudry & Lartet, 1856) [Camelopardalis duvernoyi Gaudry & Lartet, 1856]
  • †Sivatherium Falconer & Cautley, 1836 [Griquatherium Haughton, 1922; Indratherium Pilgrim, 1910; Libytherium Pomel, 1892 and Orangiatherium van Hoepen, 1932]
    • †Sivatherium giganteum Falconer & Cautley, 1836
    • †Sivatherium hendeyi Harris, 1976
    • †Sivatherium maurusium (Pomel, 1892) [Libytherium maurusium Pomel, 1892; Griquatherium cingulatum Haughton, 1922; Helladotherium olduvaiense Hopwood, 1934; Sivatherium olduvaiense (Hopwood, 1934); Libytherium olduvaiense Hopwood, 1934 and Orangiatherium vanrhyni van Hoepen, 1932]
  • †Vishnutherium Lydekker, 1876
    • †Vishnutherium iravadicum Lydekker 1876