Ornithocheiridae

Source: Wikipedia, the free encyclopedia.

Ornithocheirids
Temporal range:
Ma
Restored skeleton of
Tropeognathus mesembrinus in the National Museum of Brazil
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: †Pterosauria
Suborder: †Pterodactyloidea
Clade: †
Ornithocheiriformes
Clade: †Ornithocheirae
Family: †Ornithocheiridae
Seeley, 1870
Type species
†
Pterodactylus simus
, 1861
Genera
Synonyms
  • Criorhynchidae
    Hooley
    , 1914

Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 90 million years ago.

Ornithocheirids are generally infamous for having an enormously controversial and very confusing taxonomy. Although agreements that these animals were related, and therefore similar to

azhdarchids such as Quetzalcoatlus. Ornithocheirids were excellent fish hunters, using various flight techniques to catch their prey, and were also capable of flying great distances without flapping constantly. Paleontologists suspect that ornithocheirids were the ancestors of the pteranodontians
; this is due to many shared aspects, such as unique flying techniques, capability of long-distance flights, and most of their diet, which mainly consisted of fish.

History

Naming

junior synonym
O. platyrhinus (E to H)

The family Ornithocheiridae is without a doubt, one of the most well-known pterosaur groups, mostly due to their very controversial and convoluted taxonomic history. Most of the ornithocheirid fossil record consists of isolated teeth, as well as fragmentary bones, reaching hundreds or even thousands of remains in some localities.

ICZN.[4]
In 1874, Owen had proposed two new genera for the
Reginald Walter Hooley in 1914. In his review of Ornithocheirus, he divided the family Ornithocheiridae into two subfamilies: Ornithocheirinae and Criorhynchinae; the former consisted of Ornithocheirus and Lonchodectes, while the latter consisted of Amblydectes and Criorhynchus.[7] In his review, Hooley also considered the species Coloborhynchus clavirostris as a synonym of Criorhynchus simus.[4][8] In 1967, paleontologist Oskar Kuhn placed Criorhynchus within the family Criorhynchidae (which is now considered synonymous to Ornithocheiridae), and recognized Ornithocheirus within the family Ornithocheiridae and subfamily Ornithocheirinae. He also designated the species Coloborhynchus clavirostris as the type species of Coloborhynchus, but agreed with Hooley that it was synonymous with Criorhynchus simus.[4][6] In 1994, however, Yuong-Nam Lee revalidated the genus Coloborhynchus (with C. clavirostris as its type species), and regarded it as distinct from Criorhynchus simus.[6] Later, in 2001, paleontologist David Unwin revised the taxonomic history of the Cambridge Greensand pterosaurs, and divided Ornithocheiridae into three genera: Ornithocheirus, Coloborhynchus and Anhanguera. Unwin also designated Ornithocheirus simus as the type and only species of Ornithocheirus.[4]

In 2003, Unwin defined the family Ornithocheiridae as

eupterodactyloid by Brian Andres and colleagues,[10] was reassigned by Alexander Kellner and colleagues as the sister taxon of the former.[11]

In 2014, Andres and colleagues defined the Ornithocheiridae with a different definition: the most inclusive clade containing Ornithocheirus simus but not

phylogenetic analyses, however, contradict this name synonymy, with Ornithocheiridae and Anhangueridae classified as different families, therefore following the 2014 analysis by Andres and colleagues instead.[12][13][14] Other studies such as the ones by Kellner and colleagues in 2019 have used a different concept, classifying Coloborhynchus, Tropeognathus, as well as several other close relatives such as Ludodactylus and Caulkicephalus within the Anhangueridae, which, along with the family Hamipteridae, forms the larger group Anhangueria. They assigned Ornithocheirus outside the Anhangueria due to being undiagnosable.[4][11][15] Most recent studies have since followed this concept.[16][17][18]

Description

Comparison between the holotypes of Ornithocheirus (A and C) and Tropeognathus (B and D)

Among toothed

morphs reached wingspans of up to 8 meters (26 ft) or more.[13] Specimen NHMUK R481, a specimen that belongs to the species Coloborhynchus capito, the largest toothed pterosaur, had a wingspan that may have reached 7 meters (23 ft).[20] However, in 2013, a specimen referred to the genus Tropeognathus (MN 6594-V), a possible ornithocheirid (as it may be an anhanguerid),[21] was calculated to have had a normal wingspan of 8.26 meters (27.1 ft), with another calculated maximum wingspan reached 8.70 meters (28.5 ft), indicating that the wingspans of toothed pterosaurs could exceed 7 meters (23 ft).[21]

Skull

Skull comparisons between different anhanguerians, notice the ornithocheirids Caulkicephalus (D), Tropeognathus (E and F) and Uktenadactylus (I and J)

Ornithocheirids had elongated jaws with rounded

caniniform, which is similar to those seen in another ornithocheirid (though also recovered as an anhanguerid)[16][18] called Caulkicephalus.[23] The genus Caulkicephalus, though having similarities with other ornithocheirids, including the anterior end of the rostrum being transversely expanded, or having a low, bony sagittal crest that includes a smooth dorsal margin on the rostrum, still possesses some unique features. The most distinct characteristic of Caulkicephalus is that it bores a frontoparietal crest, a feature that is only seen in pteranodontians such as Pteranodon, and in Ludodactylus,[24] a pterosaur once assigned to the Ornithocheiridae, but some recent analysis have placed it within the more inclusive group Anhangueria, outside the Ornithocheiridae.[12][14] Other studies, however, have recovered it within the Anhangueridae instead.[11][16][18]

Postcranial skeleton

Restoration of three Tropeognathus in flight, notice their high aspect ratio

The forelimbs of ornithocheirids were proportionally enormous, around five times longer than their legs. Substantial anchorage on the body is required given the mighty arms, and accordingly, ornithocheirids have robust

pterodactyloids.[25]

In adult ornithocheirids, the

metatarsal.[26]

Classification

Holotype skull and mandible of the ornithocheirids Ferrodraco (A) and Mythunga (B), and holotype mandible of Aussiedraco (C)

The family Ornithocheiridae has had a controversial and very confusing taxonomic history; paleontologists who have studied this group seem to have had a different opinion on the composition of ornithocheirid taxonomy. A term called Anhangueridae was coined by Diogenes de Almeida Campos and Kellner in 1985 to refer to pterosaurs that belong in this family. In 2001, however, Unwin argued that the name Ornithocheiridae refers to an identical group, and should have

nomenclatural priority.[1] He therefore considered Anhangueridae a junior synonym of Ornithocheiridae in his study of pterosaur phylogeny in 2003.[9] However, in many recent studies, Anhangueridae is recovered as a separate and distinct family from Ornithocheiridae, each containing different genera.[27] The original term Ornithocheirae by Seeley had been redefined as the least inclusive clade containing Anhanguera blittersdorffi and Ornithocheirus simus,[10] therefore it is recovered as a larger group comprising the families Anhangueridae and Ornithocheiridae in recent analysis.[10][12][13][14]

Formerly assigned genera

In the past, many pterosaur genera were assigned to the Ornithocheiridae; however, following recent studies, these supposed ornithocheirids had been reclassified to other groups or families. The pterosaur

Lanceodontia in several recent studies such as the ones by Rodrigo Pêgas and colleagues,[17] and Adele Pentland and colleagues, both studies of which are from 2019.[14]

Aetodactylus is another pterosaur that was initially classified within the Ornithocheiridae; later analysis have found it outside the Ornithocheiridae: Timothy Myers in 2015 and Nicholas Longrich and colleagues in 2018 for example, are two studies that found Aetodactylus as sister taxon to the two species of Cimoliopterus (C. cuvieri and C. dunni).[12][27] In 2019, Pêgas and colleagues have found Aetodactylus, along with two other pterosaurs (Camposipterus and Cimoliopterus), within the clade Targaryendraconia, more specifically placed within the family Cimoliopteridae in a polytomy.[17]

The genus Haopterus was used to define the Ornithocheiridae in Unwin's 2003 study; however, Andres and Myers in 2013 argued that Haopterus had not been previously referred to the Ornithocheiridae besides a note added in proof to Unwin in 2001 that stated that Haopterus appeared to be a small ornithocheirid. Phylogenetic analyses since then have found Haopterus as a rogue taxon either within the Pterodactyloidea, the Ornithocheiroidea, the Pteranodontoidea, or the Istiodactylidae. In the phylogenetic analysis by Andres and Myers, Haopterus was recovered as a stable sister taxon to the group Ornithocheiroidea.[28] Some later analyses have also recovered this concept, with both Haopterus and the Ornithocheiroidea placed within the larger group Eupterodactyloidea.[10][12][29] Another study in 2019 recovered Haopterus within a different group called Mimodactylidae.[11]

Phylogeny

Different phylogenetic analysis have found Ornithocheiridae to comprise different genera, the most typical ones being Tropeognathus, Coloborhynchus and Ornithocheirus.[27][28] In 2014, Andres and colleagues created the subfamily Ornithocheirinae to contain Coloborhynchus and Ornithocheirus, as a sister taxon to Tropeognathus, and altogether formed the family Ornithocheiridae.[10] In 2018, Longrich and colleagues had included the genus Siroccopteryx in their phylogenetic analysis, specifically as a member of the ornithocheirine subfamily, sister taxon to Coloborhynchus.[12] In 2019, a study performed by Adele Pentland and colleagues had found the Ornithocheiridae to comprise more genera; while the typical Tropeognathus, Coloborhynchus and Ornithocheirus clade was included in their analysis, Pentland and colleagues found the genera Ferrodraco and Mythunga to belong the Ornithocheiridae as well, specifically sister taxa within the Ornithocheirinae, closely related to Ornithocheirus.[14] In the same year, Megan Jacobs and colleagues have recovered a different set of relationships for ornithocheirids in their analysis. Within the family, three clades emerge: the first one consists of Ornithocheirus, Tropeognathus and Siroccopteryx, the second one comprises Uktenadactylus with several Coloborhynchus species, and the third one comprises Cimoliopterus and Camposipterus. The close relationship between Siroccopteryx, Ornithocheirus and Tropeognathus is supported by several synapomorphies, such the teeth being short, straight, and relatively uniform in size.[13] Several other recent studies such as the ones by Alexander Kellner and colleagues,[11][15] or the one by Rodrigo Pêgas and colleagues, both in 2019, have recovered Coloborhynchus, Siroccopteryx, Tropeognathus and Uktenadactylus within the family Anhangueridae instead of the Ornithocheiridae.[17] In 2020, a study by Borja Holgado and Pêgas had also recovered both Ferrodraco and Mythunga within the Anhangueridae instead of this family.[18]

Cimoliopterus has generally been recovered outside the Ornithocheiridae; subsequent analyses have found it as sister taxon to Aetodactylus, as mentioned earlier.[12][27] Another possible position for Cimoliopterus is within the clade Targaryendraconia, again, closely related to Aetodactylus, and together with Camposipterus, the three formed the family Cimoliopteridae.[17] In the analysis by Jacobs and colleagues, the two Cimoliopterus species had been found as sister taxa to the three Camposipterus species (C. nasutus, C. colorhinus and C. segwickii), altogether formed an unnamed clade within the Ornithocheiridae. However, as noted by Jacobs and colleagues, support for some of these arrangements is relatively weak due to the limited number of characters that can be scored, and the levels of homoplasy are very high.[13]

Below are two cladograms showing the possible genera that are included within the ornithocheirid family. The cladogram to the left is a topology recovered by Jacobs and colleagues,[13] and the one to the right is a topology recovered by Pentland and colleagues.[14]

Paleobiology

Diet and feeding

Restoration of Cimoliopterus stealing prey from a Lonchodectes

Ornithocheirids are generally considered

skimmers, pushing their lower jaw through the water to snap up food upon impact, or fed by gleaning food from the water surface like some modern-day terns and frigatebirds. The skim-feeding hypothesis on ornithocheirids has been discounted in recent assessments of pterosaur skim-feeding, while dip-feeding is supported by a number of anatomical features. The elongated rostra of ornithocheirids are considered ideal for reaching into the water to grab swimming creatures; the rostral crests of ornithocheirids would have worked well as stabilizers for the jaws tips while being plunged into the water. Large, forward-facing eyes and well-developed flocculi are ideal for dip-feeding as well, which permits effective spotting of prey as well as judgement of distances when striking at them; as such, it seems likely that at least several ornithocheirids were efficient dip feeders. Sedate foraging methods might have also been used when hunting -- examples of these methods are: reaching food while being alighted on the water surface, and shallow surface dives.[30]

Locomotion and flight

Similar to modern-day albatrosses, most ornithocheirids used a flight technique called "dynamic soaring", which consists of travelling long distances without flapping using the vertical gradient of wind speed near the ocean surface as an advantage, at moderate flight speed. Several studies showed that most ornithocheirids sprawled their limbs to a large degree, similar to crocodiles, while other studies conclude that ornithocheirids were generally quadrupedal. Yet other studies concluded that ornithocheirids held their limbs more or less vertically extended, similar to an avian or mammalian configuration.[32] Some studies in later genera show that ornithocheirids spend much of their time at sea, and as a result, they perhaps influenced the later pteranodontids with the same piscivorous diet, as well as their well-developed flight techniques.[33][34] Analyses of limb proportions in the genus Anhanguera, however, show that some ornithocheirids were consistent with hopping, but the later genera were suggested that they most likely walked on four limbs, which consists on their wing-fingers as the front limbs, and using their hind limbs to balance.[35]

Paleoecology

Ornithocheirids were a widespread type of pterosaur, with many fossil remains found across the world.

ichthyosaurs Cetarthrosaurus, Platypterygius and Sisteronia were also found alongside the remains of ornithocheirids.[41]

Geological map of the Araripe Basin of Brazil, with the extent of the Santana Group shown in dark blue

A

crocodylomorph Araripesuchus were also found.[56] Several turtle remains were found within the formation, with some specimens referred to the genera Santanachelys, Cearachelys and Araripemys.[57][58] Many fish remains were also found, assigned to the genera Brannerion, Rhinobatos, Rhacolepis, Tharrhias and Tribodus.[59]

Ornithocheirids were also partially distributed in North America, and several specimens are thought to belong to the genus Uktenadactylus (originally Coloborhychus wadleighi).

ammonoids were thought to belong to the genera Turrilites and Scaphites,[64] and along with these, remains of the shark Leptostyrax were also found.[65]

See also

References

  1. ^ a b c d Witton 2013, p. 152.
  2. S2CID 130116778
    .
  3. ^ Seeley, H.G. (1870). "The Ornithosauria: an Elementary Study of the Bones of Pterodactyles". Cambridge: 113.
  4. ^
    PMID 23794925
    .
  5. ^
    doi:10.5194/fr-4-189-2001
    .
  6. ^ a b c d Rodrigues, Taissa; Kellner, Alexander W. A. (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Flugsaurier: Pterosaur Papers in Honour of Peter Wellnhofer. Zitteliana B. 28: 219–228.
  7. NAID 110004312505.[page needed
    ]
  8. doi:10.1080/00222931408693521
    .
  9. ^
    S2CID 86710955
    .
  10. ^
    PMID 24768054
    .
  11. ^
    PMID 31784545
    .
  12. ^
    PMID 29534059
    .
  13. ^
    S2CID 134439172
    .
  14. ^
    PMID 31582757
    .
  15. ^
    PMID 31432888
    .
  16. ^
    PMID 30894614
    .
  17. ^
    S2CID 209595986
    .
  18. ^
    doi:10.4202/app.00751.2020
    .
  19. S2CID 225569843
    .
  20. doi:10.1016/j.cretres.2011.09.003
    .
  21. ^
    PMID 23538956
    .
  22. ^ a b Witton 2013, p. 155.
  23. CiteSeerX 10.1.1.1064.6403
  24. doi:10.1016/j.cretres.2005.03.005
    .
  25. ^ Witton 2013, p. 159.
  26. ^ Witton 2013, pp. 159–160.
  27. ^
    S2CID 86099117
    .
  28. ^
    S2CID 84617119
    .
  29. PMID 26339122
    .
  30. ^ a b Witton 2013, p. 163.
  31. PMID 29877021
    .
  32. ^ Witton 2013, p. 162.
  33. ^ Witton 2013, pp. 161–162.
  34. PMID 21085624
    .
  35. ^ Habib, M. (2011). "Dinosaur Revolution: Anhanguera." H2VP: Paleobiomechanics. Weblog entry, September 20, 2011. Accessed September 28, 2011: http://h2vp.blogspot.com/2011/09/dinosaur-revolution-anhanguera.html
  36. S2CID 67809186
    .
  37. S2CID 129807044
    .
  38. S2CID 129277596
    .
  39. S2CID 129514056
    .
  40. ^ Harrison, C.J.O.; Walker, C.A. (1973). "Wyleyia: a new bird humerus from the Lower Cretaceous of England" (PDF). Palaeontology. 16 (4): 721–728. Archived from the original (PDF) on February 6, 2012.
  41. PMID 24465427
    .
  42. ^ Veldmeijer, A.J. (2006). "Toothed pterosaurs from the Santana Formation (Cretaceous; Aptian-Albian) of northeastern Brazil. A reappraisal on the basis of newly described material Archived 2012-03-17 at the Wayback Machine." Tekst. - Proefschrift Universiteit Utrecht.
  43. S2CID 128410270
    .
  44. ^ Dalla, F.V.; Ligabue, G. (1993). "On the presence of a giant pterosaur in the Lower Cretaceous (Aptian) of Chapada do Araripe (northeastern Brazil)". Bollettino della Società Paleontologica Italiana. 32 (1): 131–136.
  45. S2CID 89178816
    .
  46. PMID 25283914
    .
  47. ^ Veldmeijer, A.J.; Meijer, H.J.M.; Signore, M. (2009). "Description of Pterosaurian (Pterodactyloidea: Anhangueridae, Brasileodactylus) remains from the Lower Cretaceous of Brazil". Deinsea. 13: 9–40.
  48. ^ Leonardi, G. & Borgomanero, G. (1985). "Cearadactylus atrox nov. gen., nov. sp.: novo Pterosauria (Pterodactyloidea) da Chapada do Araripe, Ceara, Brasil." Resumos dos communicaçoes VIII Congresso bras. de Paleontologia e Stratigrafia, 27: 75–80.
  49. ^ P. H. Buisonjé. 1980. Santanadactylus brasilensis nov. gen., nov. sp., a long-necked, large pterosaurier from the Aptian of Brasil. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, B 83:145-172
  50. doi:10.1016/j.cretres.2010.12.008
    .
  51. doi:10.4202/app.2010.0057
    .
  52. S2CID 90477315
    .
  53. S2CID 140174098
    .
  54. doi:10.1127/njgpa/199/1996/151
    .
  55. S2CID 131339386
    .
  56. S2CID 85946263
    .
  57. ^ Zug, George R. "Turtle: Origin and evolution". Encyclopædia Britannica. Retrieved September 18, 2015.
  58. S2CID 45417065
    .
  59. S2CID 86285896
    .
  60. ^ Lee, Y.-N. (1994). "The Early Cretaceous Pterodactyloid Pterosaur Coloborhynchus wadleighi from North America". Palaeontology. 37 (4): 755–763.
  61. PMID 27007950
    .
  62. doi:10.1080/02724634.1995.10011231
    .
  63. ISBN 0-520-24209-2
    .
  64. ^ William A.C. (1969). The Late Cretaceous Ammonites Scaphites leei Reeside and Scaphites hippocrepis (DeKay) in the Western Interior of the United States (PDF). United States Government Printing Office. pp. 3–7.
  65. ^ Cappetta H. 1987. Chondrichthyes II. Mesozoic and Cenozoic Elasmobranchii. Schultze H.-P. (ed.), Handbook of Paleoichthyology, Volume 3B. Gustav Fischer Verlag, Stuttgart, 193 p.

Further reading

External links
Retrieved from "https://en.wikipedia.org/w/index.php?title=Ornithocheiridae&oldid=1217158810"