A comparative analysis of the ontogeny of syngnathids (pipefishes & seahorses) reveals how heterochrony contributed to their diversification

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Abstract
Background: Syngnathids are a highly derived and diverse fish clade comprising the pipefishes, pipe-horses and seahorses. They are characterized by a plethora of iconic traits that increasingly capture the attention of biologists, including geneticists, ecologists and developmental biologists. The current understanding of the origins of their derived body plan is, however, hampered by incomplete and limited descriptions of the early syngnathid ontogeny. Results: We provide a comprehensive description of the development of Nerophis ophidion , Syngnathus typhle , Hippocampus erectus from early cleavage stages to release from the male brooding organ and beyond, including juvenile development. We comparatively describe skeletogenesis with a particular focus on dermal bony plates, the snout-like jaw morphology, and appendages. Conclusions: This most comprehensive and detailed account of syngnathid development to date suggests that convergent phenotypes (e.g. reduction and loss of the caudal fins), likely arose by distinct ontogenetic means in pipefish and seahorses. Comparison of the ontogenetic trajectories of S. typhle and H. erectus provides indications that characteristic features of the seahorse body plan result from developmental truncation. Altogether this work provides a valuable resource and framework for future research to understand the evolution of the outlandish syngnathid morphology from a developmental perspective. Bullet points: Syngnathids (pipefish, pipe-horses and seahorses) are a highly derived and diverse fish taxon and many of their most iconic traits – and these traits’ variation across species - develop during their early ontogeny. This study provides a detailed description and staging table of the entire pre-release development (and beyond) of three distinct syngnathids (two phylogenetically distinct pipefishes and a seahorse) with a focus on the skeletal elements forming the cranium, axial skeleton, appendages and the plate armor. The data provided in this study suggests that developmental heterochrony, i.e. alterations in the rate and timing of developmental processes between lineages, might be a central evolutionary process shaping the morphological diversity found in syngnathids today.
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