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. 2020;139(1):5.
doi: 10.1186/s13358-020-00208-6. Epub 2020 Oct 7.

Description of a fossil camelid from the Pleistocene of Argentina, and a cladistic analysis of the Camelinae

Sinéad Lynch  1 , Marcelo R Sánchez-Villagra  1 , Ana Balcarcel  1
Affiliations

Description of a fossil camelid from the Pleistocene of Argentina, and a cladistic analysis of the Camelinae

Sinéad Lynch et al. Swiss J Palaeontol. 2020.

Abstract

We describe a well-preserved South American Lamini partial skeleton (PIMUZ A/V 4165) from the Ensenadan (~ 1.95-1.77 to 0.4 Mya) of Argentina. The specimen is comprised of a nearly complete skull and mandible with full tooth rows, multiple elements of anterior and posterior limbs, and a scapula. We tested this specimen's phylogenetic position and hypothesized it to be more closely related to Lama guanicoe and Vicugna vicugna than to Hemiauchenia paradoxa. We formulate a hypothesis for the placement of PIMUZ A/V 4165 within Camelinae in a cladistic analysis based on craniomandibular and dental characters and propose that future systematic studies consider this specimen as representing a new species. For the first time in a morphological phylogeny, we code terminal taxa at the species level for the following genera: Camelops, Aepycamelus, Pleiolama, Procamelus, and Alforjas. Our results indicate a divergence between Lamini and Camelini predating the Barstovian (16 Mya). Camelops appears as monophyletic within the Camelini. Alforjas taylori falls out as a basal member of Camelinae-neither as a Lamini nor Camelini. Pleiolama is polyphyletic, with Pleiolama vera as a basal Lamini and Pleiolama mckennai in a more nested position within the Lamini. Aepycamelus and Procamelus are respectively polyphyletic and paraphyletic. Together, they are part of a group of North American Lamini from the Miocene epoch.

Keywords: Camelinae; Lamini; North America; Phylogeny; South America.

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Conflict of interest statement

Competing interestsThe authors declare that they have no competing interests.

Figures

Fig. 1
Fig. 1
Time range of extinct species included in this study. Not to scale, species time range approximate. Species positioning in the Protolabinae follows Honey (1998) and Honey (2007). Species positioning in the Camelinae, Lamini, and Camelini follow our results. Species range for North American species following Baskin and Thomas (2016); Bravo-Cuevas and Jiménez-Hidalgo (2015); Harrison (1979); Honey and al. (1998); Honey and Taylor (1978); Pagnac (2005); Peterson (1911); Prothero (2005); Webb and Meachen (2004). Absolute ages for North American land mammal age following Woodburne (2004). Species range for South American species are uncertain and discussed by Scherer (2013). Absolute ages for South American land mammal age following Verzi et al. (2004); Soibelzon and al. (2008); Cione and Tonni (2001); Cione and Tonni (2005); Cione et al. (2015). Abbreviations: Chadron. = Chadronian; Whitn. = Whitneyan; Arikar. = Arikareean; Heming. = Hemingfordian; Barstov. = Barstovian; Clarend. = Clarendonian; Hemph. = Hemphillian; Irving. = Irvingtonian; Ranchol. = Rancholabrean; E = early; M = middle; L = late
Fig. 2
Fig. 2
Schematic drawings of L. guanicoe. a Skull, lateral view. b Skull, dorsal view. c Skull, occlusal view. d Mandible, lateral view. Abbreviations: pr = premaxillary bone; ma = maxillary bone; fr = frontal bone; la = lacrimal bone; ju = jugal bone; te = temporal bone; oc = occipital bone; sp = sphenoid bone; pt = pterygoid bone; vo = vomer bone; pl = palatine bone; pa = parietal bone; I3 = third upper incisor; I1 = first lower incisor; I2 = second lower incisor; I3 = third lower incisor; C = Upper canine; c = Lower canine; P1 = first upper premolar; P1 = first lower premolar; P2 = second lower premolar; P4 = fourth upper premolar; P4 = fourth lower premolar; M1 = first upper molar; M1 = first lower molar; M2 = second upper molar; M2 = second lower molar; M3 = third upper molar; M3 = third lower molar
Fig. 2
Fig. 2
Schematic drawings of L. guanicoe. a Skull, lateral view. b Skull, dorsal view. c Skull, occlusal view. d Mandible, lateral view. Abbreviations: pr = premaxillary bone; ma = maxillary bone; fr = frontal bone; la = lacrimal bone; ju = jugal bone; te = temporal bone; oc = occipital bone; sp = sphenoid bone; pt = pterygoid bone; vo = vomer bone; pl = palatine bone; pa = parietal bone; I3 = third upper incisor; I1 = first lower incisor; I2 = second lower incisor; I3 = third lower incisor; C = Upper canine; c = Lower canine; P1 = first upper premolar; P1 = first lower premolar; P2 = second lower premolar; P4 = fourth upper premolar; P4 = fourth lower premolar; M1 = first upper molar; M1 = first lower molar; M2 = second upper molar; M2 = second lower molar; M3 = third upper molar; M3 = third lower molar
Fig. 3
Fig. 3
Schematic drawings of L. guanicoe’s teeth. a M3, occlusal view. b M3, occlusal view
Fig. 4
Fig. 4
a, b: Char. 7: “Lower canine position (c)”. a H. macrocephala (UF 205,750), scored close from incisor (I3) (< 1 cm) (0). b L. guanicoe (ZM 17,209), scored far from incisor (I3) (≥ 1 cm) (1). c, f: Char. 18: “Metacone and paracone on upper molars (M1-M3)”. c Right M2, P. coartatus (AMNH 73,377), scored weakly developed (0). f Left M2, P. weddelli (PUN 1), scored well developed (1). d, e: Char. 19: “Anterior palatine fenestra”. d P. coartatus (AMNH 73,438), ventral view, scored absent (0). e L. guanicoe (ZM 17,209), ventral view, scored present (1). g, h, i, j: Char. 9: “First lower premolar (P1)”. g P. wilsoni (AMNH 47,130), right side, scored premolariform (0). h L. guanicoe (ZM 17,967), left side, scored absent (2). i A. robustus (OMNH 016,560), right side, scored caniniform (1). j H. macrocephala (UF 205,750), right side, scored caniniform (1). g, h, i, j: Char. 11: “Second lower premolar (P2)”. g P. wilsoni (AMNH 47,130), right side, scored premolariform (0). h L. guanicoe (ZM 17,967), left side, scored absent (2). i A. robustus (OMNH 016,560), right side, scored caniniform (1). j H. macrocephala (UF 205,750), right side, scored absent (2)
Fig. 5
Fig. 5
a, b: Char. 20: “Position of the most posterior part of the palatine process of premaxillary”. a L. guanicoe (ZM 17,209), scored closer to the third incisor (1). b A. taylori (AMNH 40,821), scored closer to the canine (0). c, d: Char. 24: “Lateral border of nasals”. c L. guanicoe (ZM 17,209), left side, scored curved (1). d T. brachydontus (AMNH 36,594), left side, scored straight (0)
Fig. 6
Fig. 6
Char. 25: Bones bordering the lacrimal vacuity. a L. guanicoe (ZM 17,209), left side, scored small, bordered by four bones (frontal, maxillary, lacrimal, nasal) (1) b P. vera (AMNH 24,670), left side, scored large, bordered by four bones (frontal, maxillary, lacrimal, nasal) (0). c A. alexandrae (UCMP 26,015), left side, scored bordered by three bones (frontal, maxillary, and lacrimal) (3). d C. hesternus (UCMP 20,040), left side, scored bordered by four bones (frontal, maxillary, lacrimal, and nasal), lacrimal border greatly reduced, frontal border greatly enlarged (4)
Fig. 7
Fig. 7
A, B, C: Char. 26: “Maxillary fossa”. a C. hesternus (UCMP 20,040), left side, scored well-developed, large pocket (0). b A. robustus (OMNH 016,560), right side, scored well-developed, small pocket (1). c L. guanicoe (ZM 17,209), left side, scored shallow or absent (2). d, e: Char. 30: “Shape of the anterior edge of the choanae”. d P. vera (AMNH 24,670), ventral view, scored V-shaped (0). e L. guanicoe (ZM 17,209), ventral view, scored U-shaped (1). f, g: Char. 31: “Anterior edge of the pterygopalatine fossa”. f M. matthewi (UCMP 31,100), scored at the level of M3 (1). g T. longirostris (CM 2498), scored posterior to M3 (0). h, i: Char. 32: “Position of the ethmoid foramen”. h Upper: above the fossa formed by a division of the lateral pterygoid muscle (0). Lower: in the fossa formed by a division of the lateral pterygoid muscle (1). i L. guanicoe (ZM 17,209), right side, scored above the fossa formed by a division of the lateral pterygoid muscle (0)
Fig. 8
Fig. 8
a, b: Char. 35: “Postglenoid process and postglenoid foramen”. a C. bactrianus (ZM 17,970), right side, scored large (1). b L. guanicoe (ZM 17,209), left side, scored small-to-absent (0). a, b: Char. 38: “Position of the lateral plate of the bullae in lateral view”. a C. bactrianus (ZM 17,970), right side, scored below external auditory meatus (0). b L. guanicoe (ZM 17,209), left side, scored anterior to external auditory meatus (1). c, d: Char. 37: “Tympanic bullae”. c C. bactrianus (ZM 17,970), ventral view, scored little inflated (1). d L. guanicoe (ZM 17,209), ventral view, scored greatly inflated (0)
Fig. 9
Fig. 9
a, b: Char. 42: “Angle of the mandible in lateral view”. a L. guanicoe (ZM 17,967), left side, scored rounded (2). b P. coartatus (AMNH 73,306), right side, scored ventrally produced with lateral flare (1). c, d: Char. 43: “Coronoid process at the level of the mandibular condyle in lateral view”. c T. brachyodontus (AMNH 36,594), right side, scored angled posteriorly (0). d L. guanicoe (ZM 17,967), left side, scored straight (1). e, f: Char. 40: “Distance between mesial end of the third lower incisors compared to the distance between the mesial end of fourth lower premolars”. e C. minidokae (UCMP 38,448), scored distance between I3 greater than distance between P4 (1). f P. grandis (UCMP 32,864), scored distance between P4 greater or equal to distance between I3 (0). g, h: Char. 39: “Anterior part of the mandible in lateral view”. g L. guanicoe (ZM 17,967), left side, scored straight (0). h T. longirostris (CM 2498), left side, scored angled dorsally (1). i Char. 41: “Portion of the mandibular ramus below the mandibular diastema in transverse section”. Left: Little narrowing dorsally, ovoid (1). Right: Strongly narrowing dorsally, tear-drop shaped (0)
Fig. 10
Fig. 10
PIMUZ A/V 4165. a Skull lateral view. b Skull occlusal view. c Mandible lateral view. d Mandible occlusal view
Fig. 11
Fig. 11
PIMUZ A/V 4165. a Left and right humeri, dorsal view. b Left and right metacarpals and carpals, ventral view. c Phalanges, ventral view. d Left scapula, lateral view. e Left radio-ulna, lateral view. f Left femur, dorsal view. g Right and left tibiae, dorsal view. h Right metatarsal and tarsals, dorsal view
Fig. 12
Fig. 12
Most parsimonious tree (TL = 149 steps, CI = 0.423, RI = 0.675, RC = 0.286). Numbers in circles indicate node numbers. Numbers below branches indicate Bremer supports. Only clade 48 and clade 52 have a Bootstrap value over 50%, respectively, at 71% and 64% (see Additional file 2, for information on the skulls, pictures not to scale)
See this image and copyright information in PMC

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