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Modularity and Complete Natural Homeoses in Cervical Vertebrae of Extant and Extinct Penguins (Aves: Sphenisciformes)

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Abstract

The cervical system of extant penguins (Aves: Sphenisciformes) is organised into morphological modules, each with its biomechanical function. Indeed, for these marine pelagic birds to acquire hydrodynamic morphology, the folding of the neck is essential. Despite a common general structure, the cervical vertebrae exhibit morphological differences depending on their positioning. These characteristics are identified as apparent cases of complete natural homeotic transformations—therefore, the composition of some modules varies. Two types of complete cervical homeoses are identified between species, but the second type can also occur within some species when the post hatching development is considered. The fossil material analysed here makes it apparent that the two modular configurations characterising the anterior part of the neck—a consequence of the first homeosis—existed 36 My and 25 My ago, for one, and circa 10 My ago, for the other. These comparisons also reveal a clear differentiation in vertebral features between the fossil species of the Oligocene–Miocene ages and the more recent and extant penguins. Ultimately, these observations make the proposal of a hypothesis in relation to the ontogenetic influence of Hox genes, and their regulators, based on the changes observed in the cervical segment of Sphenisciformes.

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Notes

  1. Adult A. patagonicus possess tiny ventral apophyses on their transitional vertebra but these cannot be regarded as the strong processus caroticus characteristic of module 5 in all species. This is why the word “true” is used.

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Acknowledgments

Carolina Acosta Hospitaleche and Claudia Tambussi (Museo de La Plata), as well as Eduardo Ruigómez (Museo Egidio Feruglio Paleontólogico), for providing photographs of the cervical vertebrae of Madrynornis mirandus, thus allowing the study of this fossil species. Daniel Ksepka (North Carolina State University) for the transmission of additional photographs of the cervical V of Icadyptes salasi, in order to confirm the first impressions on this important vertebra. Rémi Laffont (Université de Bourgogne) for his help in providing some references, René Guinard for checking the manuscript and the department of comparative anatomy of the Muséum National d’Histoire Naturelle de Paris for access to the osteological collections. Benedikt Hallgrimsson—the editor in chief—and two anonymous reviewers, whose very pertinent comments have helped to improve the manuscript.

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  1. CNRS UMR 5561, Biogéosciences, Université de Bourgogne, 6 Boulevard Gabriel, 21000, Dijon, France

    Geoffrey Guinard & Didier Marchand

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  1. Geoffrey Guinard
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Correspondence to Geoffrey Guinard.

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Guinard, G., Marchand, D. Modularity and Complete Natural Homeoses in Cervical Vertebrae of Extant and Extinct Penguins (Aves: Sphenisciformes). Evol Biol 37, 210–226 (2010). https://doi.org/10.1007/s11692-010-9097-0

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  • DOI: https://doi.org/10.1007/s11692-010-9097-0

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