Fig 1.
Location of Cerro de los Batallones.
A, Location of Cerro de los Batallones (yellow star) within Spain (Modified from [20]; B, map of Cerro de los Batallones and location of the fossil sites (Modified from [20]).
Fig 2.
Articulated giraffid specimen from BAT10.
Postcranial skeleton articulated with BAT10’13.E2-69 skull. Black, recovered during the 2013 campaign; grey, reverend during the 2014–16 campaigns. Scale bar equals 10 cm.
Fig 3.
Cranial remains of Decennatherium rex sp. nov. from BAT10.
A-C, BAT10’13.E2-69, skull in A, dorsal view; B, medial/lateral view; C, ventral view; D, caudal view. Scale bar equals 10 cm.
Fig 4.
Cranial remains of Decennatherium rex sp. nov. from BAT10.
A-C, BAT10’08.G3-91, skull in A, dorsal view; B, medial/lateral view; C, ventral view; D, ventral view of the basioccipital region. Scale bar equals 10 cm.
Fig 5.
Posterior ossicone of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’16.D4-20, right posterior ossicone in A, cranial view; B, lateral view; C, caudal view; D, medial view; E, proximal view. Scale bar equals 5 cm.
Fig 6.
Fragment of a posterior ossicone with bumps of Decennatherium rex sp. nov. from BAT10.
BAT10’07.G3-9. Scale bar equals 5 cm.
Fig 7.
Skull angles (characters 52–53) of Decennatherium rex sp. nov. from BAT10.
Defined by (8). Scale bar equals 10 cm. 3D correction by Oscar Sanisidro using ZBrush software.
Fig 8.
Dental remains of Decennatherium rex sp. nov. from BAT10.
A, BAT10’08.G3-91, left P2-M3 in occlusal view; B-D, BAT10’10.F2-18, left p2-m3 in B, occlusal view; C, lingual view; D, buccal view. Scale bar equals 5 cm.
Fig 9.
Deciduous teeth rows of Decennatherium rex sp. nov. from BAT10.
A, BAT10’09.G2-177, left DP2-DP4 in occlusal view; B, BAT10’09.F4-22, left dp2-dp4 in occlusal view. Scale bar equals 1 cm.
Fig 10.
Dental remains of Decennatherium rex sp. nov. from BAT10.
A, BAT10’07.G3-21, deciduous canine in buccal view; B-C, BAT10’12.D6-12, deciduous incisor in B, buccal view; C, lingual view; BAT10’15.D4-63, canine in buccal view; D, BAT10’07.G2-12, buccal view. Scale bar equals 1 cm.
Fig 11.
Occlusal views of the p3 of the giraffid species included in this work.
A, M26675-P80, C. sirtensis; B, from de Bonis et al., 1997, Teftfig. 3c, G. georgalasi; C, AM-127634, I. arabicum; D, AM-29946, G. punjabiensis; E, M30159, G. primaevus; F, MNCN-3439, G. camelopardalis; G, M21466, G. jumae; H, M8367, P. rouenii; I, AM-26360, P. microdon; J, MAR-669, P. coelophrys; K, MNCN-5226, O. johnstoni; L, from Schlosser, 1903 Tafel 9.10; S. sinense; M, from Hamilton, 1978, Fig 5a, S. tafeli; N, D. rex; I, BAT10’10.F2-18; II, BAT10’11.E2-21; III, BAT10’12.D6-85; IV; BAT10’11.C5-6; O, D. pachecoi, I, MNCN-43462; II, MNCN-43463; III, MNCN-43461; IV; MNCN-43465; P, from Hamilton, 1978, Fig 5, S. boisieri; Q, from Hamilton, 1978, Fig 5, A. neumayri; R, B. schaubi, I, CR2-215; II, CR2-879; III, VP-868; S, SAM-PQ-L31137 from Harris, 1976 Fig 4b, S. hendeyi; T, MNHN-1911-34, H. duvernoyi; U, KNM-ER-777 from Harris, 1991 Fig 4.22, S. maurusium; V, NHM-16163, S. giganteum; W, AM-19670, B. megacephalum. Images not to scale and modified so they have the same orientation.
Fig 12.
Atlas of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’08.G3-79, in A, cranial view; B, dorsal view; C, medial/lateral view; D, ventral view; E, caudal view. Scale bar equals 5 cm.
Fig 13.
Axis of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’14.D4-32, in A, cranial view; B, dorsal view; C, medial/lateral view; D, ventral view; E, caudal view. Scale bar equals 5 cm.
Fig 14.
Neck of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’14.E2-71-74, C3-C6 in A, dorsal view; B, medial/lateral view; C, ventral view; D, cranial view; E, caudal view. Scale bar equals 5 cm.
Fig 15.
Thoracic vertebrae of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’08.D5-10, in A, cranial view; B, medial/lateral view; C, caudal view; D, dorsal view; E, ventral view. Scale bar equals 5 cm.
Fig 16.
Lumbar vertebrae of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’13.D4-40, in A, cranial view; B, medial/lateral view; C, caudal view; D, dorsal view; E, ventral view. Scale bar equals 5 cm.
Fig 17.
Caudal vertebrae of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’14.E2-30, in A, cranial view; B, caudal view; C, medial/lateral view; D, dorsal view; E, ventral view. Scale bar equals 5 cm.
Fig 18.
Scapula of Decennatherium rex sp. nov. from BAT10.
A-D, BAT10’08.F2-51, right scapula in A, medial view; B, lateral view; C, dorsal view; D, distal view; Scale bar equals 10 cm.
Fig 19.
Forelimb of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’13.E2-67, right humerus in A, dorsal view; B, lateral view; C, palmar view; D, medial view; E, proximal view; F, distal view; G-L, BAT10’09.G2-104, right radius-ulna in G, dorsal view; H, lateral view; I, palmar view; J, medial view; K, proximal view; L, distal view. Scale bar equals 5 cm.
Fig 20.
Carpals of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2-78, left scaphoid in in A, dorsal view; B, medial view; C, palmar view; D, lateral view; E, proximal view; F, distal view; G-L, BAT10’14.E2-77, left semilunate in G, dorsal view; H, medial view; I, palmar view; J, lateral view; K, proximal view; L, distal view; M-R, BAT10’14.E2-76, left triquetrum in M, dorsal view; N, medial view; O, palmar view; P, lateral view; Q, proximal view; R, distal view; S-W, BAT10’14.E2-46, left pisiform in S, dorsal view; T, medial view; U, lateral view; V, proximal view; W, distal view. Scale bar equals 5 cm.
Fig 21.
Carpals of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2.80–74, left magnotrapezoid in in A, dorsal view; B, medial view; C, palmar view; D, lateral view; E, proximal view; F, distal view; G-L, BAT10’14.E2-80, left hamatum in G, dorsal view; H, medial view; I, palmar view; J, lateral view; K, proximal view; L, distal view. Scale bar equals 5 cm.
Fig 22.
Metacarpal III-IV of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’07.G2-28, left metacarpal III-IV in A, dorsal view; B, medial view; C, plantar view; D, proximal view; E, distal view. Scale bar equals 5 cm.
Fig 23.
Pelvis of Decennatherium rex sp. nov. from BAT10.
A-B, BAT10’09.G1-1, right pelvis fragment in A, medial view; B, lateral view. Scale bar equals 5 cm.
Fig 24.
Hindlimb of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2-10, left femur in in A, dorsal view; B, lateral view; C, plantar view; D, medial view; E, proximal view; F, distal view; G-L, BAT10’14.E2-15, right patella in G, dorsal view; H, lateral view; I, plantar view; J, medial view; K, proximal view; L, distal view; M-R, BAT10’14.E2-26, left tibia in M, dorsal view; N, lateral view; O, plantar view; P, medial view; Q, proximal view; R, distal view; S-W, BAT10’14.E2-27, right fibula in S, dorsal view; T, lateral view; U, plantar view; V, medial view; W, proximal view; X, distal view. Scale bar equals 5 cm.
Fig 25.
Tarsals of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2-17, right calcaneus in in A, dorsal view; B, medial view; C, plantar view; D, lateral view; E, proximal view; F, distal view; G-L, BAT10’14.E2-23, right astragalus in G, dorsal view; H, medial view; I, plantar view; J, lateral view; K, proximal view; L, distal view; M-R, BAT10’14.E2-32, rightnavicular-cuboid in M, dorsal view; N, medial view; O, plantar view; P, lateral view; Q, proximal view; R, distal view. Scale bar equals 5 cm.
Fig 26.
Tarsals of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2-38, right ectomesocuneiform in in A, dorsal view; B, medial view; C, plantar view; D, lateral view; E, proximal view; F, distal view; G-L, BAT10’14.E2-37, right entocuneiform in G, dorsal view; H, medial view; I, plantar view; J, lateral view; K, proximal view; L, distal view. Scale bar equals 5 cm.
Fig 27.
Metatarsal III-IV of Decennatherium rex sp. nov. from BAT10.
A-D, BAT10’12.D6-54, right metatarsal III-IV in A, dorsal view; B, medial view; C, plantar view; D, proximal view. Scale bar equals 5 cm.
Fig 28.
Lateral metapodials of Decennatherium rex sp. nov. from BAT10.
A-B, BAT10’12.D6-108, in A, dorsal view; B, plantar view; C-D, BAT10’13.E2-56 in A, lateral view; B, medial view. Scale bar equals 5 cm.
Fig 29.
Phalanges of Decennatherium rex sp. nov. from BAT10.
A-E, BAT10’14.E2-11, first phalanx of the right digit in A, distal view; B, dorsal view; C, palmar/plantar view; D, exterior view; E, proximal view; F-J, BAT10’14.E2-33, second phalanx of the right digit in F, distal view; G, dorsal view; H, palmar/plantar view; I, exterior view; J, proximal view; K-O, BAT10’14.E2-22, third phalanx of the right digit in K, distal view; L, dorsal view; M, palmar/plantar view; N, exterior view; O, proximal view; Scale bar equals 5 cm.
Fig 30.
Sesamoids of Decennatherium rex sp. nov. from BAT10.
A-F, BAT10’14.E2-42, proximal exterior sesamoid in A, cranial view; B, lateral view; C, caudal view; D, medial view; E, dorsal view; F, palmar/plantar view; G-L; BAT10’14.E2-43, proximal interdigital sesamoid in G, cranial view; H, lateral view; I, caudal view; J, medial view; K, dorsal view; L, palmar/plantar view; M-R; BAT10’14.E2-57, distal sesamoid in M, cranial view; N, lateral view; O, caudal view; P, medial view; Q, dorsal view; R, palmar/plantar view. Scale bar equals 5 cm.
Fig 31.
Phylogenetic relationships among giraffids.
MPT of 277 steps (CI = 0,671; RI = 0,780). Bremer values over each node.
Fig 32.
Ossicones of several giraffid species.
A-E, BAT10’16.D4-20, right posterior ossicone of Decennatherium rex in A, cranial view; B, lateral view; C, caudal view; D, medial view; E, proximal view; F-J, MGUV-7816,right posterior ossicone of Birgerbohlinia schaubi in F, cranial view; G, lateral view; H, caudal view; I, medial view; J, proximal view; K, SAM-PQ-L12730, left posterior ossicone of Sivatherium hendeyi in lateral view ([63]: Fig 2A, holotype); L-P, BM-39525 left posterior ossicone of Sivatherium giganteum in L, cranial view; M, medial view; N, caudal view; O, lateral view; P, proximal view; Q-R, M14955, illustration of the left posterior ossicone of Sivatherium olduvaiense in Q, posterolateral view; R, anteromedial view ([62]: Plate IIIa, b.). Scale bar equals 5 cm.
Fig 33.
Lateral views of the skulls of the giraffid species included in the analysis.
A, MNCN-5226, Okapia johnstoni; B, AMNH/WN, Schansitherium tafeli; C, BAT10’13.E2-69, Decennatherium rex; D, BAT10’08.G3-91, Decennatherium rex; E, PIK-1500, Helladotherium duvernoyi; F, AM-27016, Bramatherium megacephalum M26675-P80, G, NHM-15283, Sivatherium giganteum; H, ANK-II-127 ([91]: Fig 1), Sivatherium maurusium. Images not to scale.
Fig 34.
Lateral views of the skulls of the giraffid species included in the analysis.
A, M 26670, Canthumeryx sirtensis; B, THB-30 ([12]: Textfig. 1), Gerogiomeryx georgalasi; C, AMNH-19475, Giraffokeryx punjabiensis; D, OLD 63 EF HR, Giraffa jumae [92]; E, MNCN-3439, Giraffa camelopardalis; F, AMNH-10452, Palaeotragus rouenii; G, M1124, Palaeotragus microdon; H, Samotherium major ([1]: Fig 21.5b); I, Samotherium boissieri ([1]: Fig 21.5c). Images not to scale.
Fig 35.
Metacarpals of the giraffid species included in the analysis.
A, AMNH-51198, left and right metacarpals of Okapia johnstoni in dorsal and palmar views respectively; B, AM-948, left metacarpal of Sivatherium giganteum in dorsal and palmar views; C, BM-17102a, right metacarpal of Sivatherium giganteum in dorsal and palmar views; D, PQ-L45066, right metacarpal of Siatherium hendeyi in dorsal and palmar views; E, right metacarpal of Sivatherium maurusium in dorsal view ([64]: Fig. 7c); F, PIK-2756, left metacarpal of Palaeotragus rouenii in dorsal and palmar views; G, MNCN-42769, left metacarpal of Decennatherium pachecoi in dorsal and palmar views; H, BAT10’07.G2-28, left metacarpal of Decennatherium rex in dorsal and palmar views; I, MGUV-7806, right metacarpal of Birgerbohlinia schaubi in dorsal and palmar views; J, AM-26349, left metacarpal of S. sinense in dorsal and palmar views; K, AMNH-20758, left metacarpal of Samotherium boissieri in dorsal and palmar views; L, left metacarpal of Alcicephalus neumayri in dorsal and palmar views; M, AM-19460, left metacarpal of Bramatherium megacephalum in dorsal and palmar views; N, M11377, left metacarpal of Helladotherium duvernoyi in dorsal and palmar views; O, PIK-2756, left metacarpal of Bohlinia attica in dorsal and palmar views; P, M14957, left metacarpal of Giraffa jumae in dorsal and palmar views; Q, AMNH-53543, right and left metacarpals of Giraffa camelopardalis in dorsal and palmar views respectively. Scale bar equals 5 cm.
Fig 36.
Robustness of the metacarpal III-IV of the giraffids analyzed.
Metacarpal III-IV Robustness Index (RI) Diaphysis TD/Total Length*100. TD, transverse diameter. Measurements given in mm.
Fig 37.
Character 94. Metacarpal III-IV Robustness Index (RI).
RI = Diaphysis TD/Total Length*100. Scale bar equals 5 cm.
Fig 38.
Character 96. Metacarpal III-IV elongation.
Scale bar equals 5 cm.
Fig 39.
Character 97. Metacarpal III-IV palmar trough depth.
Scale bar equals 5 cm.
Fig 40.
Distribution of the Iberian late Miocene giraffids.
Fig 41.
Reconstruction of Decennatherium rex sp. nov. from BAT10.
A, skeletal reconstruction; B, life reconstruction of an adult female; C, life reconstruction of the head of an adult female. Illustration by Oscar Sanisidro.
Table 1.
Distribution of autapomorphic character states for the internal nodes.
Node K includes Decennatherium.
Table 2.
Body mass predictions for Decennatherium rex using a right humerus and femur and a left humerus and femur.