Plateosaurus gracilis ( von Huene, 1905 )

Fernández, Omar Rafael Regalado, Stöhr, Henrik, Kästle, Benedikt & Werneburg, Ingmar, 2023, Diversity and taxonomy of the Late Triassic sauropodomorphs (Saurischia, Sauropodomorpha) stored in the Palaeontological Collection of Tübingen, Germany, historically referred to Plateosaurus, European Journal of Taxonomy 913 (1), pp. 1-88 : 64-65

Plateosaurus gracilis ( von Huene, 1905) metataxon

Figs 16 , 18–19 , 21

Sellosaurus gracilis von Huene, 1905: 348 .

Thecodontosaurus hermannianus von Huene, 1905: 348 .

Teratosaurus trossingensis von Huene, 1907: 260 .

Sellosaurus hermannianus ( von Huene, 1905) – von Huene 1915: 3 [partim].

Plateosaurus hermannianus ( von Huene, 1905) – von Huene, 1907: 348.

Type material

Holotype of Plateosaurus gracilis

GERMANY • a partial skeleton; Heslach quarries, Stuttgart, Löwenstein Formation; SMNS 5715 .

Holotype of Thecodontosaurus hermannianus

GERMANY • a maxilla; Heslach quarries, Stuttgart, Löwenstein Formation; SMNS 4388 .

Material examined

“ GPIT Aixheim” ( Fig. 16 )

GERMANY – Aixheim, Löwenstein Formation • fragment of a diapophysis of the atlas; GPIT-PV- 60374 • centrum of the anterior dorsal vertebra; GPIT-PV- 60373 • sacral vertebra; GPIT-PV- 60377 • sacral vertebra; GPIT-PV- 60378 • anterior caudal vertebra; GPIT-PV- 60382 • anterior caudal vertebra; GPITPV-60384 • fragments of chevrons, possibly from the anterior caudals; GPIT-PV- 60391 • anterior caudal vertebra; GPIT-PV- 60383 • caudal vertebra; GPIT-PV- 60385 • centrum of caudal vertebra; GPIT-PV- 60388 • centrum of caudal vertebra; GPIT-PV- 60389 • centrum of caudal vertebra; GPIT-PV- 60387 • centrum of caudal vertebra; GPIT-PV- 60386 • centrum of caudal vertebra; GPIT-PV- 60397 • left metacarpal I; GPIT-PV- 60376 • fragments, possibly of the left fibula; GPIT-PV- 60395 • fragment, possibly of the proximal right tibia; GPIT-PV- 60393 • fragment, possibly of the distal left tibia; GPIT-PV- 60396 • distal tarsal; GPIT-PV- 60372 • astragalus; GPIT-PV- 60370 • distal tarsal; GPIT-PV- 60371 • pedal phalanx III.1; GPIT-PV- 60370 • right metatarsal I; GPIT-PV- 60366 • right metatarsal II; GPIT-PV- 60365 • right metatarsal III; GPIT-PV- 60368.

“ GPIT 18064” ( Fig. 18 )

GERMANY – Rottweil, Stuebsandstein (Löwenstein Formation) • left astragalus; GPIT-PV- 60447 • partial left foot; GPIT-PV- 60448.

“ GPIT 18318a” ( Fig. 19 )

GERMANY – Untere Mühle, Stubensandstein (Löwenstein Formation) • skull, still embedded in the matrix; GPIT-PV- 111900 • cervical 2; GPIT-PV- 60523 • cervical 3; GPIT-PV- 60535 • cervical 4; GPITPV-60537 • cervical 5; GPIT-PV- 60525 • cervical 6; GPIT-PV- 60521 • cervical 7; GPIT-PV- 60531 • cervical 8; GPIT-PV- 60522 • cervical 9; still in matrix; GPIT-PV- 60503 • cervical 10; GPIT-PV- 60536 • dorsal 1; GPIT-PV- 60530 • dorsal 1; GPIT-PV- 60534 • dorsal 3; GPIT-PV- 60257 • partial dorsal 4 and dorsal 5; GPIT-PV- 60532 • dorsal 6; GPIT-PV- 60526 • dorsal 7; GPIT-PV- 60533 • posterior half of the dorsal 8; GPIT-PV- 60528 • dorsals 11, 12 and 13; GPIT-PV- 60520 • possibly a sacral vertebra, but it was not reported as part of the same specimen; GPIT-PV- 60446 • neural arch of a dorsal vertebra; GPIT-PV- 60524 • right scapula, with a cast taken from the negative mould of the original matrix; GPITPV-60512 • left humerus; GPIT-PV- 60514 • right radius, metacarpals I to V, and phalanx I.1; GPITPV-60515 • ungual (possibly pedal ungual); GPIT-PV- 60434 • partial phalanx, the cast was a mould produced from the original negative mould where the specimen was found; GPIT-PV- 60529 • block with ribs; GPIT-PV- 60504 • block with gastralia; GPIT-PV- 60502 • rib fragments; GPIT-PV- 60442, GPIT-PV- 60443, GPIT-PV- 60444, GPIT-PV- 60439, GPIT-PV- 60441, GPIT-PV- 60438, GPIT-PV- 60437, GPIT-PV- 60436, GPIT-PV- 60440.

“ GPIT 18392” ( Fig. 21 )

GERMANY – Untere Mühle, Stubensandstein (Löwenstein Formation) • dorsal vertebra 5; GPITPV-60507 • dorsal vertebra 11; GPIT-PV- 60509 • dorsal vertebrae 12 and 13; GPIT-PV- 60508 • dorsal vertebra 10; GPIT-PV- 60517 • dorsal vertebra; GPIT-PV- 60519 • dorsal vertebra; GPIT-PV- 60518 • sacral vertebra; GPIT-PV- 60510 • dorsal vertebra; GPIT-PV- 60511 • dorsal vertebra; GPIT-PV- 60516 • anterior caudal vertebra; GPIT-PV- 60506 • three chevron from anterior caudal; GPIT-PV- 60493 • chevron from anterior caudal; GPIT-PV- 60561 • pedal phalanx; GPIT-PV- 60488 • pedal ungual and indeterminate remains; GPIT-PV- 60488 • indeterminate remains; GPIT-PV- 60489 • left dorsal rib 7; GPIT-PV- 60539 • right dorsal rib 8; GPIT-PV- 60541 • left cervical 10; GPIT-PV- 60540 • right dorsal rib 4; GPIT-PV- 60552 • right dorsal rib 1; GPIT-PV- 60553 • right dorsal rib 2; GPIT-PV- 60554 • right cervical rib 9; GPITPV-60556 • left dorsal rib 9; GPIT-PV- 60558 • left dorsal rib 12; GPIT-PV- 60555 • left dorsal rib 11; GPIT-PV- 60557 • left dorsal rib 6; GPIT-PV- 60543 • left dorsal rib 10; GPIT-PV- 60544 • left dorsal rib 5; GPIT-PV- 60545 • right dorsal rib 10; GPIT-PV- 60546 • right dorsal rib 9; GPIT-PV- 60547 • right dorsal rib 8; GPIT-PV- 60551 • right dorsal rib 5; GPIT-PV- 60550 • right dorsal rib 7; GPIT-PV- 60548 • right dorsal rib 7; GPIT-PV- 60549 • left fibula; GPIT-PV- 60566 • pedal phalanx; GPIT-PV- 60490 • pedal phalanx; GPIT-PV- 60488 • fragmentary bones; GPIT-PV- 60500 • pedal phalanx; GPIT-PV- 60494 • right pubis; GPIT-PV- 60565 • left femur proximal part; GPIT-PV- 60496 • left femur distal part; GPIT-PV- 60495 • right ischium; GPIT-PV- 60563.

Remarks

The specimen known as “GPIT Aixheim” was tentatively classified as Teratosaurus suevicus by von Huene ( von Huene 1905: 345–349; 1907: 158–168) (species 28 in Fig. 3 ). However, the maxilla NHMUK PV OR 38646, which is the holotype of Teratosaurus suevicus , has been identified as a rauisuchian ( Brusatte et al. 2009). A similar situation occurred with “GPIT 18392”, also referred to as Teratosaurus suevicus ( von Huene 1915) . Subsequently, both “GPIT Aixheim” and “GPIT 18392” were reclassified as Plateosaurus gracilis ( Sellosaurus gracilis in Hungerbühler 1998; Plateosaurus gracilis in Yates 2003) (species 20 in Fig. 3 ). “GPIT 18318a” is also referred to as cf. Plateosaurus gracilis , as it has been included as a part of Pl. gracilis following Yates (2003).

Figures

Fig. 16 (see next page). Material from Aixheim referred to as Teratosaurus ‘suevicus’. The holotype of T. suevicus Mayer, 1861 is a right maxilla currently stored in the Natural History Museum of London (NHMUK PV OR 38646). Dashed lines refer to elements that cannot be unequivocally linked to the anatomical position suggested in von Huene’s (1907) plates. A. GPIT-PV-60394, fragment of a presumed postzygapophyses from a cervical vertebra in left lateral view. B. GPIT-PV-60374, fragment of a diapophysis of the atlas in left lateral view. C. GPIT-PV-60373, centrum of the anterior dorsal vertebra in left lateral view. D. GPIT-PV-60384, centrum of a presumed posterior dorsal vertebra in left lateral view. E. GPIT-PV-60377, sacral vertebra in posterior view. F. GPIT-PV-60378, sacral vertebra in ventral view. G. GPIT-PV-60382, caudosacral vertebra in left lateral view. H. GPIT-PV-60383, anterior caudal vertebra in left lateral view. I. GPIT-PV-60381, deformed anterior caudal vertebra. J. GPIT-PV-60385, caudal vertebra in left lateral view. K. GPIT-PV-60388, centrum of caudal vertebra in right lateral view. L. GPIT-PV-60389, centrum of caudal vertebra in left lateral view. M. GPIT-PV-60387, centrum of caudal vertebra in left lateral view. N. GPIT-PV-60386, centrum of caudal vertebra in left lateral view. O. GPIT-PV-60397, centrum of caudal vertebra in left lateral view. P. GPIT-PV-60376, left metacarpal I in anterior view. Q. GPIT-PV-60395, fragments, possibly of the left fibula in lateral view. R. GPIT- PV-60396, presumed fragment of the distal left tibia in distal view. S. GPIT-PV-60372, distal tarsal in proximal view. T. GPIT-PV-60369, deformed astragalus in anterior view. U. GPIT-PV-60371, distal tarsal in proximal view. V. GPIT-PV-60370, pedal phalanx III.1 in anterior view. W. GPIT-PV-60366, right metatarsal I in anterior view. X. GPIT-PV-60365, right metatarsal II in anterior view. Y. GPIT- PV-60368, right metatarsal III in anterior view. Z. GPIT-PV-60393, presumed fragment of the proximal right tibia in left lateral view. AA. GPIT-PV-60391, fragments of chevrons, possibly from the anterior caudal vertebrae, in lateral view. AB. GPIT-PV-60364, presumed right ungual I.1, stored in the same compartment, unequivocally identified as such by von Huene (1907). The human skeleton represents a height of 1.75 m, as originally drawn by von Huene (1907: pl. 106), the relative size of the reconstructed sauropodomorph is 4.5 m. Reconstruction and photographs made by the authors, with photos taken on the labelled side or on the side that was illustrated in the past.

Fig. 18. Holotype of “Teratosaurus” trossingensis von Huene, 1907 (historical number “GPIT 18064”) from Rottweil. A. GPIT-PV-60447, astragalus, in ventral view. B. Outline of the metatarsals in proximal view. C–Q. GPIT-PV-60448, partial right pes. C. Metatarsal V in anterior view. D. Metatarsal IV in anterior view. E. Pedal phalanx IV.1 in anterior view. F. Pedal phalanx IV.2 in anterior view. G. Pedal phalanx IV.3 in anterior view.H. Pedal phalanx IV.4 in anterior view. I. pedal ungual IV.5.J. Metatarsal III in anterior view. K. Pedal phalanx III.1 in anterior view. L. Pedal phalanx III.2 in anterior view. M. Pedal phalanx III.3 in anterior view (most of the element is reconstructed). N. Metatarsal II (shaft is probably a negative cast). O. Pedal phalanx II.1 (half of the element is probably a negative cast). P. Mmetatarsal I in anterior view (shaft is probably a negative cast). Q. Pedal phalanx I.1 in anterior view.

Fig. 19 (see next page). Material of specimen “GPIT 18318a” on a reconstruction of ‘Sellosaurus gracilis’ as drawn by von Huene (1907: pl. 107); specimen “GPIT 18318a” is referred to as ‘Sellosaurus hermannianus’, and the elements reported as missing are based on what was reported by von Huene (1915). A. GPIT-PV-111900, skull, still embedded in the matrix. B. GPIT-PV-60523, cervical 2 in ventral view. C. GPIT-PV-60535, postzygapophyses of cervical 3 in right lateral view. D. GPIT-PV-60537, cervical 4 in left lateral view. E. GPIT-PV-60525, cervical 5 in right lateral view. F. GPIT-PV-60521, cervical 6 in left lateral view. G. GPIT-PV-60531, cervical 7 in left lateral view. H. GPIT-PV-60522, cervical 8 in right lateral view. I. GPIT-PV-60503, cervical 9 in dorsal view, still in matrix. J. GPIT-PV-60536, cervical 10 in right lateral view. K. GPIT-PV-60530, dorsal 1 in lateral view. L. GPIT-PV-60534, dorsal 1 in dorsal view. M. GPIT-PV-60257, dorsal 3 in ventral view. N. GPIT-PV-60532, partial dorsal 4 and dorsal 5 in ventral view. O. GPIT-PV-60526, dorsal 6 in ventral view. P. GPIT-PV-60533, dorsal 7 in ventral view.Q. GPIT-PV-60528, posterior half of the dorsal 8 in ventral view. R. GPIT-PV-60520, dorsals 11, 12 and 13 in ventral view. S. GPIT-PV-60446, possibly a sacral vertebra in ventral view, but it was not reported as part of the same specimen. T. GPIT-PV-60524, neural arch of a dorsal vertebra in dorsal view. U. GPIT-PV-60512, right scapula, with a cast taken from the negative mould of the original matrix in medial view. V. GPIT-PV-60514, left humerus in anterior view. W. GPIT-PV-60515, right radius in lateral view, metacarpals I to V in anterior view, and phalanx I.1 in anterior view. X. GPIT-PV-60529, partial phalanx, the cast was a mould produced from the original negative mould where the specimen was found. Y. GPIT-PV-60434, ungual (possibly pedal ungual). Z. GPIT-PV-60504, block with ribs. AA. GPIT-PV-60502, block with gastralia. AB. GPIT-PV-60442, rib fragment. AC. GPIT-PV-60443, rib fragment. AD. GPIT-PV-60444, rib fragment. AE. GPIT-PV-60439, rib fragment. AF. GPIT-PV-60441, rib fragment.AG. GPIT-PV-60438, rib fragment. AH. GPIT-PV-60437, rib fragment. AI. GPIT-PV-60436, rib fragment. AJ. GPIT-PV-60440, rib fragments. The human skeleton represents a height of 1.75 m, as originally drawn by von Huene (1907: pl. 107), the relative size of the reconstructed sauropodomorph is 3.5 m. Reconstruction and photographs made by the authors, with photos taken on the labelled side or on the side that was illustrated in the past. 38

Fig. 21 (see next page). Specimen “GPIT 18392”, the number which represents not a catalogue number but a number given to the finding during the excavation in 1912. A. GPIT-PV-60507, dorsal vertebra 5. B. GPIT-PV-60509, dorsal vertebra 11. C. GPIT-PV-60508, dorsal vertebrae 12 and 13. D. GPIT- PV-60517, dorsal vertebra 10. E. GPIT-PV-60519, dorsal vertebra. F. GPIT-PV-60518, dorsal vertebra. G. GPIT-PV-60510, sacral vertebra. H. GPIT-PV-60511, dorsal vertebra. I. GPIT-PV-60516, caudal vertebra. J. GPIT-PV-60506, anterior caudal vertebra. K. GPIT-PV-60493, chevron from anterior caudal. L. GPIT-PV-60561, chevron from anterior caudal. M. GPIT-PV-60493, chevron from anterior caudal. N. GPIT-PV-60493, chevron from anterior caudal. O. GPIT-PV-60488, one phalanx, possibly pedal phalanx. P. GPIT-PV-60488, pedal ungual. Q. GPIT-PV-60489, indeterminate remains. R. ribs: a = GPIT- PV-60539, left dorsal rib 7; b = GPIT-PV-60541, right dorsal rib 8; c = GPIT-PV-60540, left cervical 10; d = GPIT-PV-60552, right dorsal rib 4, e = GPIT-PV-60553, right dorsal rib 1; f = GPIT-PV-60554, right dorsal rib 2; g = GPIT-PV-60556, right cervical rib 9; h = GPIT-PV-60558, left dorsal rib 9; i = GPIT- PV-60555, left dorsal rib 12; j = GPIT-PV-60557, left dorsal rib 11; k = GPIT-PV-60543, left dorsal rib 6; l = GPIT-PV-60544, left dorsal rib 10; m = GPIT-PV-60545, left dorsal rib 5; n = GPIT-PV-60546, right dorsal rib 10; o = GPIT-PV-60547, right dorsal rib 9; p = GPIT-PV-60551, right dorsal rib 8; q = GPIT- PV-60550, right dorsal rib 5; r - GPIT-PV-60548, right dorsal rib 7; s = GPIT-PV-60549, right dorsal rib 7. S. GPIT-PV-60566, left fibula. T. GPIT-PV-60490, pedal phalanx. U. GPIT-PV-60488, pedal phalanx. V. GPIT-PV-60500, fragmentary bones: t = bone fragment identified as ʻgʼ in Hungerbühler (1998: fig. 4), u = bone fragment identified as ʻfʼ in Hungerbühler (1998: fig. 4), v = bone fragment identified as ʻeʼ in Hungerbühler (1998: fig. 4), w = bone fragment identified as ʻnʼ in Hungerbühler (1998: fig. 4). W. GPIT- PV-60494, pedal phalanx. X. GPIT-PV-60565, right pubis. Y. left femur in two parts, GPIT-PV-60496 (proximal) and GPIT-PV-60495 (distal). Z. GPIT-PV-60563, right ischium. AA. carnivorous teeth that do not belong to sauropodomorphs, GPIT-PV-60491, GPIT-PV-60497, GPIT-PV-60498, GPIT-PV-60499.

Fig. 3. Timeline showing changes in the taxonomic placement of the specimens of early-diverging sauropodomorphs considered to be Plateosaurus Meyer, 1837 from 1900 to 2004. This timeline considers only papers discussing taxonomy and papers describing and naming new species. Lines indicate continuity or change in specimen composition of species across revisions. Arrows outside the timeline indicate that the taxa were named before 1900 and/or are still valid after 2004. When more lines depart from one node, part of the specimens included in that species were referred to as different species. Lines converging in one node indicate that the specimens were lumped together into the species in the node. Stars at the end of a line indicate that in the taxonomic revision, the name became either a nomen dubium or a nomen nudum. Lines continuing after this point indicate that the specimens were referred to the new node. This timeline does not include specimens removed from the genus Plateosaurus, namely Efraasia (von Huene, 1907–1908), Ruehleia Galton, 2001, Camelotia Galton, 1985 and Tuebingosaurus Regalado Fernández & Werneburg, 2022. The green lines correspond to species that have been synonymized withPl. trossingensisFraas, 1913, and the blue lines correspond to those that have been synonymized with Pl. gracilis(von Huene, 1905). The orange-red lines refer to species removed from Sauropodomorpha Huene, 1932 altogether. Changes after 2004 are discussed in the text.