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. 2015 Feb 12;10(2):e0115477.
doi: 10.1371/journal.pone.0115477. eCollection 2015.

Ichnotaxonomic review of large ornithopod dinosaur tracks: temporal and geographic implications

Affiliations

Ichnotaxonomic review of large ornithopod dinosaur tracks: temporal and geographic implications

Ignacio Díaz-Martínez et al. PLoS One. .

Abstract

Background: Large ornithopod tracks are known from the Upper Jurassic to the uppermost Cretaceous rocks of all continents but Antarctica. They include the tracks historically called Iguanodon footprints, iguanodontid footprints, hadrosaur/hadrosaurid footprints, and other large ornithopod tracks that have been used to define ichnotaxa. More than 40 ichnospecies based on large ornithopod tracks have been defined, but the validity of many of them is questionable.

Methodology/principal findings: 34 ichnogenera and 44 ichnospecies have been analysed in this work. Many of them are considered to be invalid because they have been defined on the basis of poorly preserved tracks without diagnostic features, have an inadequate diagnosis, or are based on temporal and/or geographical criteria. Only eight ichnospecies belonging to the ichnogenera Caririchnium, Iguanodontipus and Hadrosauropodus are here regarded as valid.

Conclusions/significance: The monospecific ichnogenus Iguanodontipus (I. burreyi) is characterized by a small, rounded heel and elongate, narrow digit impressions. Its distribution is limited to the Berriasian-Valanginian of Europe. Caririchnium consists of four ichnospecies (C. magnificum [type ichnospecies], C. kortmeyeri, C. billsarjeanti and C. lotus) with a large, rounded heel and short, wide digit impressions. This ichnogenus ranges from the Berriasian-Hauterivian to the Aptian-Albian of South America, North America, Asia and Europe. Finally, Hadrosauropodus (three ichnospecies: H. langstoni [type ichnospecies], H. leonardii and H. kyoungsookimi) shows a large, bilobed heel and short, wide digit impressions. It is known from the Aptian-Albian to the Maastrichtian of North America, Asia and Europe. The ichnofamily Iguanodontipodidae includes large iguanodontian tracks characterized mainly by mesaxonic, tridactyl and subsymmetrical pes tracks that are as wide as (or wider than) long and have one pad impression in each digit and one in the heel. Its distribution is confidently limited to the Cretaceous of Europe, Asia, North America and South America.

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Conflict of interest statement

Competing Interests: The authors have declared that no competing interests exist.

Figures

Fig 1
Fig 1. Temporal distribution of the large ornithopod ichnotaxa (outline drawings of holotypic tracks).
Tracks are not to scale. 1. Camptosauropus vialovi (redrawn from [74]); 2. Sinoichnites youngi (redrawn from [46]); 3. Kharkushosauropus kharkushensis (redrawn from [56]); 4. Iguanodonichnus frenkii (redrawn from [70]); 5. Camptosaurichnus fasolae (redrawn from [70]); 6. Iguanodontipus burreyi (redrawn from [23]); 7. Wealdenichnites iguanodontoides (redrawn from [46]); 8. Caririchnium magnificum (redrawn from [11]); 9. Gigantoshiraminesauropus matsuoi (redrawn from [82]); 10. Sousaichnium pricei (redrawn from [10]); 11. Staurichnium diogenis (redrawn from [10]); 12. Brachyguanodonipus prejanensis (redrawn from [68]); 13. Hadrosaurichnoides igeensis (redrawn from [92]); 14. Iguanodonipus cuadrupedae (redrawn from [68]); 15. Shiraminesauropus reini (redrawn from [93]); 16. Shiraminesauropus hayashidaniensis (redrawn from [93]); 17. Amblydactylus gethingi (redrawn from [8]); 18. Amblydactylus kortmeyeri (redrawn from [57]); 19. Gypsichnites pacensis (redrawn from [8]); 20. Iguanodonopus xingfuensis (redrawn from [102]); 21. Iguanodontipus billsarjeanti (redrawn from [105]); 22. Ornithopodichnus masanensis (redrawn from [20]); 23. Caririchnium kyoungsookimi (redrawn from [80]); 24. Akmechetosauropus makhkamovi (redrawn from [56]); 25. Babatagosauropus bulini (redrawn from [56]); 26. Yangtzepus yipingensis (redrawn from [7]); 27. Bonaparteichnium tali (redrawn from [66]); 28. Sousaichnium monettae (redrawn from [66]); 29. Caririchnium leonardii (redrawn from [76]); 30. Caririchnium lotus (redrawn from [81]); 31. Limayichnus major (redrawn from [66]); 32. Caririchnium protohadrosaurichnos (redrawn from [78]); 33. Jiayinosauropus johnsoni (redrawn from [107]); 34. Apulosauripus federicianus (redrawn from [62]); 35. Hadrosaurichnus titicaensis (redrawn from [96]); 36. Hadrosaurichnus australis (redrawn from [93]); 37. Hadrosauripeda hauboldi (redrawn from [98]); 38. Hadrosauropodus langstoni (redrawn from [24]); 39. Hadrosauropodus nanxiongensis (redrawn from [99]); 40. Orcauichnites garumniensis (redrawn from [111]); 41. Ornithopodichnites magna (redrawn from [111]); 42. Taponichnus donottoi (redrawn from [119]); 43. Telosichnus saltensis (redrawn from [119]). The ichnogenus Goseongosauripus kimi has not been included because it was not possible to find a drawing of its holotype in the literature.
Fig 2
Fig 2. Distribution of studied large ornithopod ichnogenera.
A, temporal distribution; B, temporal distribution by continents. Based on the data from S1 Table. Discontinuous line, there are no data.?, doubtful data. Aus., Australia.
Fig 3
Fig 3. Holotypes of studied large ornithopod ichnotaxa.
A, Akmechetosauropus makhkamovi (redrawn from [56]); B, Amblydactylus gethingi (redrawn from [8]); C, Amblydactylus kortmeyeri (redrawn from [57]); D, Apulosauripus federicianus (redrawn from [62]); E, Babatagosauropus bulini (redrawn from [56]); F, Bonaparteichnium tali (redrawn from [66]); G, Brachyguanodonipus prejanensis (redrawn from [68]); H, Camptosaurichnus fasolae (redrawn from [70]); I, Camptosauropus vialovi (redrawn from [74]); J, Caririchnium magnificum (redrawn from [11]); K, Caririchnium leonardii (redrawn from [76]); L-M, Caririchnium protohadrosaurichnos (redrawn from [78]); N, Caririchnium lotus (redrawn from [81]); O, Caririchnium kyoungsookimi (redrawn from [80]); P, Gigantoshiraminesauropus matsuoi (redrawn from [82]); Q, Gypsichnites pacensis (redrawn from [8]); R, Hadrosaurichnoides igeensis (redrawn from [92]); S, Hadrosaurichnus australis (redrawn from [93]); T, Hadrosaurichnus titicaensis (redrawn from [96]); U, Hadrosauripeda hauboldi (redrawn from [98]); V, Hadrosauropodus langstoni (redrawn from [24]); W, Hadrosauropodus nanxiongensis (redrawn from [99]); X, Iguanodonichnus frenkii (redrawn from [70]); Y, Iguanodonipus cuadrupedae (redrawn from [68]).
Fig 4
Fig 4. Holotypes of studied large ornithopod ichnotaxa (cont.).
A, Iguanodonopus xingfuensis (redrawn from [102]); B, Iguanodontipus burreyi (redrawn from [23]); C, Iguanodontipus billsarjeanti (redrawn from [105]); D, Jiayinosauropus johnsoni (redrawn from [107]); E, Kharkushosauropus kharkushensis (redrawn from [56]); F, Limayichnus major (redrawn from [66]); G, Orcauichnites garumniensis (redrawn from [111]); H, Ornithopodichnites magna (redrawn from [111]); I, Ornithopodichnus masanensis (redrawn from [20]); J, Shiraminesauropus reini (redrawn from [82]); K, Shiraminesauropus hayashidaniensis (redrawn from [82]); L, Sinoichnites youngi (redrawn from [47]); M, Sousaichnium pricei (redrawn from [10]); N, Sousaichnium monettae (redrawn from [66]); O, Staurichnium diogenis (redrawn from [10]); P, Taponichnus donottoi (redrawn from [119]); Q, Telosichnus saltensis (redrawn from [119]); R, Wealdenichnites iguanodontoides (redrawn from [47]); S, Yangtzepus yipingensis (redrawn from [7]).
Fig 5
Fig 5. Groups of large ornithopod tracks classified on the basis of heel and digit impressions (see explanation in the text).
Group 1: A, holotype of Iguanodontipus burreyi (redrawn from [23]); B, Iguanodontipus burreyi (redrawn from [3]); C-E, Iguanodontipus isp. (redrawn from [101]). Group 2: F, holotype of Caririchnium magnificum (redrawn from [11]); G, holotype of Caririchnium lotus (redrawn from [81]); H, holotype of Iguanodontipus billsarjeanti (redrawn from [105]); I, holotype of Amblydactylus kortmeyeri (redrawn from [57]); J, Caririchnium isp. (redrawn from [95]). Group 3: K, holotype of Caririchnium leonardii (redrawn from [76]); L, holotype of Caririchnium kyoungsookimi (redrawn from [80]); M, Caririchnium leonardii (redrawn from [100]); N, Caririchnium isp. (redrawn from [171]); O, Caririchnium isp. (redrawn from [167]); P, Caririchnium leonardii (redrawn from [43]); Q, Caririchnium leonardii (redrawn from [172]); R, Caririchnium isp. (redrawn from [95]); S, holotype of Hadrosauropodus langstoni (redrawn from [24]); T, Hadrosauropodus langstoni (redrawn from [24]).
Fig 6
Fig 6. Tracks of Iguanodontipus.
A-C, type series of Iguanodontipus burreyi (redrawn from [23]); D, referred track of Iguanodontipus burreyi (redrawn from [3]).
Fig 7
Fig 7. Tracks of Caririchnium.
A-B, type series of Caririchnium magnificum (redrawn from [11]); C-D, type series of Caririchnium kortmeyeri (redrawn from [57]); E-F, type series of Caririchnium billsarjeanti (redrawn from [105]); G-I, type series of Caririchnium lotus (redrawn from [81]).
Fig 8
Fig 8. Tracks of Hadrosauropodus.
A, holotype of Hadrosauropodus langstoni (redrawn from [24]); B, type series of Hadrosauropodus langstoni (redrawn from [24]); C, type series of Hadrosauropodus leonardii (redrawn from [76]); D, referred track of Hadrosauropodus leonardii (redrawn from [43]); E, referred trackway of Hadrosauropodus leonardii (redrawn from [43]); F, type series of Hadrosauropodus kyoungsookimi (redrawn from [80]).
Fig 9
Fig 9. Temporal distribution of ichnogenera included in Iguanodontipodidae.
The ichnogenera in black are considered systematically valid. The ichnogenera in blue are considered systematically non-valid but the tracks belong to Iguanodontipodidae (see explanation in the text). Discontinuous line, there are no data.
Fig 10
Fig 10. Distribution of valid ichnogenera and ichnospecies of Iguanodontipodidae.
A, temporal distribution. Outline drawings of holotypic tracks (see references in Fig. 1); B, temporal distribution by continents. Discontinuous line, there are no data.

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References

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Grants and funding

This paper is a contribution to the projects: IT834-13 of the Gobierno Vasco/Eusko Jaurlaritza; CGL2010-16447 subsidized by the Spanish Ministerio de Econom��a y Competitividad (MINECO), the European Regional Development Fund, and the Government of Aragón (“Grupos Consolidados” and “Dirección General de Patrimonio Cultural”); the project CGL2010-18851/BTE (MINECO); and the project CGL2013-47521-P (MINECO). IDM is the beneficiary of a Postdoctoral grant from the Gobierno de Argentina (CONICET). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

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