PBDB Taxon

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Camelidae (camel)

Mammalia - Camelidae

Taxonomy

Camelidae was named by Gray (1821). It is extant.

It was assigned to Hydrophorae by Gray (1821); to Pecora by Bonaparte (1850); to Artiodactyla by Cope (1889) and Lambe (1908); to Ruminantii by Jaekel (1911); to Ruminantia by Leidy (1873), Gregory (1910) and Thurmond and Jones (1981); to Cameloidea by Hay (1902), Scott (1940), Carroll (1988) and Prothero (1996); to Tylopoda by Dingus (1990) and Webb and Meachen (2004); to Tylopoda by Flower (1883) and Whistler and Webb (2005); to Artiodactyla by Honey (2007); to Tylopoda by Flower and Lydekker (1891), Matthew (1908) and Agnarsson and May-Collado (2008); to Camelidamorpha by Spaulding et al. (2009); and to Artiodactyla by Kurten and Anderson (1980), Nowak (1991), Thewissen et al. (2001), Hulbert and Whitmore (2006), Baskin and Thomas (2015) and Carbot-Chanona et al. (2023).

Synonyms

Poebrotheriidae was named by Cope (1874).

It was synonymized subjectively with Poebrotheriinae by Simpson (1945).

It was assigned to Ruminantia by Cope (1874); and to Tylopoda by Flower and Lydekker (1891).
Protolabididae was named by Cope (1887).

It was synonymized subjectively with Camelidae by McKenna and Bell (1997).

It was assigned to Artiodactyla by Cope (1889).
Eschatiidae was named by Cope (1887).

It was synonymized subjectively with Camelidae by McKenna and Bell (1997).

It was assigned to Artiodactyla by Cope (1889).
Aepycamelinae was named by Webb (1965). It was considered paraphyletic by Frailey (1978) and Honey et al. (1998).

It was considered an invalid subgroup of Camelidae by Frailey (1978) and Honey et al. (1998).

It was assigned to Camelidae by Webb (1965) and McKenna and Bell (1997).
Poebrotheriinae was named by Zittel (1893). Its type is Poebrotherium. It was considered paraphyletic by Honey et al. (1998).

It was considered an invalid subgroup of Camelidae by Honey et al. (1998).

It was assigned to Camelidae by Hay (1902), Simpson (1945) and Gazin (1955).
Gomphotherium was named by Cope (1886).

It was synonymized subjectively with Protomeryx by Matthew (1901) and Hay (1902); it was considered an invalid subgroup of Camelidae by Matthew (1904), McKenna (1966) and Prothero (1996).

It was assigned to Poebrotheriidae by Cope (1886).
Altomeryx was named by Frick (1929).

It was considered a nomen nudum by Simpson (1945) and Savage (1951).
Alticamelinae was named by Simpson (1945).

It was replaced with Aepycamelinae by Webb (1965).

It was assigned to Camelidae by Simpson (1945) and Maglio (1966).
Gomphotherium serus was named by Douglass (1899) [also said to be 1900]. It is a 3D body fossil.

It was recombined as Protomeryx serus by Hay (1902); it was considered a nomen dubium by Prothero (1996) and Tabrum and Nichols (2001).
Protomeryx was named by Leidy (1856). Its type is Protomeryx halli.

It was considered a nomen dubium by Loomis (1936); it was considered a nomen dubium by McKenna (1966) and Prothero (1996).

It was assigned to Camelidae by Matthew (1901); to Poebrotheriinae by Hay (1902); and to Camelinae by Simpson (1945).

Synonymy list

Year	Name and author
1821	Camelidae Gray p. 307
1850	Camelidae Bonaparte p. 1
1856	Protomeryx Leidy
1873	Camelidae Leidy p. 317
1874	Poebrotheriidae Cope p. 26
1883	Camelidae Flower p. 184
1886	Gomphotherium Cope p. 619 fig. 10
1887	Eschatiidae Cope
1887	Protolabididae Cope
1889	Camelidae Cope p. 877
1889	Eschatiidae Cope p. 877
1889	Protolabididae Cope p. 877
1891	Camelidae Flower and Lydekker p. 90
1891	Poebrotheriidae Flower and Lydekker p. 90
1893	Poebrotheriinae Zittel
1899	Gomphotherium serus Douglass p. 12 figs. Plate 1
1901	Protomeryx Matthew p. 422
1902	Camelidae Hay p. 673
1902	Poebrotheriinae Hay p. 675
1902	Protomeryx Hay p. 676
1902	Protomeryx serus Hay p. 676
1908	Camelidae Lambe
1908	Camelidae Matthew p. 561
1910	Camelidae Gregory p. 466
1911	Camelidae Jaekel p. 234
1929	Altomeryx Frick p. 107
1940	Camelidae Scott p. 604
1945	Alticamelinae Simpson p. 150
1945	Poebrotheriinae Simpson p. 150
1945	Protomeryx Simpson p. 150
1955	Poebrotheriinae Gazin
1965	Aepycamelinae Webb p. 37
1966	Alticamelinae Maglio p. 11
1980	Camelidae Kurten and Anderson p. 301
1981	Camelidae Thurmond and Jones p. 188
1988	Camelidae Carroll
1990	Camelidae Dingus p. 13
1991	Camelidae Nowak
1996	Camelidae Prothero p. 610
1997	Aepycamelinae McKenna and Bell
2001	Camelidae Thewissen et al. p. suppl.
2004	Camelidae Webb and Meachen p. 350
2005	Camelidae Whistler and Webb p. 5
2006	Camelidae Hulbert and Whitmore p. 6
2007	Camelidae Honey
2008	Camelidae Agnarsson and May-Collado p. 974 figs. Fig. 2
2009	Camelidae Spaulding et al.
2015	Camelidae Baskin and Thomas p. 119
2023	Camelidae Carbot-Chanona et al. p. 4

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
subclass	Synapsida
—	Therapsida	()

Rank	Name	Author
infraorder	Cynodontia	()
—	Mammaliamorpha
—	Mammaliaformes
class	Mammalia
—	Cladotheria
—	Zatheria
subclass	Tribosphenida	()
subclass	Theria
—	Eutheria	()
—	Placentalia
—	Boreoeutheria
—	Laurasiatheria
—	Scrotifera
—	Euungulata
—	Artiodactylamorpha
—	Artiodactyla	()
suborder	Tylopoda	()
family	Camelidae
family	Camelidae

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Fm. Camelidae Gray 1821 [camel]

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G. †Australocamelus Patton 1969

†Australocamelus orarius Patton 1969

Subfm. Camelinae Gray 1821

G. †Alforjas Harrison 1979

†Alforjas taylori Harrison 1979

Invalid names: Pliauchenia hemphillensis Dalquest 1980 [synonym]

Tr. Camelini Webb 1965

G. †Camelops Leidy 1854

†Camelops hesternus Leidy 1873

†Camelops huerfanensis dallasi Lull 1921

Invalid names: Camelops aransas Hay 1926 [synonym], Camelops huerfanensis Cragin 1892 [synonym], Camelops sulcatus Cope 1893 [synonym], Camelops traviswhitei Mooser and Dalquest 1975 [synonym]

†Camelops kansanus Leidy 1854

†Camelops minidokae Hay 1927

Invalid names: Camelops arenarum Hay 1927 [nomen dubium], Camelops maximus Hay 1921 [nomen dubium], Camelops nevadanus Hay 1927 [nomen dubium], Camelops niobrarensis Leidy 1858 [nomen vanum], Camelops nitidus Hay 1920 [nomen dubium], Camelops vacondae Hay 1927 [nomen dubium], Eschatius Cope 1884 [nomen dubium], Eschatius conidens Cope 1884 [nomen dubium], Megalomeryx Leidy 1858 [synonym]

G. Camelus Linnaeus 1758 [camel]

†Camelus antiquus Falconer and Cautley 1836

†Camelus arctoamericanus Hay 1921

Camelus bactrianus Linnaeus 1758

Camelus dromedarius Linnaeus 1758

†Camelus grattardi Geraads 2014

†Camelus knoblochi Nehring 1901

†Camelus sivalensis Falconer and Cautley 1836

†Camelus thomasi Pomel 1893

G. †Gigantocamelus Barbour and Schultz 1939

†Gigantocamelus spatulus Cope 1893

Invalid names: Gigantocamelus fricki Barbour and Schultz 1939 [synonym]

G. †Megacamelus Frick 1929

†Megacamelus fricki Frick 1929

†Megacamelus merriami Frick 1921

G. †Megatylopus Matthew and Cook 1909

†Megatylopus cochrani Hibbard and Riggs 1949

†Megatylopus gigas Matthew and Cook 1909

Invalid names: Paracamelus brevirostris Macdonald 1959 [synonym]

†Megatylopus matthewi Webb 1965

†Megatylopus primaevus Patton 1969

G. †Paracamelus Schlosser 1903

†Paracamelus agguirrei Morales 1984

†Paracamelus alexejevi Khavesson 1950

†Paracamelus alutensis Stefanescu 1895

†Paracamelus gigas Schlosser 1903

†Paracamelus khersonensis Pavlow 1903

Invalid names: Paracamelus arenicola Reed and Longnecker 1932 [nomen nudum]

G. †Titanotylopus Barbour and Schultz 1934

†Titanotylopus nebraskensis Barbour and Schultz 1934

G. †Hesperocamelus Macdonald 1949

†Hesperocamelus stylodon Macdonald 1949

Tr. Lamini Webb 1965

G. †Aepycamelus Macdonald 1956

†Aepycamelus alexandrae Davidson 1923

†Aepycamelus bradyi Macdonald 1956

†Aepycamelus elrodi Douglass 1909

†Aepycamelus giraffinus Matthew 1909

†Aepycamelus major Leidy 1886

†Aepycamelus priscus Matthew and Cook 1909

†Aepycamelus procerus Matthew and Cook 1909

†Aepycamelus robustus Leidy 1858

Invalid names: Homocamelus caninus Leidy 1869 [synonym], Procamelus madisonius Douglass 1899 [synonym], Protolabis prehensilis Cope 1881 [synonym]

†Aepycamelus stocki Henshaw 1942

Invalid names: Alticamelus Matthew 1901 [synonym]

G. †Blancocamelus Dalquest 1975

†Blancocamelus meadei Dalquest 1975

G. †Eulamaops Ameghino 1889

†Eulamaops parallelus Ameghino 1884

G. †Hemiauchenia Gervais and Ameghino 1880

†Hemiauchenia blancoensis Meade 1945

†Hemiauchenia edensis Frick 1921

†Hemiauchenia gracilis Meachen 2005

†Hemiauchenia guanajuatensis Dalquest and Mooser 1980

†Hemiauchenia macrocephala Cope 1893

Invalid names: Camelus americanus Wortman 1898 [synonym], Lama hollomani Hay and Cook 1930 [synonym], Lama stevensi Merriam and Stock 1925 [synonym], Tanupolama seymourensis Hibbard and Dalquest 1962 [synonym]

†Hemiauchenia minima Leidy 1886

†Hemiauchenia paradoxa Gervais and Ameghino 1880

Invalid names: Tanupolama Stock 1928 [synonym]

G. Lama Cuvier 1800

Subg. Lama (Lama) Menegaz et al. 1989

†Lama castelnaudi Gervais 1855

Lama glama Linnaeus 1758

†Lama gracilis Gervais and Ameghino 1980

Lama guanicoe Müller 1776 [guanaco]

†Lama intermedia Gervais 1855

†Lama mesolithica Ameghino 1889

Lama pacos Linnaeus 1758

Invalid names: Auchenia Illiger 1811 [synonym]

G. †Palaeolama Gervais 1869

Subg. †Palaeolama (Astylolama) Churcher 1965

†Palaeolama (Astylolama) aequatorialis Hoffstetter 1952

Subg. †Palaeolama (Palaeolama) Churcher 1965

†Palaeolama (Palaeolama) paradoxa Cabrera 1935

†Palaeolama (Palaeolama) weddellii Gervais 1855

Subg. †Palaeolama (Protauchenia) Branco 1883

†Palaeolama (Protauchenia) reissi Branco 1883

†Palaeolama brevirostris Rusconi 1930

†Palaeolama castelnaudi Gervais 1854

†Palaeolama crassa Hoffstetter 1952

†Palaeolama crequii Boule and Thevenin 1920

†Palaeolama leptognata Ameghino 1889

†Palaeolama major Liais 1872

†Palaeolama mirifica Simpson 1929 [stout-legged llama]

†Palaeolama niedae Guérin and Faure 1999

†Palaeolama promesolithica Ameghino 1889

†Palaeolama weddeli Gervais 1855

†Palaeolama weddelli Gervais 1855

G. †Pleiolama Webb and Meachen 2004

†Pleiolama mckennai Webb and Meachen 2004

†Pleiolama vera Matthew 1909

G. †Pliauchenia Cope 1875

Subg. †Pliauchenia (Megatylopus) Matthew and Cook 1909

†Pliauchenia humphreysiana Cope 1875

†Pliauchenia magnifontis Gregory 1939

†Pliauchenia minima Wortman 1898

†Pliauchenia raki Frick 1921

†Pliauchenia vulcanorum Cope 1875

G. †Procamelus Leidy 1858

†Procamelus coconiensis Hay 1921

†Procamelus coconinensis Hay 1921

†Procamelus grandis Gregory 1939

Invalid names: Procamelus dakotensis Green 1956 [synonym]

†Procamelus leptocolon Matthew 1909

†Procamelus leptognathus Cope 1893

†Procamelus occidentalis Leidy 1858

Invalid names: Procamelus lacustris Douglass 1899 [synonym]

†Procamelus virginiensis Leidy 1873

Invalid names: Homocamelus Leidy 1869 [synonym]

G. Vicugna Lesson 1842

Lama (Vicugna) gracilis Gervais and Ameghino 1881

Invalid names: Auchenia frontosa Gervais and Ameghino 1880 [synonym], Auchenia lujanensis Ameghino 1889 [synonym], Auchenia minuta Burmeister 1891 [synonym], Hemiauchenia pristina Ameghino 1891 [synonym]

Vicugna vicugna Molina 1782

Invalid names: Vicugna provicugna Hoffstetter 1963 [synonym]

Invalid names: Tanupolama parvus Olson 1940 [nomen dubium]

Invalid names: Camelopini Webb 1965 [empty], Procamelus edensis raki Frick 1921 [nomen vanum], Procamelus minor Leidy 1886 [nomen dubium], Prochenia [nomen nudum], Protolabidini Webb 1965 [empty]

Subfm. †Floridatragulinae Maglio 1966

G. †Aguascalientia Stevens 1977

†Aguascalientia minuta Rincon et al. 2012

†Aguascalientia panamaensis Rincon et al. 2012

†Aguascalientia wilsoni Dalquest and Mooser 1974

G. †Floridatragulus White 1940

†Floridatragulus dolichanthereus White 1940

Invalid names: Floridatragulus barbouri White 1947 [synonym], Hypermekops olseni White 1942 [synonym]

†Floridatragulus nanus Patton 1969

†Floridatragulus texanus Patton 1969

Invalid names: Floridatragulus hesperus Patton 1969 [synonym]

Invalid names: Hypermekops White 1942 [synonym]

G. †Stevenscamelus Prothero et al. 2023

†Stevenscamelus franki Wilson 1974

G. †Matthewlabis Prothero 2011

†Matthewlabis cedrensis Matthew 1901

Invalid names: Protomeryx campester Matthew 1901 [synonym], Pseudolabis (Paralabis) matthewi Lull 1921 [synonym]

Invalid names: Paralabis Lull 1921 [replaced]

Subfm. †Miolabinae Hay 1902

G. †Capricamelus Whistler and Webb 2005

†Capricamelus gettyi Whistler and Webb 2005

G. †Cuyamacamelus Kelly 1992

†Cuyamacamelus jamesi Kelly 1992

G. †Miolabis Hay 1899

Subg. †Miolabis (Paratylopus) Matthew 1904

†Miolabis (Paratylopus) primaevus Matthew 1904

†Miolabis californicus Maxson 1930

†Miolabis fissidens Cope 1876

Invalid names: Protolabis princetonianus Sinclair 1915 [synonym]

†Miolabis fricki Kelly 1992

†Miolabis longiceps Matthew 1909

†Miolabis montanus Douglass 1899

†Miolabis transmontanus Cope 1879

G. †Nothotylopus Patton 1969

†Nothotylopus camptognathus Patton 1969

G. †Paramiolabis Kelly 1992

†Paramiolabis minutus Pagnac 2005

†Paramiolabis singularis Matthew 1918

†Paramiolabis taylori Kelly 1992

†Paramiolabis tenuis Matthew 1924

Subfm. †Nothokemadinae White 1947

G. †Gentilicamelus Loomis 1936

†Gentilicamelus cameloides Wortman 1898

†Gentilicamelus sternbergi Cope 1879

G. †Nothokemas White 1947

†Nothokemas floridanus Simpson 1932

Invalid names: Paratylopus grandis White 1940 [synonym]

†Nothokemas hidalgensis Patton 1969

†Nothokemas waldropi Frailey 1978

G. †Oxydactylus Peterson 1904

†Oxydactylus benedentatus Hay 1924

Invalid names: Procamelus concerptus Hay 1924 [synonym], Protolabis francisi Hay 1924 [synonym]

†Oxydactylus campestris Cook 1909

†Oxydactylus lacota Matthew and Macdonald 1960

†Oxydactylus longipes Peterson 1904

†Oxydactylus lulli Loomis 1911

†Oxydactylus wyomingensis Loomis 1936

G. †Palauchenia del Castillo 1869

†Palauchenia mexicana del Castillo 1869

Invalid names: Palauchenia magna Owen 1870 [synonym]

G. †Paratylopus Matthew 1904

†Paratylopus labiatus Cope 1881

†Paratylopus matthewi Lull 1921

†Paratylopus primaevus Matthew 1904

†Paratylopus wortmani Lull 1921

Invalid names: Protomeryx halli Leidy 1856 [nomen dubium]

G. †Poebrodon Gazin 1955

†Poebrodon californicus Golz 1976

†Poebrodon kayi Gazin 1955

G. †Poebrotherium Leidy 1847

†Poebrotherium chadronense Prothero 1996

†Poebrotherium eximium Hay 1902

†Poebrotherium wilsoni Leidy 1847

Invalid names: Poebrotherium andersoni Troxell 1917 [synonym]

G. †Priscocamelus Stevens et al. 1969

†Priscocamelus wilsoni Stevens et al. 1969

Subfm. †Protolabinae Zittel 1893

G. †Michenia Frick and Taylor 1971

†Michenia agatensis Frick and Taylor 1971

†Michenia deschutensis Dingus 1990

†Michenia exilis Matthew and Macdonald 1960

Invalid names: Michenia australis Stevens 1977 [synonym]

†Michenia mudhillsensis Pagnac 2005

G. †Protolabis Cope 1876

†Protolabis angustidens Cope 1874

†Protolabis barstowensis Lewis 1968

†Protolabis coartatus Stirton 1929

Invalid names: Protolabis notiochorinos Patton 1969 [synonym]

†Protolabis gracilis Leidy 1858

†Protolabis heterodontus Cope 1874

Invalid names: Procamelus foxi Macdonald 1960 [synonym]

†Protolabis inaequidens Cope and Matthew 1915

†Protolabis saxeus Matthew 1924

†Protolabis yavapaiensis Honey and Taylor 1978

G. †Tanymykter Honey and Taylor 1978

†Tanymykter brachyodontus Peterson 1904

†Tanymykter longirostris Peterson 1911

Subfm. †Stenomylinae Matthew 1910

G. †Miotylopus Schlaikjer 1935

†Miotylopus gibbi Loomis 1911

Invalid names: Dyseotylopus migrans Stock 1935 [synonym], Miotylopus bathygnathus Schlaikjer 1935 [synonym]

†Miotylopus leonardi Loomis 1911

†Miotylopus taylori Prothero 1996

Invalid names: Dyseotylopus Stock 1935 [synonym]

G. †Pseudolabis Matthew 1904

Subg. †Pseudolabis (Paralabis) Lull 1921

†Pseudolabis dakotensis Matthew 1904

Tr. †Stenomylini Prothero 1996

G. †Blickomylus Frick and Taylor 1968

†Blickomylus galushai Frick and Taylor 1968

†Blickomylus moroni Constenius and Dawson 2008

G. †Rakomylus Frick 1937

†Rakomylus raki Frick 1937

G. †Stenomylus Peterson 1907

†Stenomylus crassipes Loomis 1910

†Stenomylus gracilis Peterson 1907

†Stenomylus hitchcocki Loomis 1910

†Stenomylus keelinensis Frick and Taylor 1968

†Stenomylus taylori Cassiliano 2008

†Stenomylus tubutamensis Ferrusquia-Villafranca 1990

Invalid names: Stenomylus (Pegomylus) Frick and Taylor 1968 [invalid subgroup], Stenomylus (Stenomylus) Frick and Taylor 1968 [invalid subgroup]

G. †Wyomylus Cassiliano 2008

†Wyomylus whitei Cassiliano 2008

Invalid names: Pseudolabidinae Simpson 1945 [synonym]

G. †Stilauchenia Ameghino 1889

†Stilauchenia owenii Gervais and Ameghino 1880

G. †Teleopternus Cope 1899

†Teleopternus orientalis Cope 1899

Invalid names: Aepycamelinae Webb 1965 [invalid subgroup], Altomeryx Frick 1929 [nomen nudum], Aucheniina Bonaparte 1850 [empty], Camelina Gray 1825 [empty], Eschatiidae Cope 1887 [synonym], Gomphotherium Cope 1886 [invalid subgroup], Gomphotherium serus Douglass 1899 [nomen dubium], Merycotheriina Bonaparte 1850 [empty], Nothokematinae Prothero 1996 [empty], Poebrotheriinae Zittel 1893 [invalid subgroup], Protolabididae Cope 1887 [synonym], Protomeryx Leidy 1856 [nomen dubium]

Diagnosis

Reference	Diagnosis
D. R. Prothero 1996	he Family Camelidae is united by the following shared characters that distinguish it from the remaining Tylopoda, and all other artiodactyls: fully selenodont, transversely compressed upper molars that are relatively high- crowned (compared to other primitive Tylopoda), with straight ectolophs, and with fossettes that are closed anteriorly and posteriorly after moderate wear. Relatively long, narrow rostrum. Postorbital processes well developed, and nearly or completely closed. Inflated tympanic bulla filled with cancellous bone and indented by a deep tympanohyal groove. Angular process on mandible with a distinctive dorsal "hook." Long, unfused to fused, distally divergent middle metapodials with metacarpals II and IV and metatarsals II and V reduced to nodules. Metatarsals III and IV have flattened dorsal surfaces and their distal keels do not extend to the dorsal side. Fibular facet on the calcaneum has a proximal convexity and a dorsal concavity.