Acuticulata is a globally distributed group in the actiniarian superfamily Metridioidea comprised of taxa with ecological, economic, and scientific significance. Prominent members such as Exaiptasia diaphana and Diadumene lineata serve as model organisms for studying coral symbiosis, bleaching phenomena, and ecological invasions. Despite their importance, unresolved phylogenetic relationships and outdated taxonomic frameworks hinder a full understanding of the diversity and evolution of the taxa in this clade. In this study, we employ a targeted sequence-capture approach to construct a robust phylogeny for Acuticulata, addressing long-standing questions about familial monophyly and comparing the results to results from a more conventional five-gene dataset. Specimens from previously underrepresented families and global regions, including the Falkland Islands, were included to elucidate evolutionary interrelationships and improve resolution. Our results support the monophyly of Aliciidae, Boloceroididae, Diadumenidae, Gonactiniidae, and Metridiidae. Our results reiterate the need for taxonomic revision within the family Sagartiidae, as the specimens we included from this family were recovered in four distinct clades. Based on our results, we transfer Paraiptasia from Aiptasiidae to Sagartiidae. These findings emphasize the utility of genome-scale data for resolving phylogenetic ambiguities for morphologically problematic taxa and suggest a framework for future integrative taxonomic and ecological studies within Acuticulata.
AimTo assess global sampling coverage of sea anemones and global species richness across four different spatial resolutions, and analyse these along latitudinal gradients to investigate true bimodality and the extent to which these patterns have been influenced by uneven sampling efforts.LocationThe study encompasses a global scale.Time PeriodOccurrence data included in this study were collected from 1900 to present.TaxonSea anemones (Actiniaria).MethodsUsing 247,542 global occurrence records, we estimated species richness and sampling coverage across four resolutions of grid cells: 800 km, 600 km, 400 km and 200 km. We employed a standardised rarefaction-extrapolation approach to mitigate biases introduced by uneven sampling efforts and to ensure comparability across spatial scales, then compared these species richness estimates across latitudes.ResultsAcross all resolutions, we find a discernible peak in species richness in temperate latitudes, however, the latitudinal peak in diversity shifts dependent on the resolution; our coarsest resolution reveals the most pronounced bimodality, with peaks especially pronounced around 40 degrees N and 40 degrees S, while our finest resolution reveals species richness peaks at 40 degrees-60 degrees N and a subtler increase around 40 degrees S. We find highest observed species richness consistently in temperate regions across resolutions, particularly in southern California, United States and northern Europe. Across all resolutions, we find a discernible peak in species richness in temperate latitudes, however, the latitudinal peak in diversity shifts dependent on the resolution; our coarsest resolution reveals the most pronounced bimodality, with peaks especially pronounced around 40 degrees N and 40 degrees S, while our finest resolution reveals species richness peaks at 40 degrees-60 degrees N and a subtler increase around 40 degrees S.Main ConclusionsSea anemones display an asymmetrically bimodal pattern of global diversity and display the highest species richness at temperate latitudes around 40 degrees N and 40 degrees S. Our study underscores the need for targeted exploration in undersampled environments and understudied marine invertebrates to continue refining biogeographic theories, such as the frequency of bimodal distribution patterns.
Genomic repetitive DNA sequences (Repeatomes, REPs) are widespread in eukaryotes, influencing biological form and function. In Cnidaria, an early-diverging animal lineage, these sequences remain largely uncharacterized. This study investigates sea anemone REPs (Cnidaria: Actiniaria) in a phylogenetic context. We sequenced and assembled de novo the genome of Actinostella flosculifera and analyzed a total of 38 nuclear genomes to create the first ActiniariaREP library (Actiniaria-REPlib). We compared Actiniaria-REPlib with Repbase and RepeatModeler2 libraries, and used dnaPipeTE to annotate REPs from genomic short-read datasets of 36 species for divergence landscapes. Our study assembled and annotated the mitochondrial genomes, including 27 newly assembled ones. We re-annotated 92
Recent studies have elucidated the diversity of genes encoding venom in Sea anemones. However, most of those genes are yet to be explored in an evolutionary context. Insulin is a common peptide across metazoans and has been coopted into a predatory venom in many venomous lineages. In this study, we focus on the diversity of insulin-derived venoms in Sea anemones and on elucidating their evolutionary history. We sourced data for 34 species of Sea anemones and found sequences belonging to two venom families which have Insulin PFAM annotations. Our findings show that both families have undergone duplication events. Members of each of the independently evolving clades have consistent predicted protein structures and distinct dN/dS values. Our work also shows that sequences allied with VP302 are part of a multidomain venom contig and have experienced a secondary gain into the venom system of cuticulate Sea anemones.
Recent genomic analyses have highlighted the prevalence of speciation with gene flow in many taxa and have underscored the importance of accounting for these reticulate evolutionary processes when constructing species trees and generating parameter estimates. This is especially important for deepening our understanding of speciation in the sea where fast moving ocean currents, expanses of deep water, and periodic episodes of sea level rise and fall act as soft and temporary allopatric barriers that facilitate both divergence and secondary contact. Under these conditions, gene flow is not expected to cease completely while contemporary distributions are expected to differ from historical ones. Here we conduct range-wide sampling for Pederson’s cleaner shrimp (Ancylomenes pedersoni), a species complex from the Greater Caribbean that contains three clearly delimited mitochondrial lineages with both allopatric and sympatric distributions. Using mtDNA barcodes and a genomic ddRADseq approach, we combine classic phylogenetic analyses with extensive topology testing and demographic modeling (10 site frequency replicates x 45 evolutionary models x 50 model simulations/replicate = 22,500 simulations) to test species boundaries and reconstruct the evolutionary history of what was expected to be a simple case study. Instead, our results indicate a history of allopatric divergence, secondary contact, introgression, and endemic hybrid speciation driven by the final closure of the Isthmus of Panama and the strengthening of the Gulf Stream Current ~3.5 million years ago. The history of this species complex recovered by model-based methods that allow reticulation differs from that recovered by standard phylogenetic analyses and is unexpected given contemporary distributions. The geologically and biologically meaningful insights gained by our model selection analyses illuminate a novel pathway of species formation that resulted from one of the most biogeographically significant events in Earth’s history.
The mutualism between clownfishes (or anemonefishes) and their giant host sea anemones are among the most immediately recognizable animal interactions on the planet and have attracted a great deal of popular and scientific attention [[1][1]-[5][2]]. However, our evolutionary understanding of this iconic symbiosis comes almost entirely from studies on clownfishes— a charismatic group of 28 described species in the genus Amphiprion [[2][3]]. Adaptation to venomous sea anemones (Anthozoa: Actiniaria) provided clownfishes with novel habitat space, ultimately triggering the adaptive radiation of the group [[2][3]]. Clownfishes diverged from their free-living ancestors 25-30 MYA with their adaptive radiation to sea anemones dating to 13.2 MYA [[2][3], [3][4]]. Far from being mere habitat space, the host sea anemones also receive substantial benefits from hosting clownfishes, making the mutualistic and co-dependent nature of the symbiosis well established [[4][5], [5][2]]. Yet the evolutionary consequences of mutualism with clownfishes have remained a mystery from the host perspective. Here we use bait-capture sequencing to fully resolve the evolutionary relationships among the 10 nominal species of clownfish-hosting sea anemones for the first time ([Figure 1][6]). Using time-calibrated divergence dating analyses we calculate divergence times of less than 25 MYA for each host species, with 9 of 10 host species having divergence times within the last 13 MYA ([Figure 1][6]). The clownfish-hosting sea anemones thus diversified coincidently with clownfishes, potentially facilitating the clownfish adaptive radiation, and providing the first strong evidence for co-evolutionary patterns in this iconic partnership.### Competing Interest StatementThe authors have declared no competing interest. [1]: #ref-1 [2]: #ref-5 [3]: #ref-2 [4]: #ref-3 [5]: #ref-4 [6]: #F1
Metridium senile (Linneaus, 1761) is simultaneously one of the most familiar sea anemones in temperate shallow water and a species embroiled in taxonomic and nomenclatural confusion. Their large body size, characteristic ruff of tentacles, and ability to thrive on artificial and natural substrates make these animals conspicuous and common on rocky ledges, pilings, wharfs, shipwrecks, and bivalve reefs. Their reported range encompasses all temperate habitats in the northern hemisphere plus a handful of locales in the temperate southern hemisphere that have typically been characterized as anthropogenic introductions. Understanding the range of M. senile has been confounded by confusion about its circumscription relative to its congeners and inconsistency in the names used. Here, we summarize the taxonomy of the genus and clarify the status of M. senile. Based on results of recent phylogenetic studies, we consider Metridium to have two valid species: M. farcimen (Brandt, 1835) and M. senile. We recognize M. senile as the valid, appropriate name for all populations of Metridium in the Atlantic and for some of the populations in the Pacific. We consider M. dianthus (Ellis, 1767), M. exile Hand 1955, M. huanghaiense Pei, 1998, and M. sinensis Pei, 1998 to be junior synonyms of M. senile. We clarify the differences between M. senile and M. farcimen, and comment on the status of M. canum Stuckey, 1914.
Sea anemones (Order Actiniaria) are a diverse group of marine invertebrates ubiquitous across marine ecosystems. Despite their wide distribution and success, a knowledge gap persists in our understanding of their diversity within tropical systems, owed to sampling bias of larger and more charismatic species overshadowing cryptic lineages. This study aims to delineate the sea anemone diversity in Mo’orea (French Polynesia) with the use of a dataset from the Mo’orea Biocode’s “BioBlitz” initiative, which prioritized the sampling of more cryptic and understudied taxa. Implementing a target enrichment approach, we integrate 71 newly sequenced samples into an expansive phylogenetic framework and contextualize Mo’orea’s diversity within global distribution patterns of sea anemones. Our analysis corroborates the presence of several previously documented sea anemones in French Polynesia and identifies for the first time the occurrence of members of genera Andvakia and Aiptasiomorpha. This research unveils the diverse sea anemone ecosystem in Mo’orea, spotlighting the area’s ecological significance and emphasizing the need for continued exploration. Our methodology, encompassing a broad BLAST search coupled with phylogenetic analysis, proved to be a practical and effective approach for overcoming the limitations posed by the lack of comprehensive sequence data for sea anemones. We discuss the merits and limitations of current molecular methodologies and stress the importance of further research into lesser-studied marine organisms like sea anemones. Our work sets a precedent for future phylogenetic studies stemming from BioBlitz endeavors.
The genus Antholoba Hertwig, 1882 (Actiniaria, Enthemonae) is characterized by the presence of short and numerous tentacles, a folded oral disc in lobes or cup-shaped with an expanded distal part, transversely wrinkled body wall surface, a very long mesogloeal sphincter, parietobasilar muscles poorly developed, and the absence of acontia. Currently, the genus Antholoba is classified within the family Actinostolidae Carlgren, 1893 (superfamily Actinostoloidea Carlgren, 1932) and comprises two valid species: A. achates (Drayton in Dana, 1846) which have been recorded in Antarctica, the southwestern Atlantic, and the southeastern-western Pacific; and A. perdix (Drayton in Dana, 1846) which is distributed in the northwestern Atlantic, including the Gulf of Mexico. In recent collections along Ubatuba Bay in northern São Paulo, Brazil, we found specimens of a third, unknown species, which exhibits morphological and genetic differences from the only other species recorded from that place, A. achates (Drayton in Dana, 1846). Additionally, we examined five specimens of A. achates collected in Penha, Santa Catarina State, for morphological comparison. Our phylogenetic analyses, using molecular data, affirm the difference between the two species. Furthermore, the resultant phylogenetic trees recover the species of the genus Antholoba as a sister group to the acuticulate clade, within the superfamily Metridioidea, instead of within Actinostoloidea. We describe the material from Ubatuba as a new species, A. fabiani sp. nov., providing information and photographs of its external and internal anatomy, as well as cnidom, along with sequences of mitochondrial (12S, 16S, and COIII) and nuclear (18S and 28S) markers. Additionally, we propose placing the genus Antholoba within Metridioidea, and introduce the new family Antholobidae fam. nov.
Here we describe a new species of sea anemone from the family Aiptasiidae based on specimens collected from the Gulf of Mexico (USA: Florida & Alabama). Accounts of this species have been known since the early 1990s, primarily from an underwater field guide and hobbyist aquarium literature under the name Lightbulb Anemone. We describe it as a new species from the genus Bellactis based on anatomy, histology, and cnidom. Members of this species are small in size, with a smooth, typically contracted column divided into regions based on color and bearing rows of two or three elevated cinclides in the mid column. Their tentacles are distinctive, translucent, distally inflated and can be bulbous in shape, with sub annular rings. This description synthesizes information about Bellactis and contextualizes what is known about its diversity in light of other members of the Aiptasiidae.
The marine aquarium trade in the United States operates primarily in Florida, and though the trade’s effects on many marine taxa are largely unknown, local declines have been observed for some harvested species. To reverse local declines in the abundance of the giant Caribbean sea anemone Condylactis gigantea, state managers prohibited harvest of this popular aquarium organism in late 2012. To assess the recovery of C. gigantea following the moratorium, we monitored relative abundance at 45 fixed locations in South Florida from 2013 to 2016. Seventeen of the sites were locations provided to us by commercial aquarium-trade collectors. The remaining sites were locations at which C. gigantea was presumably not being exploited. We found increased densities at only 4 of those 17 sites, while densities at the remaining sites remained unchanged. We also examined genome-wide single nucleotide polymorphism data to assess genetic diversity and population structure of 82 individuals from seven locations (three collection and four non-collection locations) across South Florida. Observed genetic diversity was comparable in all locations. But the location in the eastern Gulf of Mexico showed a significant FIS value, suggesting inbreeding that might be attributed to a small number of occupants. Condylactis gigantea is generally well mixed across South Florida, though a relatively weak pattern of genetic differentiation was detected. These results suggest limited reproductive success and dispersal that is restricted by hydrological and geographical barriers. We highlight the importance of periodic population and genetic monitoring to assess changes in relative abundance and genetic diversity.
Cnidocytes are the explosive stinging cells unique to cnidarians (corals, jellyfish, etc). Specialized for prey capture and defense, cnidocytes comprise a group of over 30 morphologically and functionally distinct cell types. These unusual cells are iconic examples of biological novelty but the developmental mechanisms driving diversity of the stinging apparatus are poorly characterized, making it challenging to understand the evolutionary history of stinging cells. Using CRISPR/Cas9-mediated genome editing in the sea anemone Nematostella vectensis, we show that a single transcription factor (NvSox2) acts as a binary switch between two alternative stinging cell fates. Knockout of NvSox2 causes a transformation of piercing cells into ensnaring cells, which are common in other species of sea anemone but appear to have been silenced in N. vectensis. These results reveal an unusual case of single-cell atavism and expand our understanding of the diversification of cell type identity.
Members of the sea anemone genus Metridium are abundant in temperate rocky habitats and fouling communities. Their biogeographic history is expected to reflect changes in currents and habitats that have influenced benthic communities, such as the climate-influenced changes that occurred during the Last Glacial Maximum. More recently, however, anthropogenic influences such as shipping transportation and the creation of artificial habitat have altered and affected the composition of modern-day marine communities. Here we use sequence-capture data to examine the genetic structure of Metridium across its shallow-water distribution to (1) evaluate species boundaries within Metridium, (2) elucidate the dispersal history of Metridium between and among oceans, and (3) assess the influence of anthropogenic movement on modern-day populations. We find strong evidence for two species within Metridium: M. farcimen and M. senile. Dispersal from the Pacific to the Atlantic included a subsequent isolation of a small population in or above the Bering Sea, which has presumably moved southward. Within the native range of M. senile, admixture is prevalent even between oceans as a result of anthropogenic activities. The nonnative populations in Chile and the Falkland Islands came from at least two distinct introduction events originating from both coasts of the United States in the North Pacific and North Atlantic Oceans. Hybridization between M. senile and M. farcimen is documented as occurring in anthropogenically influenced habitats. The heavy influence from anthropogenic activities will continue to impact our understanding of marine organisms, particularly within the native range and for those that are easily transported across long distances.
Here we describe a new species of sea anemone from the family Aiptasiidae based on specimens collected from the Gulf of Mexico (USA: Florida & Alabama). Accounts of this species have been known since the early 1990’s, primarily from an underwater field guide and hobbyist aquarium literature under the name “Lightbulb Anemone.” We describe it as a new species from the genus Bellactis based on anatomy, histology, and cnidom. Members of this species are small in size, with a smooth, typically contracted column divided into regions based on color and bearing rows of two or three elevated cinclides in the mid column. Their tentacles are distinctive, translucent, distally inflated and can be bulbous in shape, with sub annular rings. This description synthesizes information about Bellactis and contextualizes what is known about its diversity in light of other members of the Aiptasiidae.
Here we describe a new species of sea anemone from the family Aiptasiidae based on specimens collected from the Gulf of Mexico (USA: Florida & Alabama). Accounts of this species have been known since the early 1990's, primarily from an underwater field guide and hobbyist aquarium literature under the name "Lightbulb Anemone." We describe it as a new species from the genus Bellactis based on anatomy, histology, and cnidom. Members of this species are small in size, with a smooth, typically contracted column divided into regions based on color and bearing rows of two or three elevated cinclides in the mid column. Their tentacles are distinctive, translucent, distally inflated and can be bulbous in shape, with sub annular rings. This description synthesizes information about Bellactis and contextualizes what is known about its diversity in light of other members of the Aiptasiidae.
Haloclavidae Verrill, 1899 is a family of burrowing sea anemones grouped within the superfamily Actinioidea (Rafinesque, 1815). Currently, it includes 30 species in 10 genera. Characters given for this family in descriptions of its taxa have not been consistent, with numerous exceptions to the expectations of the familial diagnosis. Previous phylogenetic analyses have shown that Haloclavidae is potentially a polyphyletic group, but resolution of relationships of the few representatives of Haloclavidae included in analyses has been problematic. Here we address questions of monophyly and affinity of Haloclavidae using three mitochondrial and two nuclear markers. We assess the monophyly of Haloclavidae in the context of all major lineages of Actiniaria Hertwig, 1882, emphasizing diversity of superfamily Actinioidea. We use parsimony-based character optimization to interpret the distribution of key traits in the superfamily. We find that Haloclavidae is not monophyletic and propose two new families, Peachiidae fam. nov. and Harenactidae fam. nov., while also retaining some species in the family Haloclavidae, so that taxonomy better reflects relationships and diversity of the group. In addition, we redescribe a species within the newly created Peachiidae, Peachia chilensis Carlgren, 1931. We use recent larval samples obtained in Antofagasta, Chile, and the histological slides from the original description to redescribe P. chilensis, to provide a complete account of cnidae, external, and internal morphology. Finally, we compare P. chilensis to other burrowing anemones found in Chile and provide an understanding of the genus Peachia that reflects recent phylogenetic perspective on diversity of anemones previously assigned to family Haloclavidae.