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

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.

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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.

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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

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).

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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

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Keywords

  • Camelidae
  • South America
  • Quaternary