The benthic polyp phase of Medusozoa (Staurozoa, Cubozoa, Scyphozoa, and Hydrozoa) has endoskeletal or exoskeletal support systems, but their composition, development, and evolution is poorly known. In this contribution the variation in synthesis, structure, and function of the medusozoan exoskeleton was examined. In addition, an evolutionary hypothesis for its origin and diversification is proposed for both extinct and extant medusozoans. We also critically reviewed the literature and included data from our own histological and microstructural analyses of some groups. Chitin is a characteristic component of exoskeleton in Medusozoa, functioning as support, protection, and a reserve for various ions and inorganic and organic molecules, which may persuade biomineralization, resulting in rigid biomineralized exoskeletons. Skeletogenesis in Medusozoa dates back to the Ediacaran, when potentially synergetic biotic, abiotic, and physiological processes resulted in development of rigid structures that became the exoskeleton. Of the many types of exoskeletons that evolved, the corneous (chitin-protein) exoskeleton predominates today in polyps of medusozoans, with its greatest variation and complexity in the polyps of Hydroidolina. A new type of bilayered exoskeleton in which there is an exosarc complementing the perisarc construction is here described.
The exoskeleton is an important source of characters for the taxonomy of Hydroidolina. It originates as epidermal secretions and, among other functions, protects the coenosarc of the polypoid stage. However, comparative studies on the exoskeletal tissue origin, development, chemical, and structural characteristics, as well as its evolution and homology, are few and fragmented. This study compares the structure and composition of the exoskeleton and underlying coenosarc in members of “Anthoathecata” and some Leptothecata, but does so mainly in bougainvilliid polyps histological analyses. We also studied the development of the exoskeleton under experimental conditions. We identified three types of glandular epidermal cells related to the origin of the exoskeleton and the secretion of its polysaccharides component. The exoskeleton of the species studied is either bilayered (perisarc and exosarc, especially in bougainvilliids) or corneous (perisarc). The exoskeleton varies in chemical composition, structural rigidity, thickness, extension, and coverage in the different regions of the colony. In bilayered exoskeletons, the exosarc is produced first and appears to be a key step in the formation of the rigid exoskeleton. The exoskeleton contains anchoring structures such as desmocytes and “perisarc extensions.”
Massive accumulations of pelagic species of Sargassum have generated recent social, economic and ecological problems along Caribbean shores. In the Mexican Caribbean, these events have prompted the study of diverse biological and ecological aspects of these macroalgae. However, studies on their associated biota, including Hydrozoa, remain scarce. This research provides important species observations in an area where data is lacking. The occurrence and percent cover of hydroids on Sargassum thalli collected on the beach at Puerto Morelos, Quintana Roo, Mexico from April 2018 to March 2019 was studied. Three pelagic species and morphotypes of Sargassum from this area were analyzed: Sargassum fluitans III, S. natans I and S. natans VIII, as well as a benthic species, S. polyceratium var. ovatum. A total of 14 taxa of hydroids, belonging to the superorders “Anthoathecata” and Leptothecata, were identified. In our study, more hydroid taxa were observed on axes of the different species of Sargassum than on leaves or aerocysts. In general, the greatest species richness of hydroids was observed from February to April. Results show that live hydrozoans attached to pelagic Sargassum are transported into the area. This should be considered in future management measures that address the recurring coastal abundance of Sargassum and its associated biota in the Caribbean region.
We present phylogenetic analyses (parsimony, maximum likelihood and Bayesian inference) for 69 lineages of anthoathecate hydroids based on 18 morphological characters (12 proposed for the first time) plus mitochondrial (16S and COI) and nuclear (18S and 28S) molecular markers. This study aims to test the monophyly of the present concept of the family Bougainvilliidae, assessing its phylogenetic position within Hydroidolina. Our working hypothesis is used as a context for inferring the evolution of certain morphological characters, focusing on the exoskeleton. Our results shed light on some phylogenetic uncertainties within Hydroidolina, delimiting eight well‐supported linages, viz. Hydroidolina, Siphonophorae, Leptothecata, Aplanulata, Filifera II, Filifera III, Capitata and Pseudothecata taxon novum, the latter supported by four morphological synapomorphies. The monophyly of several families was not supported, viz. Bougainvilliidae, Cordylophoridae, Oceaniidae, Rathkeidae and Pandeidae. Some of the genera typically considered in Bougainvilliidae, including Bougainvillia, fell into the clade Pseudothecata, which is consistently reconstructed as the sister group of Leptothecata. We formally suggest that Dicoryne be removed from Bougainvilliidae and placed in the resurrected family Dicorynidae. The exoskeleton was a key feature in the diversification of Hydroidolina, especially with the transition from the bare hydranth to one completely enveloped within the exoskeleton. In this context, bougainvilliids exhibit several intermediate states in the development of the exosarc. Although the concatenated analysis unravels some interesting hypotheses, taxon sampling is still deficient and therefore more data are necessary for achieving a more complete understanding of the evolution and ecology of bougainvilliids and their allies.
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