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. 2017 May 2:5:e3217.
doi: 10.7717/peerj.3217. eCollection 2017.

The earliest known titanosauriform sauropod dinosaur and the evolution of Brachiosauridae

Affiliations

Affiliations

  • 1 Department of Earth Science and Engineering, Imperial College London, London, United Kingdom.
  • 2 Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements, Museum National d'Histoire Naturelle, Paris, France.
  • 3 Laboratoire de Géographie Physique: Environnements Quaternaires et Actuels, CNRS/Université Paris 1 Panthéon-Sorbonne/UPEC, Paris, France.

The earliest known titanosauriform sauropod dinosaur and the evolution of Brachiosauridae

Philip D Mannion et al. PeerJ. .
. 2017 May 2:5:e3217.
doi: 10.7717/peerj.3217. eCollection 2017.

Affiliations

  • 1 Department of Earth Science and Engineering, Imperial College London, London, United Kingdom.
  • 2 Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements, Museum National d'Histoire Naturelle, Paris, France.
  • 3 Laboratoire de Géographie Physique: Environnements Quaternaires et Actuels, CNRS/Université Paris 1 Panthéon-Sorbonne/UPEC, Paris, France.

Abstract

Brachiosauridae is a clade of titanosauriform sauropod dinosaurs that includes the well-known Late Jurassic taxa Brachiosaurus and Giraffatitan. However, there is disagreement over the brachiosaurid affinities of most other taxa, and little consensus regarding the clade's composition or inter-relationships. An unnamed partial sauropod skeleton was collected from middle-late Oxfordian (early Late Jurassic) deposits in Damparis, in the Jura department of eastern France, in 1934. Since its brief description in 1943, this specimen has been informally known in the literature as the 'Damparis sauropod' and 'French Bothriospondylus', and has been considered a brachiosaurid by most authors. If correctly identified, this would make the specimen the earliest known titanosauriform. Coupled with its relatively complete nature and the rarity of Oxfordian sauropod remains in general, this is an important specimen for understanding the early evolution of Titanosauriformes. Full preparation and description of this specimen, known from teeth, vertebrae and most of the appendicular skeleton of a single individual, recognises it as a distinct taxon: Vouivria damparisensis gen. et sp. nov. Phylogenetic analysis of a data matrix comprising 77 taxa (including all putative brachiosaurids) scored for 416 characters recovers a fairly well resolved Brachiosauridae. Vouivria is a basal brachiosaurid, confirming its status as the stratigraphically oldest known titanosauriform. Brachiosauridae consists of a paraphyletic array of Late Jurassic forms, with Europasaurus, Vouivria and Brachiosaurus recovered as successively more nested genera that lie outside of a clade comprising (Giraffatitan + Sonorasaurus) + (Lusotitan + (Cedarosaurus + Venenosaurus)). Abydosaurus forms an unresolved polytomy with the latter five taxa. The Early Cretaceous South American sauropod Padillasaurus was previously regarded as a brachiosaurid, but is here placed within Somphospondyli. A recent study contended that a number of characters used in a previous iteration of this data matrix are 'biologically related', and thus should be excluded from phylogenetic analysis. We demonstrate that almost all of these characters show variation between taxa, and implementation of sensitivity analyses, in which these characters are excluded, has no effect on tree topology or resolution. We argue that where there is morphological variation, this should be captured, rather than ignored. Unambiguous brachiosaurid remains are known only from the USA, western Europe and Africa, and the clade spanned the Late Jurassic through to the late Albian/early Cenomanian, with the last known occurrences all from the USA. Regardless of whether their absence from the Cretaceous of Europe, as well as other regions entirely, reflects regional extinctions and genuine absences, or sampling artefacts, brachiosaurids appear to have become globally extinct by the earliest Late Cretaceous.

Keywords: Biogeography; Brachiosauridae; Character correlation; Cretaceous; France; Gondwana; Late Jurassic; Laurasia; Mesozoic; Ontogeny; Oxfordian; Sacral fusion.

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Conflict of interest statement

The authors declare there are no competing interests.

Figures

Figure 1

Figure 1. Simplified cladograms summarizing previous hypotheses…

Figure 1. Simplified cladograms summarizing previous hypotheses of placements of putative brachiosaurid taxa.

The four…

Figure 1. Simplified cladograms summarizing previous hypotheses of placements of putative brachiosaurid taxa.
The four cladograms are based on: (A) Royo-Torres et al. ( (using the dataset of Upchurch, Barrett & Dodson, 2004)); (B) Carballido et al. ( (using the complete strict consensus tree presented in their supplementary information)); (C) D’Emic, Foreman & Jud (2016); (D) Poropat et al. ( (using the complete strict consensus tree presented in their supplementary information)). Brachiosauridae is highlighted in red in each tree.
Figure 2

Figure 2. Location map of the quarry…

Figure 2. Location map of the quarry south-eastward of Damparis village and exact location of…

Figure 2. Location map of the quarry south-eastward of Damparis village and exact location of the palaeontological locus.
The red cross marks the place where the bones and teeth of the dinosaur were found. To the right of the cross, the path slightly turns left following the curvature of the original cliff visible in Dorlodot (: fig. 1) and Lapparent (: pl.V, fig. 1). The letter ‘T’ marks the place of the truncation (see Fig. 4). The background map is modified from http://www.geoportail.gouv.fr/ and is freely usable (http://professionnel.ign.fr/enseignement-recherche), as is the inset map of France (http://www.pacha-cartographie.com/fonds-de-carte/).
Figure 3

Figure 3. Wall constructed to allow for…

Figure 3. Wall constructed to allow for the refilling of the excavation and passage of…

Figure 3. Wall constructed to allow for the refilling of the excavation and passage of the small train on the overlying step during the excavation (A) and today (B).
The surrounding cliff did not progress since the 1930’s and marks the south-eastern limit of the quarry.
Figure 4

Figure 4. Main unconformity of the sequence…

Figure 4. Main unconformity of the sequence between the calcaires fins (above) from the calcaires…

Figure 4. Main unconformity of the sequence between the calcaires fins (above) from the calcaires massifs crayeux bioclastiques (below).
It is marked by a sedimentological truncation of the underlying undulating chalky limestone banks and underlined by a 2 cm-thick red clayey bed, the lowermost one of the sequence (Photo: O Moine). See Fig. 2 for its location in the quarry.
Figure 5

Figure 5. Original excavation plan of Vouivria…

Figure 5. Original excavation plan of Vouivria damparisensis n. gen. n. sp. (modified from Dorlodot,…

Figure 5. Original excavation plan of Vouivria damparisensis n. gen. n. sp. (modified from Dorlodot, 1934).
Figure 6

Figure 6. The locus of the discovery…

Figure 6. The locus of the discovery during the excavation (A) and today (B).

(1)…

Figure 6. The locus of the discovery during the excavation (A) and today (B).
(1) Upper banks of the calcaires massifs crayeux bioclastiques; (2) basal clayey bed separating the calcaires massifs crayeux bioclastiques (below) from the calcaires fins (above); (3) upper limit of the basal submetric limestone bank of the calcaires fins underlining the stone wall that now masks the locus of the discovery; (4) clayey bed that laterally connected with the fossil discovery; (5) southern edge of the depression infilling; (6) location of the locus of the fossil discovery; (7) upper limit of the submetric limestone bank overlying the infilling of the depression; (8) waste material and overlying railway mentioned in the original description, but today both removed (Photo: O Moine).
Figure 7

Figure 7. Chronostratigraphical position of the Damparis…

Figure 7. Chronostratigraphical position of the Damparis fossils.

Cyclostratigraphy and ammonites zones according to Hardenbol…

Figure 7. Chronostratigraphical position of the Damparis fossils.
Cyclostratigraphy and ammonites zones according to Hardenbol et al. (1998), and chronology based on Cohen et al. (2013). Sources for chronostratigraphical ranges of ammonites are given in Table S2. Dark and light blue dashed lines link maximum flooding surfaces of Tethyan and Boreal domains, respectively. Damparis and regional stratigraphic formations are separated by dashed lines, as their exact position in the ammonite biostratigraphic scheme remain to be established.
Figure 8

Figure 8. Stratigraphic column of the Damparis…

Figure 8. Stratigraphic column of the Damparis quarry.

Figure 8. Stratigraphic column of the Damparis quarry.
Figure 9

Figure 9. Teeth of Vouivria damparisensis (MNHN.F.1934.6…

Figure 9. Teeth of Vouivria damparisensis (MNHN.F.1934.6 DAM1–DAM 5).

(A–E) labial views, (F–J) mesial views,…

Figure 9. Teeth of Vouivria damparisensis (MNHN.F.1934.6 DAM1–DAM 5).
(A–E) labial views, (F–J) mesial views, (K–O) lingual views; (A, F, K) DAM 1, (B, G, L) DAM 2, (C, H, M) DAM 3, (D, I, N) DAM 4, (E, J, O) DAM 5. Scale bar equals 20 cm.
Figure 10

Figure 10. Middle cervical vertebra of Vouivria…

Figure 10. Middle cervical vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 6).

(A) Left lateral view,…

Figure 10. Middle cervical vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 6).
(A) Left lateral view, (B) anterior view, (C) right lateral view; (D) posterior view. Abbreviations: cpol, centropostzygapophyseal lamina; di, diapophysis; cprl, centroprezygapophyseal lamina; espol, expanded spinopostzygapophyseal lamina; no, notch; ns, notch; pa, parapophysis; pn, pneumatic foramen; pocdf, postzygapophyseal centrodiapophyseal fossa; podl, postzygodiapophyseal lamina; poz, postzygapophysis; prz, prezygapophysis; ri, ridge; sdf, spinodiapophyseal fossa; spol, spinopostzygapophyseal lamina; sprl, spinoprezygapophyseal lamina. Scale bar equals 10 cm.
Figure 11

Figure 11. Posterior cervical vertebra of Vouivria…

Figure 11. Posterior cervical vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 7).

(A) Left lateral view,…

Figure 11. Posterior cervical vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 7).
(A) Left lateral view, (B) right lateral view, (C) anterior view; (D) posterior view; (E) dorsal view. Abbreviations: acdl, anterior centrodiapophyseal lamina; cprf, centroprezygapophyseal fossa; cprl, centroprezygapophyseal lamina; nc, neural canal; pa, parapophysis; pcdl, posterior centrodiapophyseal lamina; pn, pneumatic foramen; ri, ridge; tprl, intraprezygapophyseal lamina. Scale bar equals 5 cm.
Figure 12

Figure 12. Anterior dorsal vertebra of Vouivria…

Figure 12. Anterior dorsal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 8).

(A) Left lateral view,…

Figure 12. Anterior dorsal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 8).
(A) Left lateral view, (B) right lateral view, (C) anterior view; (D) ventral view. Scale bar equals 5 cm.
Figure 13

Figure 13. Posterior middle dorsal vertebra of

Figure 13. Posterior middle dorsal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 9).

(A) Right lateral…

Figure 13. Posterior middle dorsal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 9).
(A) Right lateral view, (B) anterior view, (C) posterior view. Abbreviations: cprl/acpl, centroprezygapophyseal lamina or anterior centroparapophyseal lamina; nc, neural canal; pcdl, posterior centrodiapophyseal lamina; pn, pneumatic foramen; pocdf, posterior centrodiapophyseal fossa. Scale bar equals 5 cm.
Figure 14

Figure 14. Sacrum of Vouivria damparisensis (MNHN.F.1934.6…

Figure 14. Sacrum of Vouivria damparisensis (MNHN.F.1934.6 DAM 32).

(A) Dorsal view; (B) ventral view.…

Figure 14. Sacrum of Vouivria damparisensis (MNHN.F.1934.6 DAM 32).
(A) Dorsal view; (B) ventral view. Abbreviations: sv, sacral vertebrae. Scale bar equals 10 cm.
Figure 15

Figure 15. Anterior caudal vertebra of Vouivria…

Figure 15. Anterior caudal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 10).

(A) Right lateral view;…

Figure 15. Anterior caudal vertebra of Vouivria damparisensis (MNHN.F.1934.6 DAM 10).
(A) Right lateral view; (B) posterior view; (C) anterior view. Abbreviations: acdl, anterior centrodiapophyseal lamina; cprl, centroprezygapophyseal lamina; di, diapophysis; pcdl, posterior centrodiapophyseal lamina; prdl, prezygodiapophyseal lamina. Scale bar equals 10 cm.
Figure 16

Figure 16. Left scapula of Vouivria damparisensis

Figure 16. Left scapula of Vouivria damparisensis (MNHN.F.1934.6 DAM 25).

(A) Lateral view; (B) medial…

Figure 16. Left scapula of Vouivria damparisensis (MNHN.F.1934.6 DAM 25).
(A) Lateral view; (B) medial view. Abbreviations: ac, acromion; acr, acromial ridge; avp, acromial ventral process; gl, glenoid. Scale bar equals 10 cm.
Figure 17

Figure 17. Right scapula of Vouivria damparisensis

Figure 17. Right scapula of Vouivria damparisensis (MNHN.F.1934.6 DAM 26).

(A) Lateral view; (B) medial…

Figure 17. Right scapula of Vouivria damparisensis (MNHN.F.1934.6 DAM 26).
(A) Lateral view; (B) medial view. Abbreviations: ac, acromion; acr, acromial ridge; avp, acromial ventral process; gl, glenoid. Scale bar equals 10 cm.
Figure 18

Figure 18. Right coracoid of Vouivria damparisensis

Figure 18. Right coracoid of Vouivria damparisensis (MNHN.F.1934.6 DAM 27).

(A) Lateral view; (B) medial…

Figure 18. Right coracoid of Vouivria damparisensis (MNHN.F.1934.6 DAM 27).
(A) Lateral view; (B) medial view. Abbreviations: gl, glenoid. Scale bar equals 10 cm.
Figure 19

Figure 19. Right humerus of Vouivria damparisensis

Figure 19. Right humerus of Vouivria damparisensis (MNHN.F.1934.6 DAM 28).

(A) Anterior view; (B) lateral…

Figure 19. Right humerus of Vouivria damparisensis (MNHN.F.1934.6 DAM 28).
(A) Anterior view; (B) lateral view; (C) posterior view; (D) proximal view (anterior towards top); (E) distal view (anterior towards top). Abbreviations: co, condyle; dpc, deltopectoral crest; ri, ridge. Scale bar equals 10 cm.
Figure 20

Figure 20. Right ulna of Vouivria damparisensis

Figure 20. Right ulna of Vouivria damparisensis (MNHN.F.1934.6 DAM 29).

(A) Anterior view; (B) lateral…

Figure 20. Right ulna of Vouivria damparisensis (MNHN.F.1934.6 DAM 29).
(A) Anterior view; (B) lateral view; (C) posterior view; (D) medial view; (E) proximal view (anterior margin towards top); (F) distal view (anterior margin towards top). Abbreviations: amp, anteromedial process; alp, anterolateral process; icr, interosseous crest; ol, olecranon. Scale bar equals 10 cm.
Figure 21

Figure 21. Left ulna of Vouivria damparisensis

Figure 21. Left ulna of Vouivria damparisensis (MNHN.F.1934.6 DAM 30).

(A) Anterior view; (B) lateral…

Figure 21. Left ulna of Vouivria damparisensis (MNHN.F.1934.6 DAM 30).
(A) Anterior view; (B) lateral view; (C) posterior view; (D) medial view; (E) proximal view (anterior margin towards top). Abbreviations: amp, anteromedial process; alp, anterolateral process; icr, interosseous crest; ol, olecranon. Scale bar equals 10 cm.
Figure 22

Figure 22. Right carpal of Vouivria damparisensis

Figure 22. Right carpal of Vouivria damparisensis (MNHN.F.1934.6 DAM 19).

(A) Distal view; (B) medial…

Figure 22. Right carpal of Vouivria damparisensis (MNHN.F.1934.6 DAM 19).
(A) Distal view; (B) medial view; (C) proximal view; (D) anterior view. Scale bar equals 5 cm.
Figure 23

Figure 23. Left metacarpal I of Vouivria…

Figure 23. Left metacarpal I of Vouivria damparisensis (MNHN.F.1934.6 DAM 20).

(A) Lateral view; (B)…

Figure 23. Left metacarpal I of Vouivria damparisensis (MNHN.F.1934.6 DAM 20).
(A) Lateral view; (B) medial view; (C) proximal view (anterior towards top); (D) distal end view (anterior margin towards top). Scale bar equals 5 cm.
Figure 24

Figure 24. Left metacarpal II of Vouivria…

Figure 24. Left metacarpal II of Vouivria damparisensis (MNHN.F.1934.6 DAM 21).

(A) Dorsal view; (B)…

Figure 24. Left metacarpal II of Vouivria damparisensis (MNHN.F.1934.6 DAM 21).
(A) Dorsal view; (B) lateral view; (C) ventral view; (D) medial view; (E) proximal view (anterior margin towards top); (F) distal view (anterior margin towards top). Abbreviations: mc I, articular surface for metacarpal I. Scale bar equals 5 cm.
Figure 25

Figure 25. Left metacarpal III of Vouivria…

Figure 25. Left metacarpal III of Vouivria damparisensis (MNHN.F.1934.6 DAM 22).

(A) Dorsal view; (B)…

Figure 25. Left metacarpal III of Vouivria damparisensis (MNHN.F.1934.6 DAM 22).
(A) Dorsal view; (B) lateral view; (C), ventral view; (D) medial view; (E) proximal view (anterior margin towards top); (F) distal view (anterior margin towards top). Abbreviations: bp, bulge-like process. Scale bar equals 5 cm.
Figure 26

Figure 26. Left metacarpal IV of Vouivria…

Figure 26. Left metacarpal IV of Vouivria damparisensis (MNHN.F.1934.6 DAM 23).

(A) Dorsal view; (B)…

Figure 26. Left metacarpal IV of Vouivria damparisensis (MNHN.F.1934.6 DAM 23).
(A) Dorsal view; (B) lateral view; (C) ventral view; (D) medial view; (E) proximal view (anterior margin towards top); (F) distal view (anterior margin towards top). Abbreviations: fl, medial flange. Scale bar equals 5 cm.
Figure 27

Figure 27. Reconstruction of the metacarpus of

Figure 27. Reconstruction of the metacarpus of Vouivria damparisensis (MNHN.F.1934.6 DAM 20–23) in proximal view.

Figure 27. Reconstruction of the metacarpus of Vouivria damparisensis (MNHN.F.1934.6 DAM 20–23) in proximal view.
Scale bar equals 5 cm.
Figure 28

Figure 28. Manual phalanges of Vouivria damparisensis

Figure 28. Manual phalanges of Vouivria damparisensis (MNHN.F.1934.6 DAM 15-18).

Left manual phalanx I-1 (DAM16)…

Figure 28. Manual phalanges of Vouivria damparisensis (MNHN.F.1934.6 DAM 15-18).
Left manual phalanx I-1 (DAM16) in (A) dorsal view; (B) proximal view; (C) distal view. Left manual phalanx II-1 (DAM 15) in (D) dorsal view; (E) proximal view; (F) distal view. Left manual phalanx III-1 (DAM 17) in (G) dorsal view; (H) proximal view; (I) distal view. Left manual phalanx IV-1 (DAM 18) in (J) dorsal view; (K) proximal view; (L) distal view. Scale bar equals 5 cm.
Figure 29

Figure 29. Left ilium of Vouivria damparisensis

Figure 29. Left ilium of Vouivria damparisensis (MNHN.F.1934.6 DAM 33).

(A) Lateral view; (B) medial…

Figure 29. Left ilium of Vouivria damparisensis (MNHN.F.1934.6 DAM 33).
(A) Lateral view; (B) medial view. Abbreviations: act, acetabulum; pp, pubic peduncle. Scale bar equals 10 cm.
Figure 30

Figure 30. Distal end of the left…

Figure 30. Distal end of the left pubis of Vouivria damparisensis (MNHN.F.1934.6 DAM 31).

(A)…

Figure 30. Distal end of the left pubis of Vouivria damparisensis (MNHN.F.1934.6 DAM 31).
(A) Medial view; (B) distal view. Scale bar equals 10 cm.
Figure 31

Figure 31. Left ischium of Vouivria damparisensis

Figure 31. Left ischium of Vouivria damparisensis (MNHN.F.1934.6 DAM 34).

(A) Medial view; (B) anterior…

Figure 31. Left ischium of Vouivria damparisensis (MNHN.F.1934.6 DAM 34).
(A) Medial view; (B) anterior view; (C) lateral view; (D) proximal view (anterior towards right). Abbreviations: act, acetabulum; fti3, origin site for M. flexor tibialis internus; ip, iliac peduncle; pp, pubic peduncle. Scale bar equals 10 cm.
Figure 32

Figure 32. Right ischium of Vouivria damparisensis

Figure 32. Right ischium of Vouivria damparisensis (MNHN.F.1934.6 DAM 35).

(A) Lateral view; (B) medial…

Figure 32. Right ischium of Vouivria damparisensis (MNHN.F.1934.6 DAM 35).
(A) Lateral view; (B) medial view. Abbreviations: act, acetabulum; fti3, origin site for M. flexor tibialis internus; ip, iliac peduncle; pp, pubic peduncle. Scale bar equals 10 cm.
Figure 33

Figure 33. Right femur of Vouivria damparisensis

Figure 33. Right femur of Vouivria damparisensis (MNHN.F.1934.6 DAM 36).

(A) Anterior view; (B) posterior…

Figure 33. Right femur of Vouivria damparisensis (MNHN.F.1934.6 DAM 36).
(A) Anterior view; (B) posterior view; (C) distal view. Abbreviations: icr, intercondylar ridges. Scale bar equals 10 cm.
Figure 34

Figure 34. Left tibia of Vouivria damparisensis

Figure 34. Left tibia of Vouivria damparisensis (MNHN.F.1934.6 DAM 37).

(A) Anterior view; (B) lateral…

Figure 34. Left tibia of Vouivria damparisensis (MNHN.F.1934.6 DAM 37).
(A) Anterior view; (B) lateral view; (C) posterior view; (D) proximal view (anterior towards top); (E) distal view (anterior towards top). Abbreviations: cc, cnemial crest; lc, lateral condyle; mc, medial condyle; scc, second cnemial crest; tfi, tuberculum fibularis. Scale bar equals 10 cm.
Figure 35

Figure 35. Fibulae of Vouivria damparisensis (MNHN.F.1934.6…

Figure 35. Fibulae of Vouivria damparisensis (MNHN.F.1934.6 DAM 39, 40).

(A) Right fibula in anterior…

Figure 35. Fibulae of Vouivria damparisensis (MNHN.F.1934.6 DAM 39, 40).
(A) Right fibula in anterior view; (B) right fibula in lateral view; (C) right fibula in posterior view; (D) right fibula in medial view; (E) right fibula in proximal view (anterior towards top); (F) right fibula in distal view (anterior towards top); (G) left fibula in anterior view; (H) left fibula in lateral view; (I) left fibula in medial view. Scale bar equals 10 cm.
Figure 36

Figure 36. Left astragalus of Vouivria damparisensis

Figure 36. Left astragalus of Vouivria damparisensis (MNHN.F.1934.6 DAM 11).

(A) Proximal view; (B) distal…

Figure 36. Left astragalus of Vouivria damparisensis (MNHN.F.1934.6 DAM 11).
(A) Proximal view; (B) distal view; (C) lateral view; (D) posterior view; (E) anterior view. Abbreviations: ap, ascending process; pf, posterior fossa; pp, posterior process; ri, ridge. Scale bar equals 5 cm.
Figure 37

Figure 37. Left metatarsal I of Vouivria…

Figure 37. Left metatarsal I of Vouivria damparisensis (MNHN.F.1934.6 DAM 12).

(A) Ventral view; (B)…

Figure 37. Left metatarsal I of Vouivria damparisensis (MNHN.F.1934.6 DAM 12).
(A) Ventral view; (B) lateral view; (C) proximal view; (D) dorsal view; (E) medial view; (F) distal view. Scale bar equals 5 cm.
Figure 38

Figure 38. Left metatarsal III of Vouivria…

Figure 38. Left metatarsal III of Vouivria damparisensis (MNHN.F.1934.6 DAM 14).

(A) Ventral view; (B)…

Figure 38. Left metatarsal III of Vouivria damparisensis (MNHN.F.1934.6 DAM 14).
(A) Ventral view; (B) dorsal view; (C) lateral view; (D) medial view; (E) proximal view; (F) distal view. Scale bar equals 5 cm.
Figure 39

Figure 39. Strict consensus cladogram of 72…

Figure 39. Strict consensus cladogram of 72 MPTs.

Brachiosauridae is highlighted in red, and non-brachiosaurid…

Figure 39. Strict consensus cladogram of 72 MPTs.
Brachiosauridae is highlighted in red, and non-brachiosaurid taxa previously included in the clade are highlighted in blue. Note that this tree was produced following the a priori exclusion of nine unstable taxa (see text for details).
Figure 40

Figure 40. Time-calibrated phylogenetic tree, showing geographic…

Figure 40. Time-calibrated phylogenetic tree, showing geographic distribution of basal Macronaria.

Brachiosauridae is resolved after…

Figure 40. Time-calibrated phylogenetic tree, showing geographic distribution of basal Macronaria.
Brachiosauridae is resolved after the a posteriori deletion of Abydosaurus in TNT. Silhouette of Giraffatitan drawn by Matthew Wedel and available at Phylopic under a Creative Commons Attribution 3.0 Unported license (https://creativecommons.org/licenses/by/3.0/legalcode). Global palaeogeographic reconstructions from Fossilworks (http://fossilworks.org/) showing the distribution of Late Jurassic and Early Cretaceous brachiosaurids.

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