1. Evolutionary Biology

An armoured marine reptile from the Early Triassic of South China and its phylogenetic and evolutionary implications

  1. Andrzej S Wolniewicz
  2. Yuefeng Shen
  3. Qiang Li
  4. Yuanyuan Sun
  5. Yu Qiao
  6. Yajie Chen
  7. Yi-Wei Hu
  8. Jun Liu  Is a corresponding author
  1. School of Resources and Environmental Engineering, Hefei University of Technology, China
  2. Institute of Paleobiology, Polish Academy of Sciences, Poland
  3. Section Paleontology, Institute of Geosciences, University of Bonn, Germany
  4. Chengdu Center, China Geological Survey (Southwest China Innovation Center for Geosciences), China
https://doi.org/10.7554/eLife.83163
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  1. Andrzej S Wolniewicz
  2. Yuefeng Shen
  3. Qiang Li
  4. Yuanyuan Sun
  5. Yu Qiao
  6. Yajie Chen
  7. Yi-Wei Hu
  8. Jun Liu
(2023)
An armoured marine reptile from the Early Triassic of South China and its phylogenetic and evolutionary implications
eLife 12:e83163.
https://doi.org/10.7554/eLife.83163
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11 figures, 1 table and 2 additional files

Figures

Figure 1
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Locality and horizon of Prosaurosphargis yingzishanensis (HFUT YZSB-19-109).

(A) The geological map of the Nanzhang-Yuan’an region showing Yingzishan quarry, where HFUT YZSB-19-109 was collected; inset is a paleogeographic map of the South China Block in the Triassic showing the location of the Nanzhang-Yuan’an fauna (after Qiao et al., 2019). (B) Stratigraphic column showing the horizon from which HFUT YZSB-19-109 was collected. Abbreviations: Є-S, Cambrian–Silurian; D-P, Devonian–Permian; F., Formation; K-Q, Cretaceous–Quaternary; M., Middle; T1d, Daye Formation, Lower Triassic; T1j, Jialingjiang Formation, Lower Triassic; T2b, Badong Formation, Middle Triassic; T3-J, Upper Triassic–Jurassic.

Figure 2
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Holotype of Prosaurosphargis yingzishanensis (HFUT YZSB-19-109) (A) and skeletal reconstruction with known elements highlighted in white (gastralia removed for clarity) (B).
Figure 3
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Anterior vertebral column, right pectoral girdle, and right forelimb elements of Prosaurosphargis yingzishanensis.

Abbreviations: 1lg, first lateral gastral element; 2lg, second lateral gastral element; cna, cervical neural arch; cor, coracoid; cr, cervical rib; dc, dorsal centrum; dna, dorsal neural arch; dr, dorsal rib; hum, humerus; mg, median gastral element; os, osteoderm; rad, radius; sc, scapula. Scale bar = 5 cm.

Figure 4
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Dorsal ribs, osteoderms, coracoid, and humerus from the left side of the body of Prosaurosphargis yingzishanensis (A) and detail of lateral osteoderms (B).

Abbreviations: 2lg, second lateral gastral element; cor, coracoid; dr, dorsal rib; eg, ectepicondylar groove; hum, humerus; os, osteoderm, ?sc, ?scapula. Scale bar = 5 cm in (A) and 1 cm in (B).

Figure 5
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Pelvic girdle and hindlimb of Prosaurosphargis yingzishanensis.

Abbreviations: 2lg, second lateral gastral element; ast, astragalus; cal, calcaneum; dt4, distal tarsal IV; fem, femur; fib, fibula; ili, ilium; int, internal trochanter; isc, ischium; mt1, metatarsal I; mt5, metatarsal V; os, osteoderm; p1, first phalanx of digit 1; p5, first phalanx of digit 5; pub, pubis; prz, prezygapophysis; tib, tibia. Scale bar = 3 cm.

Figure 6
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Sacral and caudal vertebrae of Prosaurosphargis yingzishanensis (A), close-up of anterior caudal vertebrae (B), and close-up of more posterior caudal vertebrae (C).

Abbreviations: ch, chevron; cr2, second caudal rib; cr3, third caudal rib; crs6-7, sixth and seventh caudal ribs; dr, dorsal rib; ns, neural spine; os, osteoderm; prz, prezygapophysis; rf, rib facet; rgs, rugose surface; sr, sacral rib. Scale bar = 5 cm in (A) and 1 cm in (B) and (C).

Figure 7
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Coracoids of selected Early and Middle Triassic sauropterygians.

A - Prosaurosphargis (based on specimen HFUT YZSB-19-109); B - Largocephalosaurus (after Li et al., 2014); C - Eusaurosphargis (after Scheyer et al., 2017); D - Atopodentatus (after Cheng et al., 2014); E - Placodus (after Drevermann, 1933); F - Paraplacodus (after Rieppel, 2000b); G (photo) and H (outline) - Hanosaurus (holotype, based on specimen IVPP V 3231); I - Hanosaurus (referred specimen; after Wang et al., 2022); J - Lariosaurus sanxiaensis (after Li and Liu, 2020); K – Majiashanosaurus (after Jiang et al., 2014), L – Corosaurus (after Storrs, 1991); M - Wumengosaurus (after Wu et al., 2011); N – Anarosaurus (after Klein, 2012); O – Nothosaurus (after Ji et al., 2014).

Figure 8
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Phylogenetic relationships of Prosaurosphargis yingzishanensis within Diapsida.

The 50% majority rule consensus of 48 most parsimonious trees (MPTs) obtained from analysis of the updated dataset of Qiao et al., 2022. Numbers below nodes indicate proportion of MPTs in which the node is recovered if it is lower than 1. Abbreviations: Archosauro., Archosauromorpha; E, Eosauropterygia; Ichthyosauro., Ichthyosauromorpha; Lepidosauro., Lepidosauromorpha; Pf, Placodontiformes; Pl, Placodontia; Sa, Sauropterygia; Sr, Saurosphargidae; T., Testudines (total-group); Thala., Thalattosauria. All silhouettes are from Phylopic (http://phylopic.org).

© 2013, N Tamura. Ticinosuchus, Askeptosaurus, Utatsusaurus and Paraplacodus silhouettes were drawn by N. Tamura. Used with permission under license CC BY-NC 3.0 (https://creativecommons.org/licenses/by-nc/3.0/). Reproduction of this figure must abide by the terms of this license.

© 2013, G Monger. Keichousaurus silhouette was drawn by G Monger. Used with permission under license CC BY-NC 3.0 (https://creativecommons.org/licenses/by-nc/3.0/). Reproduction of this figure must abide by the terms of this license.

Figure 9
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Time-scaled phylogenetic tree of Sauropterygomorpha.

Abbreviations: E, Eosauropterygia; Pf, Placodontiformes; Pl, Placodontia; Sa, Sauropterygia; Sam, Sauropterygomorpha; Sr, Saurosphargidae.

Figure 10
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Evolution of the sauropterygian body plan.

Simplified phylogeny of Sauropterygomorpha with key anatomical traits (synapomorphies reconstructed from the phylogenetic analysis) indicated for important nodes.

Figure 11
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Life reconstruction of Prosaurosphargis yingzishanensis depicted in the Early Triassic shallow marine environment of the Nanzhang-Yuan’an region, Hubei Province, South China.

Tables

Table 1
Selected measurements of HFUT YZSB-19-109, holotype of Prosaurosphargis yingzishanensis.
Vertebral column
Centrum positionMaximum anteroposterior length(measured along mid-dorsoventral height)
Last dorsal18.25 mm
1st sacral19.45 mm
2nd sacral20.49 mm
3rd sacral20.50 mm
1st caudal20.50 mm
2nd caudal18.35 mm
3rd caudal19.40 mm
4th caudal20.14 mm
5th caudal18.64 mm
6th caudal19.10 mm
7th caudal17.95 mm
8th caudal18.84 mm
9th caudal18.10 mm
10th caudal17.86 mm
Pectoral girdle
Right scapula
Maximum proximodistal length68.09 mm
Maximum proximal width51.20 mm
Right coracoid
Maximum proximodistal length51.65 mm
Left coracoid
Maximum proximodistal length52.25 mm
Maximum anteroposterior length39.40 mm
Forelimb
Right humerus
Maximum proximodistal length104.44 mm
Maximum proximal width38.85 mm
Maximum distal width27.80 mm
Left humerus
Maximum anteroposterior length104.24 mm
Minimum mediolateral width20.10 mm
Maximum distal width31.50 mm
Right radius
Maximum proximodistal length
(measured along proximodistal axis)		58.50 mm
Maximum proximodistal length
(measured along anterior margin)		60.95 mm
Maximum proximodistal length
(measured along posterior margin)		51.10 mm
Pelvic girdle
Left ilium
Maximum proximodistal length23.20 mm
Maximum distal width32.99 mm
Left ischium
Maximum anteroposterior (proximodistal) length51.44 mm
Maximum mediolateral width36.59 mm
Left pubis
Maximum proximodistal length43.25 mm
Maximum anteroposterior length36.85 mm
Hindlimb
Left femur
Maximum proximodistal length93.85 mm
Maximum proximal width27.65 mm
Maximum distal width23.40 mm
Minimum mediolateral (anteroposterior) width14.25 mm
Left tibia
Maximum proximodistal length60.35 mm
Maximum proximal width18.29 mm
Maximum distal width16.39 mm
Minimum mediolateral (anteroposterior) width10.90 mm
Left fibula
Maximum proximodistal length55.59 mm
Maximum proximal width12.90 mm
Maximum distal width15.24 mm
Minimum mediolateral (anteroposterior) width7.25 mm
Astragalus
Maximum proximodistal length18.45 mm
Maximum mediolateral (anteroposterior) width18.20 mm
Calcaneum
Maximum proximodistal length14.25 mm
Maximum mediolateral (anteroposterior) width13.55 mm
Distal tarsal IV
Maximum proximodistal length9.49 mm
Maximum mediolateral (anteroposterior) width8.95 mm
Metacarpals
Metacarpal I maximum proximodistal length13.44 mm
Metacarpal II maximum proximodistal length20.89 mm
Metacarpal III maximum proximodistal length25.89 mm
Metacarpal IV maximum proximodistal length28.19 mm
Metacarpal V maximum proximodistal length24.25 mm

Additional files

MDAR checklist
https://cdn.elifesciences.org/articles/83163/elife-83163-mdarchecklist1-v2.docx
Source data 1

Character-taxon matrix used in the phylogenetic analysis.

https://cdn.elifesciences.org/articles/83163/elife-83163-data1-v2.txt

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  1. Andrzej S Wolniewicz
  2. Yuefeng Shen
  3. Qiang Li
  4. Yuanyuan Sun
  5. Yu Qiao
  6. Yajie Chen
  7. Yi-Wei Hu
  8. Jun Liu
(2023)
An armoured marine reptile from the Early Triassic of South China and its phylogenetic and evolutionary implications
eLife 12:e83163.
https://doi.org/10.7554/eLife.83163