Skip to main content

The Camelidae (Mammalia, Artiodactyla) from the Quaternary of South America: Cladistic and Biogeographic Hypotheses

Journal of Mammalian Evolution volume 20, pages 45–56 (2013)Cite this article

Abstract

Presented here is a cladistic analysis of the South American and some North American Camelidae. This analysis shows that Camelini and Lamini are monophyletic groups, as are the genera Palaeolama and Vicugna, while Hemiauchenia and Lama are paraphyletic. Some aspects of the migration and distribution of South American camelids are also discussed, confirming in part the propositions of other authors. According to the cladistic analysis and previous propositions, it is possible to infer that two Camelidae migration events occurred in America. In the first one, Hemiauchenia arrived in South America and, this was related to the speciation processes that originated Lama and Vicugna. In the second event, Palaeolama migrated from North America to the northern portion of South America. It is evident that there is a need for larger studies about fossil Camelidae, mainly regarding older ages and from the South American austral region. This is important to better undertand the geographic and temporal distribution of Camelidae and, thus, the biogeographic aspects after the Great American Biotic Interchange.

This is a preview of subscription content, access via your institution.

Access options

Buy single article

Instant access to the full article PDF.

USD 39.95

Price includes VAT (USA)
Tax calculation will be finalised during checkout.

Fig. 1
Fig. 2

References

  • Ameghino F (1884) Escursiones geológicas y paleontológicas en la Província de Buenos Aires. Bol Acad Cienc Córdoba 6:161–257

    Google Scholar 

  • Ameghino F (1889) Contribución al conocimiento de los mamíferos fósiles de la República Argentina. Actas Acad Nac Cienc Córdoba 6:1–1027

    Google Scholar 

  • Boule M, Thevenin A (1920) Mamiféres fossiles de Tarija (Mission Scientifique de Crequi-Montfort et Sénèchal de la Grange VII). Soudier, Paris

    Google Scholar 

  • Branco W (1883) Ueber eine Fossile Säugethier-Fauna Von Punin bei Riobamba in Ecuador II - Beschreibung der Fauna. Pälaontol Abh 1:57–204

    Google Scholar 

  • Breyer J (1976) Titanotylopus (=Gigantocamelus) from the Great Plains Cenozoic. J Paleontol 50:783–788

    Google Scholar 

  • Cabrera A (1931) Sobre los camélidos fósiles y actuales de la América austral. Rev Mus La Plata 33:89–117

    Google Scholar 

  • Cabrera A (1935) Sobre la osteologia de Palaeolama. An Mus Argent Cienc Nat “Bernardino Rivadavia” 66:283–312

  • Casamiquela R (1999) The Pleistocene vertebrate record of Chile. In: Rabassa J, Salemme M (eds) Quaternary of South America and Antarctic Peninsula. AA Balkema Publishers, Rotterdam, pp 91–107

    Google Scholar 

  • Castellanos A (1944) Paleontología estratigráfica de los sedimentos neógenos de la provincia de Córdoba. Publ Inst Fisiogr Geol Rosario 23:3–47

    Google Scholar 

  • Churcher CS (1965) Camelid material of the genus Palaeolama Gervais from the Talara tar-seeps, Peru, with description of a new subgenus Astylolama. Proc Zool Soc Lond 145:161–205

    Google Scholar 

  • Cione AL, Tonni EP (1995a) Chronostratigraphy and “Land-Mammal Ages” in the Cenozoic of southern South America: principles, practices and the “Uquian” problem. J Paleontol 69:135–159

    Google Scholar 

  • Cione AL, Tonni EP (1995b) Bioestratigrafía e cronología del Cenozóico Superior de la región pampeana. In: Alberdi MT, Leone G, Tonni EP (eds) Evolución biológica y climática de la región pampeana durante los últimos cinco millones de años. Un ensayo de correlación con el Mediterráneo occidental. CSIC, Madrid, pp 49–74

    Google Scholar 

  • Cione AL, Tonni EP (1999) Biostratigraphy and chronological scale of upper-most Cenozoic in the Pampean Area, Argentina. In: Rabassa J, Salemme M (eds) Quaternary of South America and Antarctic Peninsula. AA Balkema Publishers, Rotterdam, pp 23–51

    Google Scholar 

  • Cione AL, Tonni EP (2005) Bioestratigrafía basada en mamíferos del Cenozoico Superior de la Provincia de Buenos Aires, Argentina. In: Barrio RE, Etcheverry RO, Caballé MF, Llambias E (eds) Geología y recursos Minerales de la Provincia de Buenos Aires. Relatório del XVI Congreso geológico Argentino. Quick Press, La Plata, pp 183–200

    Google Scholar 

  • Cione AL, Tonni EP, Bond M, Carlini A, Pardiñas UFJ, Scillato-Yané G, Verzi D, Vucetich MG (1999) Ocurrence charts of Pleistocene Mammals in the Pampean area, eastern Argentina. In: Rabassa J, Salemme M (eds) Quaternary of South America and Antartic Peninsula. AA Balkema Publishers, Rotterdam, pp 53–59

    Google Scholar 

  • Coltorti M, Abbazzi L, Ferretti MP, Iacumin P, Paredes-Rios F, Pellegrini M, Pieruccini P, Rustioni M, Tito G, Rook L (2007) Last Glacial mammals in South America: a new scenario from Tarija (Bolivia). Naturwissenschaften 94:288–299

    PubMed  Article  CAS  Google Scholar 

  • Dalquest WW, Mooser O (1980) Late Hemphillian mammals of the Ocote local fauna, Guanajuato, Mexico. Pearce-Sellards Series 32:1–25

    Google Scholar 

  • Deraco MV, Scherer CS, Powell JE (2006) Sobre Hemiauchenia paradoxa del Pleistoceno del Río Dulce, Provincia de Santiago del Estero, Argentina. Paleontologia em Destaque 57: 47–48

    Google Scholar 

  • Deschamps CM (2005) Late Cenozoic mammal bio-chronostratigraphy in southwestern Buenos Aires Province, Argentina. Ameghiniana 42:733–750

    Google Scholar 

  • Eisenberg JF, Redford KH (1999) Mammals of the Neotropics: The Central Neotropics. The University of Chicago Press, Chicago

    Google Scholar 

  • Ferrero BS (2006) Avances en el conocimiento de la diversidad de artiodáctilos en el Pleistoceno de la Provincia de Entre Ríos, Argentina. Ameghiniana 43:38R

    Google Scholar 

  • Frenguelli J (1936) La serie geológica de la República Argentina en sus relaciones con la antigüedad del hombre. Hist Nac Arg 1:106–113

    Google Scholar 

  • Gervais H, Ameghino F (1880) Les mammifères fósiles de l’Amerique du Sud. F. Savy, Paris and Buenos Aires

    Google Scholar 

  • Gervais P (1855) Recherches sur les mammifères fósiles de l’Amérique Meridionale (Expédition dans les parties centrales de l’Amérique du Sud, de Rio de Janeiro à Lima et de Lima au Pará, exécutée par ordre du Gouvernement français pendant les années 1843 à 1847 sous la direction de F. Castelnau). Zoologie 7:1–63

    Google Scholar 

  • Goloboff P (1999) NONA ver. 2.0 Program and Documentation. Published by author, Tucuman

  • Harrison JA (1979) Revision of the Camelinae (Artiodactyla, Tylopoda) and description of the new genus Alforjas. Univ Kansas Paleontol Contrib 95:1–20

    Google Scholar 

  • Harrison JA (1985) Giant camels from the Cenozoic of North America. Smithsonian Contrib Paleobiol 57:2–39

    Google Scholar 

  • Hoffstetter R (1952) Les mammiféres pléistocènes de la République de l’Equateur. Mém Soc Géol France (Nouvelle série) 66:1–391

    Google Scholar 

  • Honey JG, Harrison JA, Prothero DR, Stevens MS (1998) Camelidae. In: Janis CM, Scott KM, Jacobs LL (eds) Evolution of Tertiary Mammals of North America, vol 1. Cambridge University Press, New York, pp 439–462

    Google Scholar 

  • Honey JG, Taylor BE (1978) A generic revision of the Protolabidini (Mammalia, Camelidae), with a description of two new Protolabidines. Bull Am Mus Nat Hist 161:367–426

    Google Scholar 

  • Kraglievich L (1946) Sobre Camélidos Chapadmalenses. Not Mus La Plata Paleontol 93:317–331

    Google Scholar 

  • Kraglievich L (1959) Contribuciones al conocimiento de la geología cuaternaria en la Argentina. IV: Nota acerca de la geología costera en la desembocadura del Arroyo Malacara (Prov. de Buenos Aires). An Mus Argent Cienc Nat "Bernardino Rivadavia" 17:3–9

  • Labarca RO, López PG (2006) Los mamíferos finipleistocénicos de la Formación Quebrada Quereo (IV Región - Chile): biogeografía, bioestratigrafía y inferencias paleoambientales. Mastozool Neotr 13:89–101

    Google Scholar 

  • Lesbre FX (1903) Recherches anatomiques sur lês Camélidés. Arch Mus D´Hist Natur Lyon 8:1–196

    Google Scholar 

  • López PM, Martínez IR, Labarca RE, Jackson DS (2005) Registro de Hippidion, Canidae, Camelidae y Edentata en el Sector de Ojo de Opache, Calama, II Región. Notic Mens Mus Nac Hist Nat 355:22–26

    Google Scholar 

  • López-Aranguren DJ (1930) Camélidos fósiles argentinos. Anal Soc Cien Arg 109:15–39, 97–126

    Google Scholar 

  • Lund PW (1837) Segunda Memória sobre a fauna das cavernas. In: Paula-Couto C (ed) Memórias sobre a Paleontologia Brasileira (Revistas e comentadas por Carlos de Paula Couto). Ministério da Educação e Saúde, Instituto Nacional do Livro, Rio de Janeiro, pp 131–203

    Google Scholar 

  • MacFadden BJ (2000) Middle Pleistocene climate change recorded in fossil mammal teeth from Tarija, Bolivia, and upper limit of the Ensenadan Land-Mammal Age. Quaternary Res 54:121–131

    Article  CAS  Google Scholar 

  • MacFadden BJ, Siles O, Zeitler P, Johnson NM, Campbell Jr KE (1983) Magnetic polarity stratigraphy of the middle Pleistocene (Ensenadan) Tarija Formation of southern Bolivia. Quaternary Res 19:172–187

    Article  Google Scholar 

  • Marshall LG, Berta A, Hoffstetter R, Pascual R, Reig OA, Bombin M, Mones A (1984) Mammals and stratigraphy: geochronology of the continental mammal-bearing Quaternary of South America. Palaeovertebrata Mém Estraord:1–76

  • Matthew WD, Cook HJA (1909) Pliocene fauna from western Nebraska. Bull Am Mus Nat Hist 26:361–414

    Google Scholar 

  • McKenna MC, Bell SK (1997) Classification of Mammals Above the Species Level. Columbia University Press, New York

    Google Scholar 

  • Menegaz AN, Goin FJ, Ortiz-Jaureguizar E (1989) Análisis morfológico e morfométrico multivariado de los representantes fósiles y vivientes del género Lama (Artiodactyla, Camelidae). Sus implicancias sistemáticas, biogeográficas, ecológicas y biocronológicas. Ameghiniana 26:153–172

    Google Scholar 

  • Mones A (1988) Nuevos registros de mamíferos fósiles de la Formación San José (Plioceno-?Pleistoceno Inferior) (Mammalia: Xenarthra, Artiodactyla, Rodentia). Commun Paleontol Mus Hist Nat 20:255–277

    Google Scholar 

  • Montellano M (1989) Pliocene Camelidae of Rancho El Ocote, Central Mexico. J Mammal 70:359–369

    Article  Google Scholar 

  • Nixon KC (1999–2002) WinClada ver. 1.0000. Published by author, Ithaca

  • Olsen ST (1988) The camel in ancient China and an osteology of the camel. Proc Acad Nat Sci Phil 140:18–58

    Google Scholar 

  • Ruez DR (2005) Earliest record of Palaeolama (Mammalia, Camelidae) with comments on “Palaeolama” guanajuatensis. J Vertebr Paleontol 25:741–744

    Article  Google Scholar 

  • Rusconi C (1933) Nuevas especies de mamíferos terciarios procedentes del Piso Chapadmalense (Plioceno medio). Anal Soc Cient Arg 115:1–101

    Google Scholar 

  • Salas R, Stucchi M, Devries TJ (2003) The presence of Plio-Pleistocene Palaeolama sp. (Artiodactyla, Camelidae) on the southern coast of Peru. Bull Inst Français Ét Andines 32:347–359

    Google Scholar 

  • Scherer CS, Ferigolo J, Ribeiro AM, Cartelle C (2007) Contribution to the knowledge of Hemiauchenia paradoxa (Artiodactyla, Camelidae) from the Pleistocene of southern Brazil. Rev Bras Paleontol 10:35–52

    Article  Google Scholar 

  • Socorro OAA (2006) Tesoros paleontológicos de Venezuela, el Cuaternario del Estado Falcón. Instituto del Patrimonio Cultural, Taima Taima

    Google Scholar 

  • Tauber AA (1999) Hallazgo de uma vicuña em el Pleistoceno de la Província de Córdoba, República Argentina. Ameghiniana 36:55–62

    Google Scholar 

  • Tonni EP, Politis GG (1980) La distribución del guanaco (Mammalia, Camelidae) en la Província de Buenos Aires durante el Pleistoceno Tardio y Holoceno. Los factores climáticos como causas de su retracción. Ameghiniana 1:53–66

    Google Scholar 

  • Ubilla M, Perea D (1999) Quaternary vertebrates of Uruguay: a biostratigraphic and climatic overview. In: Rabassa J, Salemme M (eds) Quaternary of South America and Antarctic Peninsula. AA Balkema Publishers, Rotterdam, pp 75–89

    Google Scholar 

  • Webb SD (1965) The osteology of Camelops. Bull Los Angeles County Mus Sci 1:1–54

    Google Scholar 

  • Webb SD (1972) Locomotor evolution in camels. Forma et Functio 5:99–111

    Google Scholar 

  • Webb SD (1974) Pleistocene llamas of Florida with a brief review of the Lamini. In: Webb SD (ed) Pleistocene Mammals of Florida. University of Florida Press, Gainsville, pp 170–213

    Google Scholar 

  • Webb SD (1985) Late Cenozoic mammal dispersals between the Americas. In: Stehli FG, Webb SD (eds) The Great American Biotic Interchange. Plenum Press, New York, pp 357–386

    Chapter  Google Scholar 

  • Webb SD, Meachen J (2004) On the origin of lamine Camelidae including a new genus from the late Miocene of the High Plains. Bull Carnegie Mus Nat Hist 36:349–362

    Article  Google Scholar 

  • Webb SD, Stehli FG (1995) Selenodont Artiodactyla (Camelidae and Cervidae) from the Leisey Shell Pits, Hills Borough County, Florida. Bull Florida Mus Nat Hist 37:621–643

    Google Scholar 

  • Woodburne MO (2010) The Great American Biotic Interchange: dispersals, tectonics, climate, sea level and holding pens. J Mammal Evol 17: 245–264

    PubMed  Article  Google Scholar 

Download references

Acknowledgements

I thank Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the financial grant and the Florida Museum of Natural History for the International Travel Grant. Also, UFRGS and FZBRS for the physical support to develop this work; to J. Ferigolo, A.M. Ribeiro, and T.V. Oliveira for their critical reading and help. To the follow persons and institutions for permission to access the respective fossil and extant collections: B. MacFadden and R. Hulbert (FLMNH/UF), M. Reguero and I. Olivares (MLP), A. Kramarz (MACN), A. Dondas (MMCNLS), B. Mamani and J. Vargas (MNHN), F. Paredes (MNPA), J.L. Roman-Carrion (EPN), M. Trindade (MACN, Salto), J. Solovyi (MHD), N. Guidon (FUMDHAM), C. Cartelle (PUCMG), J. Ferigolo and M. Jardim (FZBRS), K. Aaris (ZMUC), UFAC and DNPM. I thank also L. Avilla and J. Wible for the opportunity participation of this volume and the anonymous referees of this paper for their important contributions.

Author information

Authors and Affiliations

  1. Universidade Federal do Recôncavo da Bahia, Centro de Ciências Agrárias, Ambientais e Biológicas, Rua Rui Barbosa 710, Campus Universitário, Centro, Cruz das Almas, Bahia, 44.380-000, Brazil

    Carolina Saldanha Scherer

Authors
  1. Carolina Saldanha Scherer
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Carolina Saldanha Scherer.

Appendices

Appendix I. Characters Used in Cladistic Analysis

After the character states, inside parentheses, the papers where the characters were previously applied in cladistic analysis are represented. Those followed by “mod” were modified in this work. The remaining characters were described by other authors (e.g., López-Aranguren 1930; Cabrera 1931, 1935; Hoffstetter 1952; Webb 1974; Cartelle, unpublished work; Menegaz, unpublished work), but included in a cladistic analysis for the first time here.

  1. 1.

    Lacrimal vacuity: present (0); absent (1). (Harrison 1979, 1985; Honey et al. 1998)

  2. 2.

    Maxillary fossa: present (0); absent (1). (Harrison 1979, 1985; Honey et al. 1998)

  3. 3.

    Nasal shape in transverse section: straight (0); arched (1). (Harrison 1979, 1985; Honey et al. 1998)

  4. 4.

    Zygomatic arch in lateral view: curved (0); straight (1). (Harrison 1979, 1985)

  5. 5.

    Rostrum length (calculated as “rostrum length ratio,” RLR: ratio between the dentary depth at P4 and distance of P4 to the canine; according to Cabrera 1931): elongated, RLR less than or equal to 0.5 (0); short, RLR greater than or equal to 0.6 (1). (Honey and Taylor 1978 mod.; Harrison 1979, 1985 mod.; Honey et al. 1998)

  6. 6.

    Rostrum width (calculated as “rostrum width ratio,” RWR: ratio between the rostrum’s narrowest point and the cranium’s widest point on the zygomatic arches): broad, RWR greater than 0.2 (0); narrow, RWR less than 0.2 (1). (Honey and Taylor 1978 mod.; Harrison 1979, 1985 mod; Honey et al. 1998)

  7. 7.

    Anterior edge of choanae: at the level of or posterior to M3 distal lobe (0); between the M3 mesial and distal lobes (1); at the level of M2 distal lobe or between M2 and M3 (2).

  8. 8.

    Anterior margin of dentary angle: well marked (0); weakly marked or absent (1). (Harrison 1979, 1985; Honey et al. 1998)

  9. 9.

    Crest of mandibular diastema: strong and acute (0); reduced and rounded (1). (Harrison 1979, 1985; Honey et al. 1998)

  10. 10.

    First upper incisor: present (0); absent (1). (Honey and Taylor 1978; Harrison 1979, 1985; Honey et al. 1998)

  11. 11.

    Second upper incisor: present (0); absent (1). (Honey and Taylor 1978; Harrison 1979, 1985; Honey et al. 1998)

  12. 12.

    Lower incisor shape: spatulated crown, small wear facets, quite imbricated, thick enamel layer on the lingual side (0); cylindrical crown with trapezoidal section, large chisel-shaped wear facets, little imbricated or not imbricated, thin enamel layer or absence of enamel on the lingual side (1). (Harrison 1985; Honey et al. 1998)

  13. 13.

    Canine transverse section: elliptical or laterally compressed (0); circular or rounded (1). (Harrison 1979, 1985; Honey et al. 1998)

  14. 14.

    First upper premolar: present (0); absent (1). (Harrison 1979, 1985; Honey et al. 1998)

  15. 15.

    First lower premolar: present (0); absent (1). (Honey and Taylor 1978; Harrison 1979, 1985; Honey et al. 1998)

  16. 16.

    Second upper premolar: present (0); absent (1). (Harrison 1979, 1985; Honey et al. 1998)

  17. 17.

    Second lower premolar: present (0); absent (1). (Honey and Taylor 1978; Harrison 1979, 1985; Honey et al. 1998)

  18. 18.

    Third upper premolar: present (0); absent (1). (Harrison 1979; Honey et al. 1998 mod)

  19. 19.

    Third lower premolar: present (0); absent (1). (Harrison 1979, 1985; Honey et al. 1998 mod)

  20. 20.

    Fourth lower premolar shape: triangular, with fossettid only in the distal lobe (0); quadrangular, with fossetids in the mesial and distal lobes.

  21. 21.

    Molar lingual lophs and labial lophids: V-shaped or triangular (0); U-shaped or rounded (1).

  22. 22.

    Molar labial styles and lingual stylids (“ribs”): well developed (0); poorly developed (1).

  23. 23.

    Enamel folds in molars: present (0); absent (1).

  24. 24.

    Proto- and parastylids (“lama buttress”): small or absent (0); greatly developed (1). (Harrison 1979, 1985; Honey et al. 1998)

  25. 25.

    Ratio between humerus and radius-ulna lengths (RUR): long humerus, RUR greater than 0.8 (0); short humerus, RUR less than 0.8 (1).

  26. 26.

    Femur and tibia lengths: length comparable or femur longer (0); femur shorter (1).

  27. 27.

    Metacarpal and metatarsal lengths: metacarpal shorter (0); metacarpal longer or lengths comparable (1). (Harrison 1979, 1985 mod; Honey et al. 1998)

  28. 28.

    Metacarpal and humerus lengths: metacarpal shorter or lengths comparable (0); metacarpal longer (1).

  29. 29.

    Ratio between metatarsal and femur lengths (RMF): short metatarsal, RMF less than 0.8 (0); long metatarsal, RMF greater than 0.8 (1).

  30. 30.

    Radio-ulna length: short, less than 370 mm (0); long, greater than 390 mm (1).

  31. 31.

    Tibia length: short, less than 340 mm (0); long, greater than 370 mm (1).

  32. 32.

    Metapodial length: short, less than 270 mm (0); long, greater than 280 mm (1). (Honey et al. 1998 mod)

  33. 33.

    Limb segment contribution to the limb total length: stylopodial, zygopodial, and metapodial with similar lengths or metapodial shorter (0); zygopodial and metapodial longer than stylopodial (1).

  34. 34.

    Radio-ulna gracility (calculated as “gracility index,” GI; the ratio between the radio-ulna proximal width and its total length): gracile, GI less than 0.13 (0); robust, GI greater than 0.13 (1).

  35. 35.

    Metapodial gracility (calculated as for the radio-ulna): gracile, GI less than 0.15 (0); robust, GI greater than 0.15 (1). (Honey and Taylor 1978 mod)

Appendix II. Data Matrix Used in Cladistic Analysis

In the matrix “A” is the polymorphic condition (0 + 1) and “?” means unknown condition.

Table 1
Full size table

Appendix III. Synapomorphies in the Strict Consensus Tree

  • Node 2. Protolabinae + Camelinae: 29(1) (ambiguous).

  • Node 3. Protolabinae: 6(1).

  • Node 5. Procamelus + Camelinae: 7(1) (ambiguous), 10(1), 11(1), 27(1), 30(1), 31(1), 32(1), 34(1) (ambiguous), 35(1) (ambiguous).

  • Node 6. Camelini + Lamini: 16(1), 17(1).

  • Node 7. Camelini: 1(1) (convergent with clade of node 18), 2(1) (convergent with clade of node 16), 9(1), 13(1).

  • Node 8. Lamini: 3(1), 7(2), 29(0) (reversion).

  • Node 10. Unnamed node: 14(1), 22(1) (convergent with Procamelus).

  • Node 11. Unnamed node: 15(1).

  • Node 12. Palaeolama: 20(1).

  • Node 13. Palaeolama major + P. mirifica: 22(0) (reversion).

  • Node 14. Unnamed node: 8(1) (convergent with Michenia), 21(1) (convergent with Procamelus, Megatylopus and Camelus), 23(1) (convergent with Camelus), 25(1) (convergent with Megacamelus).

  • Node 15. Unnamed node: 29(1) (ambiguous), 28(1) (convergent with Camelus).

  • Node 16. Unnamed node: 2(1) (convergent with Camelini), 24(1) (convergente with Aepycamelus), 35(0) (reversion).

  • Node 17. Hemiauchenia paradoxa + H. macrocephala: 33(1), 26(1) (convergente with Megacamelus and Vicugna spp.)

  • Node 18. Unnamed node: 18(1), 1(10) (convergent with Camelini), 19(1) (convergent with Camelus, H. blancoensis and Camelops), 31(0) (reversion), 34(0) (reversion).

  • Node 19. Hemiauchenia gracilis + H. edensis: 23(0) (reversion), 27(0) (reversion).

  • Node 20. Lama castelnaudi + (L. guanicoe + (Vicugna)): 5(1), 30(0) (reversion), 32(0) (reversion).

  • Node 21. L. guanicoe + (Vicugna vicugna + V. provicugna) 25(0) (reversion), 28(0) (reversion).

  • Node 22. Vicugna: 12(1), 26(1) (convergent with Megacamelus and Hemiauchenia paradoxa + H. macrocephala).

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Scherer, C.S. The Camelidae (Mammalia, Artiodactyla) from the Quaternary of South America: Cladistic and Biogeographic Hypotheses. J Mammal Evol 20, 45–56 (2013). https://doi.org/10.1007/s10914-012-9203-4

Download citation

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10914-012-9203-4

Keywords

  • Camelidae
  • South America
  • Quaternary