Marnes d'Auzas Formation

Marnes d'Auzas Formation
Stratigraphic range: Late Cretaceous (Late Maastrichtian)
~
Sandstones and marls of the Tricouté 4 site in the middle part of the Marnes d’Auzas Formation
TypeGeological formation
UnderliesAurignac Formation
OverliesNankin Limestone Formation, Calcarenites du Jadet Formation, Grès de Labarre Formation
ThicknessAbout 100 m (330 ft) ; up to 285 m (935 ft)?
Lithology
PrimaryMarls
OtherSandstones, limestones
Location
Coordinates43°25′N 1°30′E
RegionOccitania
CountryFrance
Marnes d'Auzas Formation is located in France
Marnes d'Auzas Formation
Marnes d'Auzas Formation (France)
Marnes d'Auzas Formation is located in Occitanie
Marnes d'Auzas Formation
Marnes d'Auzas Formation (Occitanie)
Marnes d'Auzas Formation is located in Haute-Garonne
Marnes d'Auzas Formation
Marnes d'Auzas Formation (Haute-Garonne)

The Marnes d'Auzas Formation (Auzas Marls Formation) is a geological Formation in southwestern France whose strata date back to the Late Maastrichtian, approximately 67.5 to 66 million years ago. It consists primarily of marls with some interbeds of sandstones and limestones. These sediments correspond to varied paleoenvironments where lagoons, coastal marshes and more continental environments (lakes, rivers and alluvial plains) coexist and/or follow one another. The Marnes d’Auzas Formation has yielded numerous fossils of aquatic and terrestrial invertebrates and vertebrates, including dinosaurs.

Location

Location of the Marnes d'Auzas Formation within the Maastrichtian deposits organization of the Petites-Pyrénées (Haute-Garonne department).
Location of the Marnes d'Auzas Formation within the Maastrichtian deposits organization of the Petites-Pyrénées (Haute-Garonne).

The Marnes d’Auzas Formation outcrops in the Aurignac, Saint-Martory and Plagne anticlines in the central part of the Haute-Garonne department (Petites Pyrénées), as well as in the Montfa anticline in Ariège department (Plantaurel massif) in Occitania region. The Marnes d’Auzas Formation conformably overlies the Nankin limestone Formation in the Aurignac anticline, the Calcarenites du Jadet Formation in the Saint-Martory and Plagne anticlines,[nb 1] and the Grès de Labarre Formation in the Montfa anticline. The Marnes d’Auzas Formation is conformably overlain by the Lower Paleocene (Danian) Aurignac Formation.[3][4][5][1]

Lithostratigraphy and paleoenvironment

The Marnes d’Auzas Formation is composed mainly of grey to multi-coloured clayey marls (ochre, green, red, or wine-coloured). The marls contain locally several limestone levels (calcarenites, sandy limestones, marly limestones, chalky limestones) and sandstone beds, sometimes cross-bedded.[6][7] In the type locality of the formation, Bilotte assigns it a thickness of 285 m (935 ft).[6] Other publications indicate a thickness of 250 m (820 ft)[8] or only about 100 m (330 ft).[9]

The Marnes d’Auzas Formation corresponds to sediments whose depositional environment evolved from the paralic domain (coastal lagoons, tidal marsh, tidal muddy channels, mudflats) at the base of the formation, towards a more continental domain (freshwater marshes and lakes, alluvial plain, fluvial channels) in its upper part.[10][11][12][13][9][14] The Marnes d’Auzas Formation was deposited on the west coast of the former Ibero-Armorican island, which included a large part of France and Spain.[15]

The rocks of the Marnes d’Auzas Formation and the underlying Ausseing Group attest the alternation of shallow marine sediments and coastal continental deposits (estuarine and fluvial) corresponding to the fluctuations in sea level and the variations in the coastline during the upper Maastrichtian in this part of the Ibero-Armorican island.[16]

Age

The Marnes d’Auzas Formation is dated to the late Maastrichtian on the basis of its paleontological content, which notably combines marine and continental microfauna and microflora characteristic of this age,[17] but also thanks to the age of the underlying geological formations, and on correlations with the open-ocean deposits of the blue marls of Saint-Loup and yellow marly limestones of Gensac, two western lateral equivalents of the Marnes d’Auzas, which yielded ammonites and foraminifera characteristic of the late Maastrichtian.[18][19][20] Furthermore, at Larcan, the yellow marly limestones of Gensac Formation yielded remains of the hadrosaur Canardia garonnensis (also known in the Marnes d’Auzas Formation) in a stratum located less than one meter below the iridium level marking the Cretaceous–Paleogene boundary (and therefore between the Maastrichtian and the Danian).[21][22]

Paleontology

Stratigraphic distribution of lambeosaurine dinosaurs localities in the Pyrenees, including the Marnes d'Auzas Formation.
Stratigraphic distribution of lambeosaurine dinosaurs localities in the Pyrenees, including the Marnes d'Auzas Formation.

The Marnes d’Auzas Formation contains a wide variety of fossils related to the diversity of marginal-littoral and continental paleoenvironments that meet and/or succeed each other.[23] The formation has yielded numerous foraminifera, ostracods, bivalves, gastropods and charophytes characteristic of different environments.[24][25] Vertebrate remains are also abundant, including numerous dinosaur bones, among which the lambeosaurine hadrosaurids predominate[26][21].These dinosaurs are represented only by juvenile specimens. According to Yves Laurent, this could be explained by a taphonomic bias due to particuliar transport conditions that would have resulted in the deposition of small materials.[27]

Vertebrates of the Marnes d'Auzas Formation

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
 Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Fish

Fish of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Alestidae indet.[28] Indeterminate Tricouté 4 Middle[29] A lower tooth A characiform fish
Chondrichthyes indet.[30] Indeterminate Marsoulas Teeth
Coupatezia[31] C. fallax Peyrecave Top of Lower[32] A tooth A dasyatid stingray. Specimen originally assigned to an undetermined species of Coupatezia closely resembling C. fallax.[33] Subsequently, Cappetta and Corral assigned this tooth to this species.[34]
Lepisosteidae indet. Indeterminate Cassagnau 1 and 2 Middle[35][36] Scales, vertebrae[37] A gar
cf. Lepisosteus cf. L. sp. Cassagnau 1 and 2 Middle Teeth[38] A gar
Palaeogaleus[39] P. sp. Peyrecave Top of Lower Two teeth[40] A triakid shark
Paratrygonorrhina[41] P. amblysoda Peyrecave Top of Lower Teeth A guitarfish. The only tooth figured in 1997 was first attributed to Rhinobatos sp., then to P. amblysoda in 2007.[42][41]
Phyllodontidae Indeterminate Cassagnau 1 Middle Teeth[38]
cf. Pycnodus cf. P. sp. Mérigon 2 Upper Teeth[38]
Rhombodus R. binkhorsti[43] Peyrecave Top of Lower Teeth A stingray of the extinct family Rhombodontidae Life restoration of Rhombodus binkhorsti.
Sparidae indet.[44] Indeterminate Cassagnau 1 and Marsoulas Middle Teeth

Amphibians

Amphibians of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Albanerpetontidae indet. Indeterminate Cassagnau 1 Middle MDE-Cas1-156, a dentary fragment[45]
Anura indet.[46] Indeterminate Peyrecave Top of Lower Fragments of long bone, scapula, and possibly vertebrae and maxillae (cited but not described)
Urodela indet.[47] Indeterminate Peyrecave Top of Lower Unspecified (cited but not described)[39]

Squamates

Squamates of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Amphisbaenia? indet.[48][49] Indeterminate Tricouté 1 Lower[50][51][nb 2] A vertebra
Teiidae indet.[52][49] Indeterminate Peyrecave Top of Lower Tooth
Teiiodea indet.[53] Indeterminate Tricouté 4 Middle A partial dentary A possible barbatteiid?=
Varanoidea indet. Indeterminate Cassagnau 2 Middle MDE-Ca2-124, a dorsal vertebra[54][55] A probable freshwater mosasauroid[56]

Turtles

Turtles of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
?Foxemys ?F. sp. Cassagnau 2 Middle MDE-Cas2-212, right epiplastron + entoplastron[57] A bothremydid turtle
Iberoccitanemys I. atlanticum Cassagnau 2 Middle MDE-Cas2-259, an almost complete shell A bothremydid turtle. This specimen, first assigned to a new species of the genus Elochelys (E. convenarum),[58] was later referred to the new genus Iberoccitanemys with the new combination I. convenarum.[59] This species was subsequently synonymyzed with Polysternon atlanticum (which does not belong to the genus Polysternon) under the new combination I. atlanticum.[60] Shell of Iberoccitanemys atlanticum from Cassagnau 2.
Pleurodira indet. Indeterminate Cassagnau 1 and 2, Mérigon 1 Middle and Upper Dermal plates, girdle and limb elements[61]

Crocodilians

Crocodilians of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Crocodyliformes indet. Indeterminate Auzas, Cassagnau 1 and 2, Mérigon 1, Tricouté 1 and 3, Peyrecave Lower to Upper
Eusuchia indet. Indeterminate Cassagnau 2 Middle MDE-Cas2-229, dorsal (thoracic?) vertebra ; MDE-Cas2-31, 218, dorsal (lumbar?) vertebrae ; MDE-Cas-32, caudal vertebra[63] Vertebrae possibly belonging to Thoracosaurus[64]
Musturzabalsuchus? M sp.? Cassagnau 1 and 2 Middle Teeth[65][66] A probable allodaposuchid eusuchian[nb 3]
Thoracosaurus T. neocesariensis? Cassagnau 1 and 2 Middle
  • MDE-Cas1-199, almost complete skull lacking mandible
  • Teeth
  • MDE-Cas2-262, left humerus
  • MDE-Cas2-33, 202, two cervical vertebrae
  • MDE-Cas1-167 and MDE-Cas2-126, osteoderms[68][69][70]
 Skull initially assigned to Thoracosaurus neocesariensis.[68] According to Christopher Brochu, this specimen belongs to a taxon disctinct from T. neocesariensis.[71] However, this material has not been re-studied since. Skull (MDE-Cas1-199) of Thoracosaurus neocesariensis from Cassagnau 1.

Dinosauria

Dinosauria of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Abelisauridae indet. Indeterminate Cassagnau 1 Middle MDE-Cas1-210, a tooth 4.4 centimetres (2 in) high[72] Initially referred to a Theropoda indet.,[72] this tooth has been reinterpreted as belonging to an indeterminate abelisaurid.[73][74]
Canardia[21] C. garonnensis Tricouté 3 Lower[50][75] A lambeosaurine hadrosaurid. Prieto-Márquez et al. initially assigned Canardia to a clade of basal lambeosaurines, the aralosaurines.[77] Later, Longrich et al. classified Canardia and all European and North African lambeosaurines into a more derived clade, the arenysaurines.[78] Alternatively, Ramírez-Velasco did not recognize either the aralosaurine or the arenysaurine clades and positioned Canardia as the sister taxon to the clade Corythosauria.[79] Holotype right maxilla of Canardia garonnensis from Tricouté 3. Skull reconstruction of Canardia garonnensis showing known elements in white.
Dromaeosauridae indet. Indeterminate Cassagnau 1 and 2 Middle
  • MDE-Cas1-92, tooth)[80][81]
  • MDE-Cas2-28, tooth (the largest tooth 2.6 centimetres (1 in) high and 1.7 centimetres (1 in) wide) [82][83]
  • MDE-Cas2-232[84]
Enantiornithes? indet. Indeterminate Cassagnau 1 Middle MDE-Cas1-152, a proximal end of a scapula[85] An enantiornithean bird
Euronychodon? E. sp. Marsoulas Middle MDE-Mar-03, tooth[86]
Hadrosauridae indet. Indeterminate Auzas MDE-Auz1-01, a proximal humerus ; a distal femur[87]
Indeterminate Mérigon 1 Upper[88] Dentary, tooth[89]
Indeterminate Peyrecave Top of Lower Fragmentary teeth[39]
Indeterminate Tricouté 2 Lower[50]
  • MDE-Ma2-01, fragment of dentary dental battery including four teeth[90]
  • caudal vertebrae
  • tibiae (including MDE-Ma1-02, a 47 centimetres (19 in) long specimen)[91]
  • MDE-Ma1-03, a right metatarsal IV[92]
 The teeth possibly belong to a lambeosaurine[93]
Lambeosaurinae indet. Indeterminate Cassagnau 1 Middle
  • MDE-Cas1-200, proximal region of a left ischium [94][95]
  • MDE-Cas1-14, MDE-Cas1-215 et 216, three ungual phalanges[96]
  • MDE-Cas1-03, maxillary tooth crown[97][98]
  • three dentary tooth crowns (including MDE-Cas1-01 et MDE-Cas1-11)[99][100]
 Fragmentary ischium and three isolated teeth of indeterminate Lambeosaurinae from Cassagnau 1 and Tricouté 1.
Indeterminate Cassagnau 2 Middle
  • MDE-Cas2-138 and MDE-Cas2-248, a predentary and a complete left dentary belonging to a single individual
  • MDE-Cas2-02, a partial left dentary
  • MDE-Cas2-11, a set of dentary tooth crowns
  • MDE-Cas2-223, an isolated dentary tooth crown
  • MDE-Cas2-207, a distal end of left humerus
  • MDE-Cas2-23 and MDE-Cas2-249, tibiae
  • MDE-Cas2-250, a right fibula

An articulated partial postcranial skeleton[nb 5], several ribs (including MDE-Cas2-260), a chevron (MDE-Cas-2-239), and centra of caudal vertebrae were also found.

 Remains first attributed to an indeterminate Euhadrosauria[103] then to an indeterminate lambeosaurine.[104] The pubis of the Cassagnau 2 hadrosaurid exhibits typical lambeosaurine characteristics and in particular shows similarities with the pubis of Parasaurolophus cyrtocristatus, Corythosaurus casuarius, Magnapaulia laticaudus, and, among Ibero-Armorican lambeosaurines, with Arenysaurus.[105] The presence of accessory ridges on at least some dentary teeth from this locality also suggests that they belong to a lambeosaurine.[105] Since the maxilla is not preserved in the Cassagnau 2 lambeosaurine, it is not known whether the material from this site belongs to Canardia (known both in older and younger strata of the same region) or to another lambeosaurine. Skeletal elements (predentary in dorsal and ventral views, two lower jaws, teeth, and a pubis) of an indeterminate lambeosaurine from Cassagnau 2. Partial skeleton of a juvenile lambeosaurine from Cassagnau 2.
Indeterminate Tricouté 1 Lower MDE-Ma1-01, dentary tooth crown ; another tooth is cited and figured by Gheerbrant et al.[106] MDE-Ma1-01 was first attributed to an indeterminate hadrosaurid, then to an indeterminate lambeosaurine.[107]
Titanosauria indet. Indeterminate Cassagnau 1 and 2 Middle A titanosaurian sauropod. In comparison to older titanosaurs from the Ibero-Armorican island, the teeth of this form are different from those of Ampelosaurus and Lirainosaurus,[113][27] and resemble those of Atsinganosaurus.[114]

Pterosaurs

Pterosaurs of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
Azhdarchidaeindet. Indeterminate Mérigon 1 Upper MDE-Me1-04, a nearly complete cervical vertebra with part of both ends missing.[88] A quetzalcoatline azhdarchid belonging to the clade Serpennata, which grouped together quetzalcoatlines possessing highly elongated cervical vertebrae.[115] As preserved, this vertebra measures 46 centimetres (18 in) long, but its total length is estimated at 55 centimetres (22 in).[88] It belongs to a giant pterosaur with an estimated wingspan of 8.5 to 9 metres (28 to 30 ft).[116][117][118]
Azhdarchid cervical vertebra from Mérigon 1.
Azhdarchid cervical vertebra from Mérigon 1.

Mammals

Mammals of the Marnes d'Auzas Formation
Genus Species Location Stratigraphic position Material Notes Images
cf. Azilestes cf. A. yvettae[119] Tricouté 4 Middle A left upper molar[120] A zhelestid eutherian
Hainina H. cassagnauensis[121] Tricouté 1 and 4 Middle One tooth from TRCT 1 and six teeth from TRCT 4[122] A kogaionid multituberculate.
Theria indet. Indeterminate Peyrecave Top of Lower Five fragmentary teeth[123][124]

See also

Notes

  1. ↑ In the Saint-Martory and Plagne anticlines, the layers equivalent to the Nankin Limestone Formation of the Aurignac anticline are represented by the Ausseing Group which contains two carbonate formations, the Nankin 2 Limestone Formation at the base and the Jadet Calcarenites Formation at the top, separated from each other by a marly intercalation, the Lestaillats Marls Formation.[1][2]
  2. ↑ Curiously, Gheerbrant and colleagues place the Tricouté 1 site in the middle part of the Marnes d’Auzas Formation,[29] while other authors place it at the base of the formation.[50][51]
  3. ↑ Musturzabalsuchus is probably a junior synonym of Allodaposuchus.[67]
  4. ↑ MDE-Ma3-21 is the most complete scapula; MDE-Ma3-12 is less complete than MDE-Ma3-21 but preserves a more complete scapular blade showing distal expansion.[76]
  5. ↑ This partial skeleton consists of MDE-Cas2-04, 05, 06, 07, 08, 09 and 10, seven complete cervical vertebrae ; MDE-Cas2-01, a complete left pubis ; MDE-Cas2-12 and 15, left and right nearly complete femora ; MDE-Cas2-13, left tibia ; MDE-Cas2-14 and MDE-Cas-16, left and right fibulae ; MDE-Cas2-17, 18 and 19, left metatarsals II, III and IV ; MDE-Cas2-21 and 22, two phalanges.[101][102]

References

  1. 1 2 Laurent, Bilotte & Le Loeuff 2002, p. 123 (fig.2).
  2. ↑ Laurent 2003, p. 30-31 ; 31 (fig.7).
  3. ↑ Bilotte 1980, p. 58.
  4. ↑ Bilotte, Tambareau & Villatte 1983, p. 271 (tab.1).
  5. ↑ Lepicard 1985, p. 17 (fig.6).
  6. 1 2 Bilotte 1980, p. 59-61 ; 60 (fig.1).
  7. ↑ Lepicard 1985, p. 79 ; 84 ; 87 ; 243.
  8. ↑ Bilotte 1985, p. 264.
  9. 1 2 Laurent, Bilotte & Le Loeuff 2002, p. 123.
  10. ↑ Bilotte 1980, p. 62.
  11. ↑ Lepicard 1985, p. 87-98.
  12. ↑ Bilotte 1985, p. 266.
  13. ↑ Bilotte 1991, p. 418 ; 421.
  14. ↑ Laurent 2003, p. 31.
  15. ↑ Csiki-Sava et al. 2015, p. 9 ; 25 ; 40.
  16. ↑ Laurent 2003, p. 30-31, 31 (fig.7).
  17. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 124 ; 126.
  18. ↑ Bilotte 1991, p. 417 (fig.7) ; 420.
  19. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 124-126.
  20. ↑ Bilotte, Laurent & Téodori 2010, p. 2-8.
  21. 1 2 3 4 Prieto-Márquez et al. 2013, p. 5-13 ; 17-18.
  22. ↑ Bilotte, Laurent & Téodori 2010, p. 6-7.
  23. ↑ Bilotte & Andreu 2006, p. 311-312.
  24. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 123-124.
  25. ↑ Bilotte & Andreu 2006, p. 311-312 ; 313-314.
  26. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 126-131.
  27. 1 2 Laurent 2003, p. 47.
  28. ↑ Gheerbrant, Allain & Laurent 2026, p. 4 ; 6 (fig.3a-c).
  29. 1 2 Gheerbrant, Allain & Laurent 2026, p. 4 (text and fig.2.
  30. ↑ Laurent 2003, p. 35 ; plate 1 (fig.2-3).
  31. ↑ Gheerbrant, Abrial & Cappetta 1997, p. planche 1 (fig.6) ; 262.
  32. ↑ Gheerbrant, Abrial & Cappetta 1997, p. 259 (text and fig.2).
  33. ↑ Gheerbrant, Abrial & Cappetta 1997, p. 261 (plate 1) ; 262.
  34. ↑ Cappetta & Corral 1999, p. 364.
  35. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 124.
  36. ↑ Prieto-Márquez et al. 2013, p. 4 (fig.2) ; 31.
  37. ↑ Laurent 2003, p. 36 ; plate 2 (fig.5).
  38. 1 2 3 Laurent 2003, p. 36.
  39. 1 2 3 Gheerbrant, Abrial & Cappetta 1997, p. 260.
  40. ↑ Gheerbrant, Abrial & Cappetta 1997, p. 260 ; plate 1 (fig.1-2).
  41. 1 2 Kriwet, Soler-Gijón & López-Martínez 2007, p. 1059 ; 1067.
  42. ↑ Gheerbrant, Abrial & Cappetta 1997, p. plate 1 (fig.3) ; 262.
  43. ↑ Gheerbrant, Abrial & Cappetta 1997, p. plate 1 (fig.4-5) ; 262.
  44. ↑ Laurent 2003, p. 37.
  45. ↑ Laurent 2003, p. 37 ; plate 5 (fig. 16).
  46. ↑ Laurent 2003, p. 21 ; 37.
  47. ↑ Laurent 2003, p. 21.
  48. ↑ Gheerbrant, Abrial & Cappetta 1997, p. plate 2 (fig.7).
  49. 1 2 Laurent 2003, p. 41.
  50. 1 2 3 4 Laurent 2003, p. 17 (stratigraphic log of Tricouté 1,2, and 3).
  51. 1 2 Prieto-Márquez et al. 2013, p. 2 ; 4 (fig.2).
  52. ↑ Gheerbrant, Abrial & Cappetta 1997, p. plate 2 (fig.6).
  53. ↑ Gheerbrant, Allain & Laurent 2026, p. 5-6 ; 6 (fig.3d-f.
  54. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 125 (plate 1F) ; 126.
  55. ↑ Laurent 2003, p. 41 ; plate 11 (fig.39).
  56. ↑ Garcia et al. 2015, p. 502.
  57. ↑ Laurent 2003, p. 40 ; plate 10 (fig.33).
  58. ↑ Laurent, Tong & Claude 2002, p. 465-471.
  59. ↑ Pérez-García, Ortega & Murelaga 2012, p. 223-228.
  60. ↑ Pérez-García, Ortega & Murelaga 2021, p. 667-675.
  61. ↑ Laurent 2003, p. 38.
  62. ↑ Laurent 2003, p. 41-42 ; plate 12 (fig.41 ;43) ; plate 13 (fig.44-45 ; 47) ; plate 14 (fig.48-49).
  63. ↑ Laurent 2003, p. 42-43; plate 16 (fig.54, 55) ; plate 17 (fig. 56, 57).
  64. ↑ Laurent 2003, p. 42.
  65. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 126 ; 127.
  66. ↑ Laurent 2003, p. 45 ; plate 21 (fig.64).
  67. ↑ Martin et al. 2016, p. 626-628.
  68. 1 2 Laurent, Buffetaut & Le Loeuff 2000, p. 19-27.
  69. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 126 ; 127 (plate 2A).
  70. ↑ Laurent 2003, p. 42; 43-45 ; plate 13 (fig.46); plate 18 (fig.58); plate 19 (fig.60-61) ; plate 20 (fig.62-63).
  71. ↑ Brochu 2004, p. 621-622.
  72. 1 2 Laurent 2003, p. 46 ; plate 22 (fig.70).
  73. ↑ Csiki-Sava et al. 2015, p. 44-45.
  74. ↑ Isasmendi & Malafaia 2025, p. 472 (fig.9) ; 474.
  75. ↑ Prieto-Márquez et al. 2013, p. 4 (fig.2) ; 5.
  76. ↑ Prieto-Márquez et al. 2013, p. 11 (fig.9A and C) ; 17.
  77. ↑ Prieto-Márquez et al. 2013, p. 5-13 ; 17-18 ; 35 (fig.25).
  78. ↑ Longrich et al. 2021, p. fig.10.
  79. ↑ Ramírez-Velasco 2022, p. 6 (fig.3) ; 7 (fig.4) ; 9.
  80. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 127, plate II (fig.B) ; 129.
  81. ↑ Laurent 2003, p. 46 ; plate 23 (fig. 73).
  82. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 127, plate II (fig.C) ; 129.
  83. ↑ Laurent 2003, p. 46 ; plate 23 (fig. 74).
  84. ↑ Laurent 2003, p. 46 ; plate 22 (fig. 72).
  85. ↑ Laurent 2003, p. 61 ; plate 56 (fig.163).
  86. ↑ Laurent 2003, p. 46-47 ; plate 23 (fig.75).
  87. ↑ Laurent 2003, p. 53 ; plate 38 (fig.117).
  88. 1 2 3 Buffetaut et al. 1997, p. 553.
  89. ↑ Laurent 2003, p. 22 ; 48.
  90. ↑ Laurent 2003, p. 49-50 ; plate 30 (fig.94).
  91. ↑ Laurent 2003, p. 55-56 ; plate 44 (fig.132).
  92. ↑ Laurent 2003, p. 56 ; plate 48 (fig.140).
  93. ↑ Prieto-Márquez et al. 2013, p. 31.
  94. ↑ Laurent 2003, p. 55 ; plate 41 (fig.125).
  95. ↑ Prieto-Márquez et al. 2013, p. 31 ; 32 (fig.23B).
  96. ↑ Laurent 2003, p. 57 ; plate 49 (fig.144-146).
  97. ↑ Laurent 2003, p. 49 ; plate 29 (fig.88).
  98. ↑ Prieto-Márquez et al. 2013, p. 31 ; 32 (fig.23D).
  99. ↑ Laurent 2003, p. 49 ; plate 29 (fig.89) ; plate 30 (fig.92).
  100. ↑ Prieto-Márquez et al. 2013, p. 31 ; 32 (fig.23C).
  101. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 128 (plate III) ; 129 ; 130 (plate IV).
  102. ↑ Laurent 2003, p. 50 ; 55-57 ; plate 30 (fig.94).
  103. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 129-131.
  104. ↑ Prieto-Márquez et al. 2013, p. 31-33.
  105. 1 2 Prieto-Márquez et al. 2013, p. 32-33.
  106. ↑ Gheerbrant, Abrial & Cappetta 1997, p. 264; 266; plate 2 (fig.8).
  107. ↑ Prieto-Márquez et al. 2013, p. 31 ; 32 (fig.23E).
  108. ↑ Laurent 2003, p. 47 ; plate 24 (fig.76).
  109. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 127 (plate II D-F ; 129).
  110. ↑ Laurent 2003, p. 47 ; plate 24 (fig.77).
  111. ↑ Laurent 2003, p. 47 ; plate 24 (fig.78).
  112. ↑ Laurent 2003, p. 47 ; plate 25 (fig. 79).
  113. ↑ Laurent, Bilotte & Le Loeuff 2002, p. 129).
  114. ↑ Garcia et al. 2010, p. 274-275.
  115. ↑ Thomas & McDavid 2025, p. 16 (fig.4) ; 17 ; 24.
  116. ↑ Buffetaut et al. 1997, p. 553-554.
  117. ↑ Laurent 2003, p. 61 ; plate 55 (fig. 162).
  118. ↑ Buffetaut 2008, p. 252.
  119. ↑ Gheerbrant, Allain & Laurent 2026, p. 13-20.
  120. ↑ Gheerbrant, Allain & Laurent 2026, p. 14.
  121. ↑ Gheerbrant, Allain & Laurent 2026, p. 6-13.
  122. ↑ Gheerbrant, Allain & Laurent 2026, p. 6-7.
  123. ↑ Gheerbrant, Abrial & Cappetta 1997, p. 261 (plate 1) ; 263-264.
  124. ↑ Laurent 2003, p. 62.

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