Systema Naturae includes representatives of every major lineage of the animal phylum Cnidaria. However, Linnaeus did not classify the members of the phylum as is now done, and the diversity of the group is not well represented. We contrast the Linnaean perspective on cnidarian diversity with the modern, phylogenetic perspective. For each order, we detail diversity at the family level, providing phylogenetic context where possible.
The widespread assumption that COI and other mitochondrial genes will be ineffective DNA barcodes for anthozoan cnidarians has not been well tested for most anthozoans other than scleractinian corals. Here we examine the limitations of mitochondrial gene barcoding in the sub-class Octocorallia, a large, diverse, and ecologically important group of anthozoans. Pairwise genetic distance values (uncorrected p) were compared for three candidate barcoding regions: the Folmer region of COI; a fragment of the octocoral-specific mitochondrial protein-coding gene, msh1; and an extended barcode of msh1 plus COI with a short, adjacent intergenic region (igr1). Intraspecific variation was <0.5%, with most species exhibiting no variation in any of the three gene regions. Interspecific divergence was also low: 18.5% of congeneric morphospecies shared identical COI barcodes, and there was no discernible barcoding gap between intra- and interspecific p values. In a case study to assess regional octocoral biodiversity, COI and msh1 barcodes each identified 70% of morphospecies. In a second case study, a nucleotide character-based analysis correctly identified 70% of species in the temperate genus Alcyonium. Although interspecific genetic distances were 2× greater for msh1 than COI, each marker identified similar numbers of species in the two case studies, and the extended COI + igr1 + msh1 barcode more effectively discriminated sister taxa in Alcyonium. Although far from perfect for species identification, a COI + igr1 + msh1 barcode nonetheless represents a valuable addition to the depauperate set of characters available for octocoral taxonomy.
Cnidaria, the sister group to Bilateria, is a highly diverse group of animals in terms of morphology, lifecycles, ecology, and development. How this diversity originated and evolved is not well understood because phylogenetic relationships among major cnidarian lineages are unclear, and recent studies present contrasting phylogenetic hypotheses. Here, we use transcriptome data from 15 newly-sequenced species in combination with 26 publicly available genomes and transcriptomes to assess phylogenetic relationships among major cnidarian lineages. Phylogenetic analyses using different partition schemes and models of molecular evolution, as well as topology tests for alternative phylogenetic relationships, support the monophyly of Medusozoa, Anthozoa, Octocorallia, Hydrozoa, and a clade consisting of Staurozoa, Cubozoa, and Scyphozoa. Support for the monophyly of Hexacorallia is weak due to the equivocal position of Ceriantharia. Taken together, these results further resolve deep cnidarian relationships, largely support traditional phylogenetic views on relationships, and provide a historical framework for studying the evolutionary processes involved in one of the most ancient animal radiations.
The anthozoan sub-class Octocorallia, comprising approximately 3000 species of soft corals, gorgonians, and sea pens, remains one of the most poorly understood groups of the phylum Cnidaria. Efforts to classify the soft corals and gorgonians at the suprafamilial level have long thwarted taxonomists, and the subordinal groups in current use are widely recognized to represent grades of colony forms rather than clades. Molecular phylogenetic analyses of the sub-class do not support either the current morphologically based subordinal or familial-level taxonomy. To date, however, the resolution necessary to propose an alternative, phylogenetic classification of Octocorallia or to elucidate patterns of morphological evolution within the group is lacking. Attempts to understand boundaries between species and interspecific or intraspecific phylogenetic relationships have been hampered by the very slow rate of mitochondrial gene evolution in Octocorallia, and a consequent dearth of molecular markers with variation sufficient to distinguish species (or sometimes genera). A review of the available ITS2 sequence data for octocorals, however, reveals a yet-unexplored phylogenetic signal both at sequence and secondary-structure levels. In addition, incongruence between mitochondrial and nuclear gene trees suggests that hybrid speciation and reticulate evolution may be an important mechanism of diversification in some genera. Emerging next-generation genomic-sequencing technologies offer the best hope for a breakthrough in our understanding of phylogenetic relationships and of evolution of morphological traits in Octocorallia. Genome and transcriptome sequencing may provide enough characters to resolve relationships at the deepest levels of the octocoral tree, while simultaneously offering an efficient means to screen for new genetic markers variable enough to distinguish species and populations.
Background Our ability to investigate processes shaping the evolutionary diversification of corals (Cnidaria: Anthozoa) is limited by a lack of understanding of species boundaries. Discerning species of corals has been challenging due to a multitude of factors, including homoplasious and plastic morphological characters and the use of molecular markers that are either not informative or have not completely sorted. Hybridization can also blur species boundaries by leading to incongruence between morphology and genetics. We used traditional DNA barcoding and restriction-site associated DNA sequencing combined with coalescence-based and allele-frequency methods to elucidate species boundaries and simultaneously examine the potential role of hybridization in a speciose genus of octocoral, Sinularia. Results Species delimitations using two widely used DNA barcode markers, mtMutS and 28S rDNA, were incongruent with one another and with the morphospecies identifications. When mtMutS and 28S were concatenated, a 0.3% genetic distance threshold delimited the majority of morphospecies. In contrast, 12 of the 15 examined morphospecies formed well-supported monophyletic clades in both concatenated RAxML phylogenies and SNAPP species trees of > 6000 RADSeq loci. DAPC and Structure analyses also supported morphospecies assignments, but indicated the potential for two additional cryptic species. Three morphologically distinct species pairs could not, however, be distinguished genetically. ABBA-BABA tests demonstrated significant admixture between some of those species, suggesting that hybridization may confound species delimitation in Sinularia. Conclusions A genomic approach can help to guide species delimitation while simultaneously elucidating the processes generating coral diversity. Results support the hypothesis that hybridization is an important mechanism in the evolution of Anthozoa, including octocorals, and future research should examine the contribution of this mechanism in generating diversity across the coral tree of life. Electronic supplementary material The online version of this article (10.1186/s12862-019-1427-y) contains supplementary material, which is available to authorized users.
Octocorals, especially gorgonians, are conspicuous on Caribbean coral reefs, but there is no consensus regarding species relationships. Mitochondrial protein-coding genes [NADH-dehydrogenase subunits 2 (ND2) and 6 (ND6), and mutS homolog (msh1), 1633 bp] from 28 shallow-water species were sequenced to develop the first molecular phylogeny for Caribbean octocorals. The specimens were collected primarily in the Caribbean or off Brazil in 1999-2001. Morphological characters (sclerites and axial ultrastructure) were also examined in order to map them onto the molecular phylogeny. Analyses of both nucleotide and amino acid substitutions using maximum parsimony and likelihood (including maximum-likelihood and Bayesian analysis) generated very similar results, with most nodes having high levels of support. These molecular results were significantly different from the generally accepted classification. Neither Plexauridae nor Gorgoniidae were monophyletic. Plexaurella spp., nominal plexaurids, were basal to the gorgoniids, sharing many morphological characters with them. This corroborates previous findings using secondary metabolites and biosynthetic pathways. The sea fans, Gorgonia spp. and Pacifigorgia spp., as well as the pinnate gorgonians, Muriceopsis flavida and Pseudopterogorgia spp., did not have sea fan or pinnate relatives, suggesting there has been convergent evolution of colony form. Caribbean plexaurids appeared more derived and/or recently evolved according to both morphological and molecular data (e.g. Eunicea spp. and Plexaura spp.). Molecular phylogenetics is a promising approach for reconstructing phylogenetic relationships among octocorals as well as to understand their complex morphology. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s00227-003-1018-7. On that page (frame on the left side), a link takes you directly to the supplementary material. IntroductionOctocorals, sessile cnidarians, are found in marine habitats ranging from intertidal to abyssal waters and are distributed from the Arctic to the Antarctic (Bayer 1961). Gorgonians, octocorals containing a sclero-proteinaceous axis, are ecologically important and the dominant macrofauna on many Caribbean reefs. There can be up to 40 octocoral species on a single Caribbean coral reef (Sa´nchez et al. 1997 Sa´nchez 1999), and octocoral communities produce dense and colorful gardens, which provide three-dimensional structures for many reef dwellers and scenic value for underwater activities (Fig. 1). Despite their importance, evolutionary relationships among Caribbean octocorals have received Marine Biology (2003) 142: 975-987
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