Palaeontology

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Setting the record straight for fossil flying fishes versus non-flying ones: a comment on Xu et al. (2015)

Comment

Andrea Tintori

Cite this article: Tintori A. 2015 Setting the record straight for fossil flying fishes versus non-flying ones: a comment on Xu et al. (2015). Biol. Lett. 11: 20150179. http://dx.doi.org/10.1098/rsbl.2015.0179

Received: 6 March 2015 Accepted: 10 July 2015

Author for correspondence: Andrea Tintori e-mail: [email protected]

The accompanying reply can be viewed at http://dx.doi.org/10.1098/rsbl.2015.0604.

Dipartimento di Scienze della Terra, Universita` degli Studi di Milano, 20133 Milano, Italy

Fossil flying fishes have been known for 150 years from the Upper Triassic of the Alps [1] and recently from the late Middle Triassic of China [2]. So far, five species belonging to Thoracopterus have been described and included in the Peltopleuriformes family Thoracopteridae [2]. Now, Wushaichthys exquisitus [3], possibly from the Xingyi Fauna (Guizhou Province, China) of Late Ladinian age, has been described as belonging in this family. The systematic palaeontology in Xu et al. [3] appears at minimum contradictory, as the supposed synapomorphies of Thoracopteridae are actually plesiomorphies widespread among Peltopleuriformes, most of them not considered in their analysis. The original diagnosis of Thoracopteridae [4] was amended [2,5] after careful revisions of all available specimens, leading to the correction of earlier [5,6] anatomical misinterpretations. A new diagnosis was not proposed [3], and the traditional thoracopterid characteristics were not questioned [3,7], so it is curious that the authors have not realized that Wushaichthys does not possess the actual family synapomorphies. Furthermore, the generic diagnosis is totally uninformative and, because of its combination of characteristics, W. exquisitus could well be assigned to already existing Peltopleuridae genera, for whom extensive recent literature is available [5,8,9]. The phylogenetic analysis presented does not include a significant sampling of the order Peltopleuriformes. Thus, its results are biased by the exclusion of taxa that may be significant to the conclusions drawn. Gradual achievement of the flight capability is also questionable, as all five Thoracopterus species show a very similar ratio in elongation of the paired fins, independently from the scale covering of the body [2,5]. Wushaichthys exhibits sexual dimorphism [3], as do several other Triassic fishes [2,5,8–11]. Modified anal fins do not imply internal fertilization, ovoviviparity and, even less, viviparity, especially if the anal fin is not modified in an actual gonopodium as in extant poeciliid [12], which are ovoviviparous and not viviparous. Wushaichthys, and all other Peltopleuriformes [2,8–10], have no gonopodium-like structure, the anal fin being large, not narrow and elongated, and with hooklets all along the posterior margin, not just at the tip. The provenience and age of the material described is also problematic. Regarding the origin of the specimens, they are said to come from the Wusha area. Therefore, the only exhaustive excavation that conforms to the scientific standards around Wusha is the one near Nimaigu village [2]: as those specimens are not from that excavation, the outcrop and fossiliferous level/bed they are from, and their exact age, should be specified. The proposed ‘Middle Triassic’ age [3] is too vague, when the precise locality is also obscure. Finally, Potanichthys xingyiensi [7] has been included as a valid species [3]. However, it has been already synonymized [13] to Thoracopterus wushaensis [2], following the priority rule of the International Code of Zoological Nomenclature, where two names relate to the same species. However, a question remains to be solved about the site and stratigraphic level yielding Potanichthys. When the paper by Xu et al. [7] was published online, after the authors were aware of Thoracopterus specimens from the Nimaigu excavation [2], their

& 2015 The Author(s) Published by the Royal Society. All rights reserved.

supposed synapomorphies for thoracopterids sensu [3] are actually widespread characters among all Peltopleuriformes, and the remaining two have not been described in both Wushaichthys and Peripeltopleurus spp. Data accessibility. No new data have been used for this comment. Competing interests. The author declares no conflict of interest. Funding. The author received no funding for this study.

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Bronn HG. 1858 Beitra¨ge zur triassischen Fauna und Flora der bitumino¨sen Schiefer von Raibl. Neues Jahrb. Min., Geol. Pala¨ont. 1, 1–32. Tintori A, Sun Z-Y, Lombardo C, Jiang D-Y, Ji C, Ryosuke M. 2012 A new ‘flying’ fish from the Late Ladinian (Middle Triassic) of Wusha (Guizhou Province, southern China). Gortania 33, 39 –50. Xu G-H, Zhao L-J, Shen C-C. 2015 A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early rayfinned fishes. Biol. Lett. 11, 20140960. (doi:10. 1098/rsbl.2014.0960) Griffith J. 1977 The Upper Triassic fishes from Polzberg bei Lunz, Austria. Zool. J. Linn. Soc. 60, 1–93. (doi:10.1111/j.1096-3642.1977. tb00834.x) Tintori A, Sassi D. 1992 Thoracopterus Bronn (Osteichthyes: Actinopterygii): a gliding fish from

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the Upper Triassic of Europe. J. Vert. Paleont. 12, 265 –283. (doi:10.1080/02724634.1992.10011459) Lehman JP. 1979 Note sur les poissons du Trias de Lunz. I. Thoracopterus Bronn. Ann. Naturhist. Mus. Wien 82, 53– 66. Xu G-H, Zhao L-J, Gao K-Q, Wu F-X. 2013 A new stem-neopterygian fish from the Middle Triassic of China shows the earliest over-water gliding strategy of the vertebrates. Proc. R. Soc. B 280, 20122261. (doi:10.1098/rspb.2012.2261) Bu¨rgin T. 1992 Basal Ray-finned fishes (Osteichthyes; Actinopterygii) from the Middle Triassic of Monte San Giorgio (C. Tessin, Switzerland). Schweiz. Pala¨ont. Abh. 114, 1–164. Lombardo C. 1999 Sexual dimorphism in a new species of the Actinopterygian Peltopleurus from the Triassic of Northern Italy. Palaeontology 42, 741 –760. (doi:10.1111/1475-4983.00095)

10. Bu¨rgin T. 1990 Reproduction in Middle Triassic actinopterygians; complex fin structures and evidence of viviparity in fossil fishes. Zool. J. Linn. Soc. 100, 379 –391. (doi:10.1111/j.1096-3642.1990. tb01866.x) 11. Tintori A, Lombardo C. 1996 Gabanellia agilis gen.n.sp.n. (Actinopterygii, Perleidiformes) from the Calcare di Zorzino of Lombardy (N.Italy). Riv. It. Paleont. Strat. 102, 227–236. 12. Rosen DE, Gordon M. 1953 Functional anatomy and evolution of male genitalia in poeciliid fishes. Zoologica 38, 1– 47. 13. Tintori A, Hitij T, Jiang D-Y, Lombardo C, Sun Z-Y. 2014 Triassic actinopterygian fishes: the recovery after the end-Permian crisis. Integr. Zool. 9, 394–411. (doi:10.1111/1749-4877. 12077)

Biol. Lett. 11: 20150179

References

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specimens were described as being from the Ladinian of Xiemi, Wusha, from where no other fossils have previously been published, not even Wushaichthys, which is much more common than Thoracopterus in the same fossiliferous horizon ( personal observation in Nimaigu excavation, 2012). In conclusion, the morphological features of W. exquisitus are very similar to species of Peltopleurus and Peripeltopleurus, closely related genera of Peltopleuridae [5,8,9]: four of the

Setting the record straight for fossil flying fishes versus non-flying ones: a comment on Xu et al. (2015).

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