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Volume 27.1
January–April 2024
Full table of contents
ISSN: 1094-8074, web version;
1935-3952, print version
Recent Research Articles
See all articles in 27.1 January-April 2024
See all articles in 26.3 September-December 2023
See all articles in 26.2 May-August 2023
See all articles in 26.1 January-April 2023
María Ríos. Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-Consejo Superior de Investigaciones Científicas, 2 Gutiérrez Abascal, Madrid, 28006, Spain. maria.rios.iba@mncn.csic.es
María Ríos is a PhD. granted candidate at the Museo Nacional de Ciencias Naturales-Consejo Superior de Investigaciones Científicas, Madrid (Spain). Her studies deal with the systematics, taxonomy, morphometry, phylogeny, paleobiology and evolution of the fossil spanish giraffids, mainly focusing on the extraordinary giraffid collections from the Late Miocene site Batallones-10 (Cerro de los Batallones, Madrid, Spain).
Melinda Danowitz. Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine, 8000 Northern Boulevard, Old Westbury, NY 11568, USA. mdanowit@nyit.edu
Melinda Danowitz is a medical student and Academic Medicine Scholar. She is studying to become a pediatrician, but plans to continue paleontological research and pursue a career combining clinical medicine with academia. Her research focuses on ruminant skeletal morphology, with an emphasis on giraffid evolutionary anatomy and paleoecology.
Nikos Solounias. Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine, 8000 Northern Boulevard, Old Westbury, NY 11568, USA and Department of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA. nsolouni@nyit.edu
Dr. Nikos Solounias is a professor of anatomy at NYIT College of Osteopathic Medicine. An expert in modern and paleontological ungulate anatomy and biology, he has more than 40 years of experience in the research and teaching of mammalian anatomy and evolution. Dr. Solounias has published more than 80 articles in peer-reviewed journals and has been awarded four NSF grants. He received his B.S. in Biology from Cornell University and an M.A. in Biology-Embryology from Clark University. He earned his Ph. D. in Geology-Paleontology from the University of Colorado, Boulder and has held postdoctoral positions at Yale, Harvard, and Johns Hopkins universities. He studied anatomy and embryology at the Harvard Medical School.
FIGURE 1. Drawing indicating the different features described. Terminology using the right metapodials of Okapia johnstoni. AMNH 51218: 1, metacarpal palmar view; 2, metatarsal plantar view; and Samotherium major from Samos, MGL S 382; 3, pygmaios. Scale bar equals 50 mm.
FIGURE 2. Measurements applied to the giraffid metapodials, based on those proposed by Quiralte (2011), using the same specimen as in figure 1.
FIGURE 3. Helladotherium duvernoyi from Pikermi. 1, left metacarpal (reversed) MNHN 1658; 2, right metatarsal MNHN 1564.
FIGURE 4. Three selected metacarpal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.
FIGURE 5. Palmar views of the right metacarpals (left) and plantar views of metatarsals (right) of certain Giraffidae. 1, Helladotherium duvernoyi, late Miocene, Pikermi, NHM 11382 (metacarpal), MNHNP 1554 (metatarsal) (both reversed); 2, Bramatherium megacephalum, late Miocene-Pliocene, Siwaliks, AMNH 29820, (metacarpal), AMNH 19668 (metatarsal); 3, Sivatherium giganteum, Pleistocene, Siwaliks, NHM 39553 (metacarpal), NHM 39752 (metatarsal); 4, Decennatherium pachecoi, late Miocene, Los Valles de la Fuentidueña, MNCN 42769 (metacarpal), MNCN 42764 (metatarsal) (both reversed); 5, Birgerbohlinia schaubi, late Miocene, Piera, IPS 5060 (metacarpal), Crusafont, 1952, fig. 15-5 (metatarsal), (both reversed); 6, Okapia johnstoni, Zaire, AMNH 51196 (metacarpal), AMNH 51196 (metatarsal). Scale bar equals 200 mm.
FIGURE 6. Representative cross sections of right metacarpals. 1, Helladotherium duvernoyi, late Miocene, Pikermi, MNHNP 1527; 2, Birgerbohlinia schaubi, late Miocene, Samos, AMNH 20610 (reversed); 3, Okapia johnstoni, AMNH 15218 (reversed); 4, Samotherium major, late Miocene, Samos, AMNH 22790; 5, Bohlinia attica, late Miocene, Pikermi, PA 1474-1991; 6, Giraffa camelopardalis, AMNH 70016. Scale bar equals 80 mm.
FIGURE 7. Three additional metacarpal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.
FIGURE 8. Bramatherium megacephalum from the Upper Siwaliks. 1, Right metacarpal AMNH 29820; 2, Right metatarsal AMNH 19688.
FIGURE 9. Sivatherium giganteum from the Upper Siwaliks. 1, Right metacarpal NHM UK 39533; 2, Left metatarsal (reversed) NHM UK 39752.
FIGURE 10. Metacarpal proximal morphology. Birgerbohlinia schaubi, late Miocene, Samos, AMNH 20610. Abbreviations: HA, os hamatum facet; TC, os trapezoideocapitatum facet; S, synovial fossa. Image not to scale.
FIGURE 11. Samotherium major from Samos. 1, Right metacarpal NHMBe 711; 2, Right metatarsal AMNH 22967.
FIGURE 12. Palmar views of the right metacarpals (left) and plantar views of the metatarsals (right) of certain Giraffidae. 1, Samotherium major, late Miocene, Samos, NHM Be Samos 711 (metacarpal), AMNH 22967 (metatarsal) (both reversed); 2, Palaeotragus rouenii , late Miocene, Pikermi, MNHNP 1692 (metacarpal), (reversed); MNHNP 1691 (metatarsal). Scale bar equals 200 mm.
FIGURE 13. Bohlinia attica from Pikermi. 1, Right metacarpal MNHN 52780; 2, Right metatarsal MNHN 1639.
FIGURE 14. Palmar views of the right metacarpals (left) and plantar views of the metatarsals (right) of certain Giraffidae. 1, Canthumeryx sirtensis, early-middle Miocene, Moruorot Hill, NMK Mo 41 (metatarsal); 2, Bohlinia attica, late Miocene, Pikermi, MNHNP 27561 (metacarpal), MNHNP 2357 (metatarsal) (reversed); 3, Giraffa camelopardalis, Zaire, AMNH 53550 (metacarpal), AMNH 53550 (metatarsal) (both reversed). Scale bar equals 200 mm.
FIGURE 15. Giraffa camelopardalis from Kenya. 1, Left metacarpal (reversed) AMNH 53543; 2, Left metatarsal (reversed) AMNH 53550.
FIGURE 16. Three selected metatarsal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.
FIGURE 17. Three selected metatarsal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.
FIGURE 18. Metatarsal proximal morphology. 1, Helladotherium duvernoyi, late Miocene, Pikermi, M 1138; 2, Bramatherium megacephalum, late Miocene-Pliocene, Siwaliks, AMNH 19688; 3, Decennatherium pachecoi, late Miocene, Los Valles de Fuentidueña, MNCN 42764; 4, Birgerbohlinia schaubi, late Miocene, Piera, IPS 5090. Abbreviations: NC, os naviculocuboideum facet; CIL, os cuneiforme intermediolaterale facet; CM, os cuneiforme mediale facet. Images not to scale.
FIGURE 19. Box-and-whiskers plot with the metacarpal RI of the Giraffidae analyzed.
FIGURE 20. Dispersion plot with the absolute metacarpal length of the Giraffidae analyzed.
FIGURE 21. Dispersion plot showing the total length of the metacarpals versus the midshaft minimum transverse diameter of the Giraffidae analyzed.
FIGURE 22. Box-and-whiskers plot with the metatarsal RI of the Giraffidae analyzed.
FIGURE 23. Dispersion plot with the absolute metatarsal length of the Giraffidae analyzed.
FIGURE 24. Dispersion plot showing the total length of the metatarsals versus the midshaft minimum transverse diameter of the Giraffidae analyzed.
TABLE 1. Dimensions in mm of the metacarpal III-IV of Giraffidae ( DT, transversal diameter; Diaph., diaphysis; RI, robustness index).
Species | Specimen nº | Total Length | Diaph. TD | RI | Reference |
Canthumeryx sirtensis | UCB V.4899/42058 | 290 | 17,5 | 6,03 | Hamilton, 1978 |
Canthumeryx sirtensis | BU 20128 | 317 | Hamilton, 1973, 1978 | ||
Helladotherium duvernoyi | MNHN PIK1658 | 395 | 46 | 11,65 | |
Helladotherium duvernoyi | MNHN PIK1527 | 405 | 68 | 16,79 | |
Helladotherium duvernoyi | MNHN PIK1525 | 420 | 56 | 13,33 | |
Helladotherium duvernoyi | MNHN PIK1526 | 410 | 60 | 14,63 | |
Helladotherium duvernoyi | M 11382 | 408 | 56,61 | 13,88 | |
Helladotherium duvernoyi | M 11377 | 420 | 66,42 | 15,81 | |
Helladotherium duvernoyi | 400 | Gaudry et al., 1873 | |||
Helladotherium duvernoyi | 1972/9 | 420 | 67 | 15,95 | Melentis, 1974 |
Helladotherium duvernoyi | 1972/11 | 413 | 65 | 15,74 | Melentis, 1974 |
Helladotherium duvernoyi | MNHN PIK-1525 | 427 | Geraads, 2009 | ||
Helladotherium duvernoyi | MNHN PIK-1526 | 410 | Geraads, 2009 | ||
Helladotherium duvernoyi | BMNH M 11382 | 418 | Geraads, 2009 | ||
Helladotherium duvernoyi | MNHN PIK-1527 | 407 | Geraads, 2009 | ||
Helladotherium duvernoyi | BMNH M 11377 | 426 | Geraads, 2009 | ||
Helladotherium duvernoyi | AMNH 20610 | 445 | 65,21 | 14,65 | |
Helladotherium duvernoyi | MTLA248 | 425 | Kostopoulos, 2009 | ||
Helladotherium duvernoyi | MTLA249 | 438 | 70 | 15,98 | Kostopoulos, 2009 |
Helladotherium duvernoyi | MNHN MAR916 | 455 | 54 | 11,87 | |
Helladotherium duvernoyi | MNHN MAR914 | 460 | 66 | 14,35 | |
Helladotherium duvernoyi | MNHN, sans numero | 465 | Geraads, 2009 | ||
Helladotherium duvernoyi | HD-5504 | 435 | Geraads, 2009 | ||
Helladotherium duvernoyi | HD-5492 | 430 | Geraads, 2009 | ||
Helladotherium duvernoyi | sans numero | 418 | Geraads, 2009 | ||
Helladotherium duvernoyi | sans numero | 435 | Geraads, 2009 | ||
Helladotherium duvernoyi | sans numero | 470 | Geraads, 2009 | ||
Helladotherium duvernoyi | 408 | Geraads, 2009 | |||
Helladotherium duvernoyi | 436 | Geraads, 2009 | |||
Helladotherium duvernoyi | MNHN FLUB824 | 410 | 58 | 14,15 | |
Helladotherium duvernoyi | 410 | Gaudry et al., 1873 | |||
Bramatherium megacephalum | AM 29820 | 427 | 56 | 13,11 | Colbert, 1935 |
Bramatherium megacephalum | AM 19460 | 432 | 58 | 13,43 | Colbert, 1935 |
Bramatherium megacephalum | AM 19460 | 398 | 61 | 15,33 | Colbert, 1935 |
Sivatherium giganteum | OR 17102 | 320 | 57,53 | 17,98 | |
Sivatherium giganteum | OR 39605 | 320 | 66,85 | 20,89 | |
Sivatherium giganteum | OR 17102a | 312 | 59,69 | 19,13 | |
Sivatherium giganteum | OR 39600 | 327 | 72,83 | 22,27 | |
Sivatherium giganteum | OR 39533 | 340 | 69,1 | 20,32 | |
Sivatherium giganteum | OR 11089 | 302 | 59,54 | 19,72 | |
Sivatherium giganteum | OR 17101 | 313 | 68,16 | 21,78 | |
Sivatherium giganteum | OR 39603 | 310 | 72,83 | 23,49 | |
Sivatherium giganteum | OR 17092 | 69,07 | |||
Sivatherium giganteum | BM 39533 | 355 | Singer and Boné, 1960 | ||
Sivatherium giganteum | 348 | Dietrich, 1942 | |||
Decennatherium pachecoi | MNCN-42769 | 391 | 36,17 | 9,25 | |
Decennatherium pachecoi | 44,5 | Crusafont, 1952 | |||
Birgerbohlinia schaubi | IPS-935 | 373 | 48,89 | 13,11 | |
Birgerbohlinia schaubi | IPS-7640 | 382 | 48,3 | 12,64 | |
Birgerbohlinia schaubi | IPS-5060 | 374 | 47,28 | 12,64 | |
Birgerbohlinia schaubi | IPS-5059 | 381 | 51,01 | 13,39 | |
Birgerbohlinia schaubi | IPS-5061 | 54,58 | |||
Birgerbohlinia schaubi | IPS-5010 | 51,14 | |||
Birgerbohlinia schaubi | IPS-5014 | 63,4 | |||
Birgerbohlinia schaubi | IPS-7629 | 52,12 | |||
Birgerbohlinia schaubi | MGUV-7806 | 391 | 62,22 | 15,91 | |
Okapia johnstoni | AMNH 51218 | 320 | |||
Okapia johnstoni | AMNH 51903 | 321 | 32,98 | 10,27 | |
Okapia johnstoni | AMNH 51903 | 322 | 33,96 | 10,55 | |
Okapia johnstoni | AMNH 51200 | 316 | 30,27 | 9,58 | |
Okapia johnstoni | AMNH 51196 | 305 | 32,02 | 10,50 | |
Okapia johnstoni | AMNH 51200 | 315 | 31,31 | 9,94 | |
Okapia johnstoni | AMNH 51197 | 330 | 34,11 | 10,34 | |
Okapia johnstoni | AMNH 2268 | 317 | 32,13 | 10,14 | |
Okapia johnstoni | AMNH 51196 | 306 | 31,97 | 10,45 | |
Okapia johnstoni | AMNH 51197 | 328 | 34,26 | 10,45 | |
Okapia johnstoni | AMNH 51198 | 312 | 32,05 | 10,27 | |
Okapia johnstoni | AMNH 2268 | 321 | 33,47 | 10,43 | |
Okapia johnstoni | AMNH 51198 | 313 | 31,73 | 10,14 | |
Okapia johnstoni | MNCN-11425 | 295,3 | 31,32 | 10,61 | |
Okapia johnstoni | MA (6) O (I) | 307,5 | 32,9 | 10,70 | Churcher, 1970 |
Samotherium major | AMNH 10742 | 390 | 48,64 | 12,47 | |
Samotherium major | AMNH 20758 | 395 | 54,18 | 13,72 | |
Samotherium major | AMNH 22969 | 399 | 56,69 | 14,21 | |
Samotherium major | AMNH 20758 | 399 | 53,48 | 13,40 | |
Samotherium major | AMNH 22845 | 405 | 56,53 | 13,96 | |
Samotherium major | AMNH 22843-A | 405 | 57,14 | 14,11 | |
Samotherium major | AMNH 95122 | 409 | 51,22 | 12,52 | |
Samotherium major | AMNH 20595 | 412 | 54,11 | 13,13 | |
Samotherium major | AMNH 95121 | 415 | 52,73 | 12,71 | |
Samotherium major | AMNH 22973 | 416 | 54,58 | 13,12 | |
Samotherium major | AMNH 22970 | 416 | 53,33 | 12,82 | |
Samotherium major | AMNH 20614 | 423 | 49,71 | 11,75 | |
Samotherium major | AMNH 22971 | 428 | 57,82 | 13,51 | |
Samotherium major | AMNH 20636 | 471 | 52,48 | 11,14 | |
Samotherium major | MGL S32 | 415 | 47,6 | 11,47 | Kostopoulos, 2009 |
Samotherium major | PIM329 | 423 | 99 | 23,40 | Kostopoulos, 2009 |
Samotherium major | PMMS68 | 423 | 50,4 | 11,91 | Kostopoulos, 2009 |
Samotherium major | MYT41 | 397 | 52,4 | 13,20 | Kostopoulos, 2009 |
Samotherium major | AMNH20595 Q1 | 425 | 54,1 | 12,73 | Kostopoulos, 2009 |
Samotherium major | AMNH95121 Q1 | 415 | 53,4 | 12,87 | Kostopoulos, 2009 |
Samotherium major | MGL S35 | 423 | 50,3 | 11,89 | Kostopoulos, 2009 |
Samotherium major | PIM328 | 440 | 55 | 12,50 | Kostopoulos, 2009 |
Samotherium major | AMNH20614 Q1 | 432 | 49,3 | 11,41 | Kostopoulos, 2009 |
Samotherium major | MTLA76 | 425 | 58,7 | 13,81 | Kostopoulos, 2009 |
Samotherium major | AMNH95122 Q1 | 414 | 52 | 12,56 | Kostopoulos, 2009 |
Samotherium major | MTLA450 | 427 | 54,2 | 12,69 | Kostopoulos, 2009 |
Samotherium major | AMNH22973 Q5 | 420 | 41,9 | 9,98 | Kostopoulos, 2009 |
Samotherium major | PIM330 | 445 | 52 | 11,69 | Kostopoulos, 2009 |
Samotherium major | AMNH22971 Q5 | 440 | 57,2 | 13,00 | Kostopoulos, 2009 |
Samotherium major | PIM327 | 435 | 98 | 22,53 | Kostopoulos, 2009 |
Samotherium major | PIM336 | 413 | 51,7 | 12,52 | Kostopoulos, 2009 |
Samotherium major | PIM337 | 405 | 49,3 | 12,17 | Kostopoulos, 2009 |
Samotherium major | PIM324 | 427 | 47 | 11,01 | Kostopoulos, 2009 |
Samotherium major | AMNH22969 Q5 | 412 | 55,6 | 13,50 | Kostopoulos, 2009 |
Samotherium major | AMNH nn Q6 | 423 | 57,5 | 13,59 | Kostopoulos, 2009 |
Samotherium major | AMNH22970 Q5 | 420 | 43,2 | 10,29 | Kostopoulos, 2009 |
Samotherium major | PIM326 | 423 | 102 | 24,11 | Kostopoulos, 2009 |
Samotherium major | MTLA75 | 413 | 56,3 | 13,63 | Kostopoulos, 2009 |
Samotherium major | AMNH20758 Q4 | 400 | 53,5 | 13,38 | Kostopoulos, 2009 |
Samotherium major | MTLA117 | 57,3 | Kostopoulos, 2009 | ||
Samotherium major | MTLA398 | 51 | Kostopoulos, 2009 | ||
Samotherium major | MTLA399 | 55,3 | Kostopoulos, 2009 | ||
Samotherium major | MNHN SLQ690 | 410 | 40 | 9,76 | |
Samotherium major | MNHN SLQ691 | 408 | 41 | 10,05 | |
Samotherium major | KTA133 | 408 | 50 | 12,25 | Geraads, 1994 |
Samotherium major | KTB61 | 421 | 55 | 13,06 | Geraads, 1994 |
Samotherium major | 410 | Bohlin, 1926 | |||
Palaeotragus rouenii | M 11406 | 403 | 34,98 | 8,68 | |
Palaeotragus rouenii | M 11406 | 34,35 | |||
Palaeotragus rouenii | MTLB155 | 434 | 38,3 | 8,82 | Kostopoulos, 2009 |
Palaeotragus rouenii | MGL S781 | 410 | 37,4 | 9,12 | Kostopoulos, 2009 |
Palaeotragus rouenii | KTD26 | 405 | 33,5 | 8,27 | Geraads, 1994 |
Palaeotragus rouenii | KTA7 | 32,5 | Geraads, 1994 | ||
Palaeotragus rouenii | KTD27 | 32 | Geraads, 1994 | ||
Palaeotragus rouenii | MNHN PIK1692 | 432 | 22 | 5,09 | |
Palaeotragus rouenii | AMPG 1040/91 | 428 | 26 | 6,07 | |
Palaeotragus rouenii | BM M 11406 | 398 | Bohlin, 1926 | ||
Palaeotragus rouenii | 11406b | 32,8 | Churcher, 1970 | ||
Palaeotragus rouenii | BM a | 403 | 31,8 | 7,89 | Churcher, 1970 |
Bohlinia attica | AMNH 10453 | 710 | 48,87 | 6,88 | |
Bohlinia attica | M 11403 | 700 | 58,24 | 8,32 | |
Bohlinia attica | M 11401 | 50,12 | |||
Bohlinia attica | MNHN PIK27561 | 715 | 48 | 6,71 | |
Bohlinia attica | 710 | 47 | 6,62 | Geraads et al., 2005 | |
Bohlinia attica | VTK 80 | 661 | 45 | 6,81 | Geraads, 1979 |
Bohlinia attica | VTK 79 | 693 | 50 | 7,22 | Geraads, 1979 |
Bohlinia attica | 690 | Geraads, 1998 | |||
Bohlinia attica | 730 | 57 | 7,81 | Geraads et al., 2005 | |
Bohlinia attica | 700 | 60 | 8,57 | Geraads et al., 2005 | |
Bohlinia attica | 695 | 60,5 | 8,71 | Geraads et al., 2005 | |
Bohlinia attica | 50 | Geraads et al., 2005 | |||
Bohlinia attica | AMPG 1923/91 | 655 | 47 | 7,18 | |
Bohlinia attica | AMPG 1925/91 | 720 | 40 | 5,56 | |
Giraffa camelopardalis | AMNH 70016 | 693 | 45,74 | 6,60 | |
Giraffa camelopardalis | AMNH 70016 | 695 | 47,04 | 6,77 | |
Giraffa camelopardalis | AMNH 14135 | 622 | 44,09 | 7,09 | |
Giraffa camelopardalis | AMNH 14135 | 621 | 45,31 | 7,30 | |
Giraffa camelopardalis | MNCN-3439 | 605 | 39,61 | 6,55 | |
Giraffa camelopardalis | AMNH 53543 | 650 | 47,32 | 7,28 | |
Giraffa camelopardalis | AMNH 53550 | 753 | 54,09 | 7,18 | |
Giraffa camelopardalis | AMNH 53550 | 745 | 52,15 | 7,00 | |
Giraffa camelopardalis | AMNH 82001 | 730 | 57 | 7,81 | |
Giraffa camelopardalis | KNM OM 2278 | 667 | Harris, 1976 | ||
Giraffa camelopardalis | KNM OM 2269 | 735 | Harris, 1976 | ||
Giraffa camelopardalis | KNM OM 2272 | 624 | Harris, 1976 | ||
Giraffa camelopardalis | KNM OM 2217 | 738 | Harris, 1976 |
TABLE 2. Descriptive parameters of the metacarpal robustness index of the Giraffidae ( Max., maximum; Min., minimum; SD, standard deviation; Ntotal, total sample size).
RI | |||||
Max. | Mean | Min. | SD | N total | |
Birgerbohlinia schaubi | 15.91 | 13.54 | 12.64 | 1.36 | 5 |
Bohlinia attica | 8.71 | 7.31 | 5.56 | 0.96 | 11 |
Bramatherium megacephalum | 15.33 | 13.96 | 13.11 | 1.20 | 3 |
Canthumeryx sirtensis | 6.03 | 6.03 | 6.03 | 1 | |
Decennatherium pachecoi | 9.25 | 9.25 | 9.25 | 1 | |
Giraffa camelopardalis | 7.81 | 7.06 | 6.55 | 0.39 | 9 |
Helladotherium duvernoyi | 16.79 | 14.51 | 11.65 | 1.65 | 12 |
Okapia johnstoni | 10.70 | 10.31 | 9.58 | .29 | 14 |
Palaeotragus rouenii | 9.12 | 7.71 | 5.09 | 1.53 | 7 |
Samotherium major | 24.11 | 13.24 | 9.76 | 3.02 | 43 |
Sivatherium giganteum | 23.49 | 20.70 | 17.98 | 1.79 | 8 |
TABLE 3. Descriptive parameters of the metacarpal length of the Giraffidae (M ax., maximum; Min., minimum; SD, standard deviation; Ntotal, total sample size).
Length | |||||
Mean | Max. | Min. | SD | N total | |
Canthumeryx sirtensis | 303.50 | 317.00 | 290.0 | 19.09 | 2 |
Helladotherium duvernoyi | 424.79 | 470.00 | 395.00 | 19.56 | 29 |
Bramatherium megacephalum | 419.00 | 432.00 | 398.00 | 18.36 | 3 |
Sivatherium giganteum | 324.70 | 355.00 | 302.00 | 17.56 | 10 |
Decennatherium pachecoi | 391.00 | 391.00 | 391.00 | 1 | |
Birgerbohlinia schaubi | 380.20 | 391.00 | 373.00 | 7.26 | 5 |
Okapia johnstoni | 315.25 | 330.00 | 295.30 | 9.19 | 15 |
Samotherium major | 417.61 | 471.00 | 390.00 | 14.79 | 44 |
Palaeotragus rouenii | 414.13 | 434.00 | 398.00 | 14.71 | 8 |
Bohlinia attica | 698.25 | 730.00 | 655.00 | 22.17 | 12 |
Giraffa camelopardalis | 682.92 | 753.00 | 605.00 | 54.31 | 13 |
TABLE 4. Dimensions in mm of the metatarsal III-IV of Giraffidae ( DT, transversal diameter; Diaph., diaphysis; RI, robustness index).
Species | Specimen nº | Total Length | Diaph. TD | RI | Reference | |
Canthumeryx sirtensis | KNH MO 41 | 278 | 23 | 8.27 | ||
Canthumeryx sirtensis | KNM .M0.41B | 280 | 18.5 | 6.61 | Hamilton, 1978 | |
Canthumeryx sirtensis | IPP 1933-9 | 260 | 18 | 6.92 | Hamilton, 1978 | |
Canthumeryx sirtensis | BU 20117 | 349 | Hamilton, 1978 | |||
Canthumeryx sirtensis | UCB. V. 4899/42058 | 260 | 17 | 6.54 | Hamilton, 1978 | |
Helladotherium duvernoyi | MNHN PIK1564 | 450 | 48 | 10.67 | ||
Helladotherium duvernoyi | MNHN PIK1563 | 440 | 55 | 12.50 | ||
Helladotherium duvernoyi | MNHN PIK1565 | 435 | 40 | 9.20 | ||
Helladotherium duvernoyi | MNHN PIK1554 | 440 | 58 | 13.18 | ||
Helladotherium duvernoyi | M 11382 | 436 | 49.36 | 11.32 | ||
Helladotherium duvernoyi | M 11384 | 441 | 50.47 | 11.44 | ||
Helladotherium duvernoyi | M 8364 | 434 | 49.38 | 11.38 | ||
Helladotherium duvernoyi | M 11384 | 450 | 53.29 | 11.84 | ||
Helladotherium duvernoyi | M 11383 | 456 | 50.59 | 11.09 | ||
Helladotherium duvernoyi | M 11383 | 456 | 50.59 | 11.09 | ||
Helladotherium duvernoyi | M 11381 | 435 | 62.57 | 14.38 | ||
Helladotherium duvernoyi | M 11381 | 435 | 62.57 | 14.38 | ||
Helladotherium duvernoyi | 1972/10 | 458 | 56 | 12.23 | Melentis, 1974 | |
Helladotherium duvernoyi | MNHN PIK1566 | 455 | 55 | 12.09 | ||
Helladotherium duvernoyi | AMNH 27805 | 495 | 60 | 12.12 | ||
Helladotherium duvernoyi | MTLA85 | 480 | 52 | 10.83 | Kostopoulos, 2009 | |
Helladotherium duvernoyi | BM M 11382 | 440 | Bohlin, 1926 | |||
Helladotherium duvernoyi | RMS | 465 | Bohlin, 1926 | |||
Helladotherium duvernoyi | 440 | Pilgrim, 1911 | ||||
Helladotherium duvernoyi | BM M 11381 | 440 | Bohlin, 1926 | |||
Bramatherium megacephalum | AMNH 19770 | 452 | 50.6 | 11.19 | ||
Bramatherium megacephalum | AMNH 19464 | 382 | 40.64 | 10.64 | ||
Bramatherium megacephalum | AMNH 19831 | 442 | 42.92 | 9.71 | ||
Bramatherium megacephalum | AMNH 19688 | 421 | 52.27 | 12.42 | ||
Bramatherium megacephalum | 465 | 61.5 | 13.23 | Geraads & Güleç, 1999 | ||
Sivatherium giganteum | OR 39752 | 410 | 52.77 | 12.87 | ||
Sivatherium giganteum | OR 39752 | 410 | 52.77 | 12.87 | ||
Sivatherium giganteum | OR 17089 | 56.2 | ||||
Sivatherium giganteum | OR 17090 | 59.84 | ||||
Sivatherium giganteum | OR 39753 | 58.73 | ||||
Sivatherium giganteum | OR 17090 | 59.84 | ||||
Sivatherium giganteum | OR 39753 | 58.73 | ||||
Sivatherium giganteum | OR 17089 | 56.2 | ||||
Sivatherium giganteum | 415 | Bohlin, 1926 | ||||
Decennatherium pachecoi | MNCN-42764 | 427.5 | 35.35 | 8.27 | ||
Decennatherium pachecoi | MNCN-42765 | 406.14 | 42.46 | 10.45 | ||
Decennatherium pachecoi | MNCN-42771 | 38.4 | ||||
Decennatherium pachecoi | MNCN-42778 | 34.94 | ||||
Decennatherium pachecoi | MNCN-42770 | 36.17 | ||||
Decennatherium pachecoi | MNCN-42766 | 36.59 | ||||
Decennatherium pachecoi | MNCN-42767 | 35.88 | ||||
Decennatherium pachecoi | MNCN-42768 | 33.38 | ||||
Decennatherium pachecoi | 40 | Crusafont, 1952 | ||||
Birgerbohlinia schaubi | IPS-5015 | 48.51 | ||||
Birgerbohlinia schaubi | IPS-4981 | 47.48 | ||||
Birgerbohlinia schaubi | Nº48 | 410 | Crusafont, 1952 | |||
Birgerbohlinia schaubi | Nº183 | 384 | 46.5 | 12.11 | Crusafont, 1952 | |
Birgerbohlinia schaubi | IPS-5090 | 48.08 | ||||
Okapia johnstoni | AMNH 51218 | 330 | 32 | 9.70 | ||
Okapia johnstoni | AMNH 51903 | 325 | 30.6 | 9.42 | ||
Okapia johnstoni | AMNH 51903 | 324 | 29.98 | 9.25 | ||
Okapia johnstoni | AMNH 51198 | 327 | 29.62 | 9.06 | ||
Okapia johnstoni | AMNH 51200 | 322 | 27.73 | 8.61 | ||
Okapia johnstoni | AMNH 51196 | 319 | 30.62 | 9.60 | ||
Okapia johnstoni | AMNH 51196 | 318 | 30.63 | 9.63 | ||
Okapia johnstoni | AMNH 51200 | 321 | 28.15 | 8.77 | ||
Okapia johnstoni | AMNH 2268 | 332 | 31.89 | 9.61 | ||
Okapia johnstoni | AMNH 51197 | 335 | 31.6 | 9.43 | ||
Okapia johnstoni | AMNH 51197 | 334 | 31.6 | 9.46 | ||
Okapia johnstoni | AMNH 51198 | 326 | 29.6 | 9.08 | ||
Okapia johnstoni | MNCN-11425 | 317 | 30.34 | 9.57 | ||
Okapia johnstoni | MA(6) O (I) | 316.2 | 29.3 | 9.27 | Churcher, 1970 | |
Samotherium major | PIM350 | 465 | 50 | 10.75 | Kostopoulos, 2009 | |
Samotherium major | PIM344 | 460 | 49 | 10.65 | Kostopoulos, 2009 | |
Samotherium major | MGL S95 | 473 | 48.7 | 10.30 | Kostopoulos, 2009 | |
Samotherium major | AMNH20636 Q1 | 495 | 48 | 9.70 | Kostopoulos, 2009 | |
Samotherium major | MGL S94 | 465 | 51 | 10.97 | Kostopoulos, 2009 | |
Samotherium major | AMNH28845B Q4 | 480 | 55.1 | 11.48 | Kostopoulos, 2009 | |
Samotherium major | PIM360 | 470 | 48 | 10.21 | Kostopoulos, 2009 | |
Samotherium major | AMNH20615 Q1 | 447 | 49.7 | 11.12 | Kostopoulos, 2009 | |
Samotherium major | AMNH20633 Q1 | 500 | 56.4 | 11.28 | Kostopoulos, 2009 | |
Samotherium major | PIM359 | 460 | Kostopoulos, 2009 | |||
Samotherium major | AMNH22697 Q5 | 465 | 56.2 | 12.09 | Kostopoulos, 2009 | |
Samotherium major | AMNH28845A Q4 | 470 | 53.1 | 11.30 | Kostopoulos, 2009 | |
Samotherium major | AMNH20595 Q1 | 465 | 53.5 | 11.51 | Kostopoulos, 2009 | |
Samotherium major | MTLA279 | 503 | 55 | 10.93 | Kostopoulos, 2009 | |
Samotherium major | MTLB401 | 475 | 50 | 10.53 | Kostopoulos, 2009 | |
Samotherium major | MTLA387 | 463 | 57.6 | 12.44 | Kostopoulos, 2009 | |
Samotherium major | MTLA247 | 495 | 59.3 | 11.98 | Kostopoulos, 2009 | |
Samotherium major | PIM346 | 475 | 57.7 | 12.15 | Kostopoulos, 2009 | |
Samotherium major | MTLA72 | 480 | 47.4 | 9.88 | Kostopoulos, 2009 | |
Samotherium major | SLQ690 | 460 | Kostopoulos, 2009 | |||
Samotherium major | PIM48 | 460 | 54 | 11.74 | Kostopoulos, 2009 | |
Samotherium major | MTLA470 | 475 | 52.1 | 10.97 | Kostopoulos, 2009 | |
Samotherium major | AMNH22966 Q5 | 450 | 54.6 | 12.13 | Kostopoulos, 2009 | |
Samotherium major | PIM352 | 430 | Kostopoulos, 2009 | |||
Samotherium major | PIM355 | 447 | 49.5 | 11.07 | Kostopoulos, 2009 | |
Samotherium major | MTLA342 | 56.4 | Kostopoulos, 2009 | |||
Samotherium major | MGL S382b | 47 | Kostopoulos, 2009 | |||
Samotherium major | MGL S1063 | 52 | Kostopoulos, 2009 | |||
Samotherium major | MAR571 | 420 | 45 | 10.71 | ||
Samotherium major | KTA134 | 462 | 53 | 11.47 | Geraads, 1994 | |
Palaeotragus rouenii | PIK1691 | 420 | 25 | 5.95 | ||
Palaeotragus rouenii | 445 | 32.8 | 7.37 | Kostopoulos, 2009 | ||
Palaeotragus rouenii | 32.5 | Kostopoulos, 2009 | ||||
Palaeotragus rouenii | 30 | Kostopoulos, 2009 | ||||
Palaeotragus rouenii | 377 | 30 | 7.96 | Geraads, 1994 | ||
Bohlinia attica | MNHN PIK2357 | 714 | 50 | 7.00 | ||
Bohlinia attica | M 11403 | 52.97 | ||||
Bohlinia attica | M 11403 (6) | 49.36 | ||||
Bohlinia attica | M 11403 | 49.4 | ||||
Bohlinia attica | M 11403 | 48.33 | ||||
Bohlinia attica | 690 | Geraads et al., 2005 | ||||
Bohlinia attica | MNHN SLQ682 | 709 | 48 | 6.77 | ||
Bohlinia attica | 680 | 46 | 6.76 | Geraads et al., 2005 | ||
Bohlinia attica | NKT-24 | 670 | 55.5 | 8.28 | Kostopoulos et al., 1996 | |
Bohlinia attica | NKT-171 | 680 | 51.8 | 7.62 | Kostopoulos et al., 1996 | |
Bohlinia attica | AUT | 658 | 49 | 7.45 | Geraads et al., 2005 | |
Bohlinia attica | 695 | 52.5 | 7.55 | Geraads et al., 2005 | ||
Bohlinia attica | FM2025 | 56 | Geraads et al., 2005 | |||
Bohlinia attica | MNHN MAR3254 | 692 | 50 | 7.23 | ||
Bohlinia attica | 620 | Mecquenem, 1924 | ||||
Bohlinia attica | 675 | 52 | 7.70 | Geraads et al., 2005 | ||
Bohlinia attica | AMNH 10453 | 702 | 48.43 | 6.90 | ||
Bohlinia attica | AMPG 1972/63 | 680 | 49 | 7.21 | ||
Giraffa camelopardalis | AMNH 82001 | 742 | 52 | 7.01 | ||
Giraffa camelopardalis | AMNH 53543 | 658 | 45 | 6.84 | ||
Giraffa camelopardalis | MNCN-3439 | 638 | 39.24 | 6.15 | ||
Giraffa camelopardalis | AMNH 14135 | 625 | 39.32 | 6.29 | ||
Giraffa camelopardalis | AMNH 53550 | 765 | 50.86 | 6.65 | ||
Giraffa camelopardalis | AMNH 14135 | 626 | 40.02 | 6.39 | ||
Giraffa camelopardalis | AMNH 53550 | 764 | 49.52 | 6.48 | ||
Giraffa camelopardalis | AMNH 70016 | 685 | 45.43 | 6.63 | ||
Giraffa camelopardalis | AMNH 70016 | 686 | 44.75 | 6.52 | ||
Giraffa camelopardalis | OM 2278 | 673 | Harris, 1976 | |||
Giraffa camelopardalis | 660 | Harris, 1976 | ||||
Giraffa camelopardalis | OM 2272 | 632 | Harris, 1976 | |||
Giraffa camelopardalis | OM 2217 | 732 | Harris, 1976 | |||
Giraffa camelopardalis | 755 | Harris, 1976 | ||||
Giraffa camelopardalis | OM 2269 | 748 | Harris, 1976 | |||
Giraffa camelopardalis | 755 | 45 | 5.96 | Harris, 1976 |
TABLE 5. Descriptive parameters of the metatarsal robustness index of the Giraffidae ( Max., maximum; Min., minimum; SD, standard deviation; Ntotal, total sample size).
RI | |||||
Max. | Mean | Mín. | SD | N total | |
Birgerbohlinia schaubi | 12.11 | 12.11 | 12.11 | 1 | |
Bohlinia attica | 8.28 | 7.32 | 6.76 | .46 | 11 |
Bramatherium megacephalum | 13.23 | 11.44 | 9.71 | 1.40 | 5 |
Canthumeryx sirtensis | 8.27 | 7.09 | 6.54 | .81 | 4 |
Decennatherium pachecoi | 10.45 | 9.36 | 8.27 | 1.55 | 2 |
Giraffa camelopardalis | 7.01 | 6.49 | 5.96 | .31 | 10 |
Helladotherium duvernoyi | 14.38 | 11.86 | 9.20 | 1.33 | 16 |
Okapia johnstoni | 9.70 | 9.32 | 8.61 | .33 | 14 |
Palaeotragus rouenii | 7.96 | 7.09 | 5.95 | 1.03 | 3 |
Samotherium major | 12.44 | 11.14 | 9.70 | .73 | 24 |
Sivatherium giganteum | 12.87 | 12.87 | 12.87 | 0.00 | 1 |
TABLE 6. Descriptive parameters of the metatarsal length of the Giraffidae ( Max., maximum; Min., minimum; SD, standard deviation; Ntotal, total sample size).
Length | |||||
Mean | Max. | Min. | SD | N total | |
Canthumeryx sirtensis | 285 .4 | 349 | 260 | 36 .81 | 5 |
Helladotherium duvernoyi | 449 .05 | 495 | 434 | 16 .22 | 20 |
Bramatherium megacephalum | 432 .4 | 465 | 382 | 32 .44 | 5 |
Sivatherium giganteum | 411 .67 | 415 | 410 | 2 .89 | 3 |
Decennatherium pachecoi | 416 .82 | 427 .5 | 406 .14 | 15 .1 | 2 |
Birgerbohlinia schaubi | 397 | 410 | 384 | 18 .38 | 2 |
Okapia johnstoni | 324 .73 | 335 | 316 .2 | 6 .27 | 14 |
Samotherium major | 467 .04 | 503 | 420 | 19 .07 | 27 |
Palaeotragus rouenii | 414 | 445 | 377 | 34 .39 | 3 |
Bohlinia attica | 681 .92 | 714 | 620 | 24 .34 | 13 |
Giraffa camelopardalis | 696 .5 | 765 | 625 | 53 .76 | 16 |
First comprehensive morphological analysis on the metapodials of Giraffidae
Plain Language Abstract
Giraffids are a group of ruminants with only two living members, the giraffe and the okapi. This family, however, was much more diverse in the past, with more than 20 species that were variable in body size, neck length, and skeletal features. We describe characteristics on the limbs of living and extinct giraffids, and find that each species exhibits a unique combination of features. Our forelimb and hindlimb descriptions allow for genus identification based on a limb specimen, and for comparisons between the different taxa. We find both phylogenetic and environmental signals in the limb anatomical features, allowing for studies involving cladistics as well as paleoecology. We also evaluate measurements of the limbs, and find that most species exhibit a front-loaded body weight distribution, consistent with the elongated necks and robust ossicones. The morphological features of the giraffid metapodials, as well as the limb lengths and proportions prove to be a useful tool for genus identifications and comparisons.
Resumen en Español
Primer análisis morfológico exhaustivo sobre los metapodiales de Giraffidae
Los jiráfidos son un grupo de rumiantes del grupo Pecora relictos con sólo dos taxones vivos. Durante el Mioceno, sin embargo, este grupo fue mucho más diverso, con más de 20 especies diferentes que mostraban una amplia gama de variabilidad. Además de en muchas otras partes del esqueleto, esta variabilidad también se muestra en sus metapodiales. Encontramos diferencias anatómicas inter-específicas en los metapodiales de la familia Giraffidae; cada taxón evaluado posee una combinación única de morfologías de las extremidades. La superficie metapodial próximo-palmar/plantar proporciona características útiles y permite identificaciones y comparaciones a nivel de género. Describimos el canal central del eje metapodial; cuando se combina con la longitud total de la extremidad, la profundidad de esta depresión permite una mejor separación entre taxones. Encontramos que el índice de robustez de los metacarpianos supera al de los metatarsianos en todos los jiráfidos evaluados excepto en uno, soportando una distribución de peso corporal con carga frontal, consistente con cervicales alargadas o grandes osiconos observados en muchos taxones. Las características morfológicas de los metapodiales de los jiráfidos, así como las longitudes y proporciones de las extremidades, pueden constituir una herramienta útil para el análisis filogenético.
Palabras clave: Giraffidae; Metapodial; Morfología; Extremidades; Ruminantia; Anatomía
Traducción: Enrique Peñalver (Sociedad Española de Paleontología)
Résumé en Français
Première analyse globale de la morphologie des métapodes de Giraffidae
Les giraffidés sont un groupe relique de ruminants Pecora avec seulement deux taxons actuels. Cependant, pendant le Miocène, ce groupe était beaucoup plus diversifié, avec plus de 20 espèces différentes montrant une grande variabilité de nombreuses parties du squelette, notamment les métapodes. Nous décrivons des différences anatomiques inter-spécifiques sur les métapodes de giraffidés. Chaque taxon évalué possède une combinaison unique de caractères morphologiques des membres. La surface proximo-palmaire/plantaire des métapodes fournit des caractères utiles permettant d'identifier et de comparer les genres. Nous décrivons les différences au niveau du sillon longitudinal de la diaphyse du métapode. Combinée avec la longueur absolue du membre, la profondeur de ce sillon permet de mieux distinguer les taxons. Nous observons que l'indice de robustesse des métacarpes dépasse celui des métatarses chez tous les giraffidés évalués, à l'exception d'un taxon. Cela suggère une distribution de la masse corporelle plus importante vers l'avant, ce qui est cohérent avec les cervicales allongées et les ossicônes de grande taille observés chez de nombreux taxons. Les caractères morphologiques des métapodes de giraffidés, ainsi que les longueurs et les proportions des membres, peuvent être utiles pour les analyses phylogénétiques.
Mots-clés : Giraffidae ; métapodes ; morphologie ; membres ; Ruminantia ; anatomie
Translator: Antoine Souron
Deutsche Zusammenfassung
Die erste umfassende morphologische Analyse der Giraffidae-Metapodien
Giraffiden sind eine Gruppe reliktischer Stirnwaffenträger-Ruminanten mit nur zwei heute noch lebenden Taxa. Während des Miozäns war diese Gruppe jedoch viel diverser und zeigte mit mehr als 20 unterschiedlichen Arten eine große Variabilität. Neben vielen anderen Skelettteilen spiegelt sich diese Variabilität auch in den Metapodien dieser Tiere wieder. Wir finden interspezifische anatomische Unterschiede bei giraffiden Metapodien; jedes evaluierte Taxon besitzt eine einzigartige Kombination seiner Beinmorphologie. Die proximo-palmar/plantare Oberfläche bietet sinnvolle Merkmale und erlaubt Gattungsidentifikation und Vergleiche. Wir beschreiben den zentralen Trog des Metapodium-Schafts; eine Kombination der Trogtiefe mit der absoluten Beinlänge erlaubt eine bessere Teilung der Taxa. Wir fanden heraus, dass der Metacarpale-Robustheits-Index den Metatarsale-Robustheits-Index bei allen bis auf einer evaluierten Giraffide übertrifft. Dies unterstützt eine im vorderen Körperbereich konzentrierte Gewichtsverteilung, was mit den verlängerten Halswirbeln oder den großen Ossicones mancher Taxa einhergeht. Die morphologischen Merkmale der giraffiden Metapodien können zusammen mit den Beinlängen-und Proportionen ein brauchbares Werkzeug für die phylogenetische Analyse sein.
Schlüsselwörter: Giraffidae; Metapodium; Morphologie; Beine; Ruminantia; Anatomie
Translator: Eva Gebauer
Arabic
Translator: Ashraf M.T. Elewa
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Review: The Princeton Field Guide to Mesozoic Sea Reptiles
The Princeton Field Guide to Mesozoic Sea Reptiles
Article number: 26.1.1R
April 2023