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First comprehensive morphological analysis on the metapodials of Giraffidae

María Ríos, Melinda Danowitz, and Nikos Solounias

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

 

 

riosMaría Ríos. Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-Consejo Superior de Investigaciones Científicas, 2 Gutiérrez Abascal, Madrid, 28006, Spain. This email address is being protected from spambots. You need JavaScript enabled to view it.

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).

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danowitzMelinda Danowitz. Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine, 8000 Northern Boulevard, Old Westbury, NY 11568, USA. This email address is being protected from spambots. You need JavaScript enabled to view it.

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.

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solouniasNikos 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. This email address is being protected from spambots. You need JavaScript enabled to view it.

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.

 figure1

FIGURE 2. Measurements applied to the giraffid metapodials, based on those proposed by Quiralte (2011), using the same specimen as in figure 1.

 figure2

 

FIGURE 3. Helladotherium duvernoyi from Pikermi. 1, left metacarpal (reversed) MNHN 1658; 2, right metatarsal MNHN 1564.

 figure3

FIGURE 4. Three selected metacarpal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.

 figure4

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.

 figure5

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.

 figure6

FIGURE 7. Three additional metacarpal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.

 figure7

FIGURE 8. Bramatherium megacephalum from the Upper Siwaliks. 1, Right metacarpal AMNH 29820; 2, Right metatarsal AMNH 19688.

 figure8

FIGURE 9. Sivatherium giganteum from the Upper Siwaliks. 1, Right metacarpal NHM UK 39533; 2, Left metatarsal (reversed) NHM UK 39752.

figure9 

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.

 figure10

FIGURE 11. Samotherium major from Samos. 1, Right metacarpal NHMBe 711; 2, Right metatarsal AMNH 22967.

 figure11

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.

 figure12

FIGURE 13. Bohlinia attica from Pikermi. 1, Right metacarpal MNHN 52780; 2, Right metatarsal MNHN 1639.

figure13 

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.

 figure14

FIGURE 15. Giraffa camelopardalis from Kenya. 1, Left metacarpal (reversed) AMNH 53543; 2, Left metatarsal (reversed) AMNH 53550.

 figure15

FIGURE 16. Three selected metatarsal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.

 figure16

FIGURE 17. Three selected metatarsal characteristics. This simplified table can be used to facilitate specimen identifications and differentiate between different giraffid taxa.

figure17 

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.

 figure18

FIGURE 19. Box-and-whiskers plot with the metacarpal RI of the Giraffidae analyzed.

 figure19

FIGURE 20. Dispersion plot with the absolute metacarpal length of the Giraffidae analyzed.

 figure20

FIGURE 21. Dispersion plot showing the total length of the metacarpals versus the midshaft minimum transverse diameter of the Giraffidae analyzed.

 figure21

FIGURE 22. Box-and-whiskers plot with the metatarsal RI of the Giraffidae analyzed.

 figure22

FIGURE 23. Dispersion plot with the absolute metatarsal length of the Giraffidae analyzed.

figure23 

FIGURE 24. Dispersion plot showing the total length of the metatarsals versus the midshaft minimum transverse diameter of the Giraffidae analyzed.

figure24

 

 

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