The importance of molecular characters when morphological variability hinders diagnosability: systematics of the moon jellyfish genus Aurelia (Cnidaria: Scyphozoa).

Cryptic species have been detected across Metazoa, and while no apparent morphological features distinguish them, it should not impede taxonomists from formal descriptions. We accepted this challenge for the jellyfish genus Aurelia, which has a long and confusing taxonomic history. We demonstrate that morphological variability in Aurelia medusae overlaps across very distant geographic localities. Even though some morphological features seem responsible for most of the variation, regional geographic patterns of dissimilarities are lacking. This is further emphasized by morphological differences found when comparing lab-cultured Aurelia coerulea medusae with the diagnostic features in its recent redescription. Previous studies have also highlighted the difficulties in distinguishing Aurelia polyps and ephyrae, and their morphological plasticity. Therefore, mostly based on genetic data, we recognize 28 species of Aurelia, of which seven were already described, 10 are formally described herein, four are resurrected and seven remain undescribed. We present diagnostic genetic characters for all species and designate type materials for newly described and some resurrected species. Recognizing moon jellyfish diversity with formal names is vital for conservation efforts and other studies. This work clarifies the practical implications of molecular genetic data as diagnostic characters, and sheds light on the patterns and processes that generate crypsis.

Is it possible to use behavior characters for evolutionary reconstruction in marine invertebrates? A methodological approach using Ethokit Logger.

The use of behavioral data is quite common in studies of chordate animals and some groups of arthropods; however, these data are usually used in ecological and conservation studies. Their use remains uncommon in phylogenetic reconstructions, especially for non-model groups in behavioral studies. This study aims to evaluate the methodological use of behavioral (feeding process) data with EthoKit Logger in the phylogenetic reconstruction of the Cnidaria, a group in the so-called 'lower' Metazoa. The results indicate considerable cohesion with reconstructions based on molecular data available in previous studies. We therefore suggest that the use of behavioral characters can possible be a useful secondary tool or a proof test for molecular evolutionary reconstructions.

Transcriptomic Analysis of Four Cerianthid (Cnidaria, Ceriantharia) Venoms.

Tube anemones, or cerianthids, are a phylogenetically informative group of cnidarians with complex life histories, including a pelagic larval stage and tube-dwelling adult stage, both known to utilize venom in stinging-cell rich tentacles. Cnidarians are an entirely venomous group that utilize their proteinaceous-dominated toxins to capture prey and defend against predators, in addition to several other ecological functions, including intraspecific interactions. At present there are no studies describing the venom for any species within cerianthids. Given their unique development, ecology, and distinct phylogenetic-placement within Cnidaria, our objective is to evaluate the venom-like gene diversity of four species of cerianthids from newly collected transcriptomic data. We identified 525 venom-like genes between all four species. The venom-gene profile for each species was dominated by enzymatic protein and peptide families, which is consistent with previous findings in other cnidarian venoms. However, we found few toxins that are typical of sea anemones and corals, and furthermore, three of the four species express toxin-like genes closely related to potent pore-forming toxins in box jellyfish. Our study is the first to provide a survey of the putative venom composition of cerianthids and contributes to our general understanding of the diversity of cnidarian toxins.

Ceriantharia (Cnidaria) of the World: an annotated catalogue and key to species.

The diversity of Ceriantharia is known from studies formally describing species from the late 18th Century onwards. However, no nomenclators including a list and discussion of all valid species have been produced since a list discussed by Carlgren in 1912. The present nomenclator presents a complete list of adult species of Ceriantharia of the World, including a discussion on each species. It includes the three families (Arachnactidae, Botrucnidiferidae, Cerianthidae) and the currently accepted 54 species based on their adult form. This study serves as a presentation of the "state-of-the-art" list of species of Ceriantharia, and includes a species identification key to support taxonomic identification. Additional in-depth species-by-species investigations for almost all cerianthid species is still needed, as the information available for most of these species is quite superficial.

Knock knock, who's there?: marine invertebrates in tubes of Ceriantharia (Cnidaria: Anthozoa).

This study reports on the fauna found in/on tubes of 10 species of Ceriantharia and discusses the characteristics of these occurrences, as well as the use of mollusc shells in ceriantharian tube construction. A total of 22 tubes of Ceriantharia from Argentina, Brazil, Japan, Norway, Portugal and the United States were analysed, revealing 58 species of marine invertebrates using them as alternative substrates. Based on a literature review and analyses of the sampled material, we report new occurrences for Photis sarae (Crustacea), Microgaza rotella (Mollusca), Brada sp., Dipolydora spp., Notocirrus spp., and Syllis garciai (Annelida). The use of mollusc shells in tube construction increases the tubes' structural resistance and strength. Ceriantharian tubes are suitable alternative substrates for the dwelling of numerous tubicolous and infaunal species that usually burrow into sediments or anchor on fixed or mobile habitats seeking shelter, thus playing a relevant role as local biodiversity hotspots.

Revision of the genus Ceriantheomorphe (Cnidaria, Anthozoa, Ceriantharia) with description of a new species from the Gulf of Mexico and northwestern Atlantic.

The present study presents a revision of the genus Ceriantheomorphe Carlgren, 1931, including redescriptions of the two presently recognized species, Ceriantheomorphe ambonensis (Kwietniewski, 1898) and Ceriantheomorphe brasiliensis (Mello-Leitão, 1919), comb. nov., and a description of the new species Ceriantheomorphe adelita sp. nov.

Exotic species dominate marinas between the two most populated regions in the southwestern Atlantic Ocean.

Human occupation of coastal areas promotes the establishment of non-native species but information on bioinvasions is usually biased toward the Northern Hemisphere. We assessed non-native species' importance in sessile communities at six marinas along the most urbanized area of the Southwestern Atlantic coastline. We found 67 species, of which 19 are exotic. The most frequent species was the exotic polychaete Branchiomma luctuosum, while the most abundant was the exotic bryozoan Schizoporella errata that monopolized the substrata in three marinas. Along with S. errata, the exotic polychaete Hydroides elegans and ascidian Styela plicata dominated space in the three remaining marinas, while native species were in general rare. We show that communities associated with artificial substrata along this Brazilian urbanized area are dominated by exotic species and that using abundance data along with species identity can improve our understanding of the importance of exotic species for the dynamics of biological communities.

Linear Mitochondrial Genome in Anthozoa (Cnidaria): A Case Study in Ceriantharia.

Sequences and structural attributes of mitochondrial genomes have played a critical role in the clarification of relationships among Cnidaria, a key phylum of early-diverging animals. Among the major lineages of Cnidaria, Ceriantharia ("tube anemones") remains one of the most enigmatic in terms of its phylogenetic position. We sequenced the mitochondrial genomes of two ceriantharians to see whether the complete organellar genome would provide more support for the phylogenetic placement of Ceriantharia. For both Isarachnanthus nocturnus and Pachycerianthus magnus, the mitochondrial gene sequences could not be assembled into a single circular genome. Instead, our analyses suggest that both species have mitochondrial genomes consisting of multiple linear fragments. Linear mitogenomes are characteristic of members of Medusozoa, one of the major lineages of Cnidaria, but are unreported for Anthozoa, which includes the Ceriantharia. The inferred number of fragments and variation in gene order between species is much greater within Ceriantharia than among the lineages of Medusozoa. We identify origins of replication for each of the five putative chromosomes of the Isarachnanthus nocturnus mitogenome and for each of the eight putative chromosomes of the Pachycerianthus magnus mitogenome. At 80,923 bp, I. nocturnus now holds the record for the largest animal mitochondrial genome reported to date. The novelty of the mitogenomic structure in Ceriantharia highlights the distinctiveness of this lineage but, because it appears to be both unique to and diverse within Ceriantharia, it is uninformative about the phylogenetic position of Ceriantharia relative to other Anthozoa. The presence of tRNAMet and tRNATrp in both ceriantharian mitogenomes supports a closer relationship between Ceriantharia and Hexacorallia than between Ceriantharia and any other cnidarian lineage, but phylogenetic analysis of the genes contained in the mitogenomes suggests that Ceriantharia is sister to a clade containing Octocorallia + Hexacorallia indicating a possible suppression of tRNATrp in Octocorallia.

Revision of the genus Carybdea (Cnidaria: Cubozoa: Carybdeidae): clarifying the identity of its type species Carybdea marsupialis.

While records of Carybdea marsupialis in the literature suggest a worldwide distribution of this species, the validity of some of these records has been questioned recently, as has the validity of some nominal Carybdea species. We inspected material of all known species of Carybdea from multiple locations (i.e. Spain, Algeria, Tunisia, Puerto Rico, California, Hawaii, Australia, South Africa, and Japan) using morphological and genetic tools to differentiate Carybdea species as well as understand their evolutionary relationships. We observed morphological differences between adult medusae of Mediterranean and Caribbean C. marsupialis; the most obvious differences were the structure of the phacellae, the structure of the pedalial canal knee bend, and the number and structure of the velarial canals. The characters of the adult Mediterranean specimens agree with the description provided by Claus (1878) for individuals of C. marsupialis from the Adriatic Sea (Italy); specimens from the Caribbean (Puerto Rico) agreed with the description of C. xaymacana by Conant (1897). Significant differences between both species were also observed in the newly released medusa stage. Further, we resolved a discord about the undefined polyp culture originating from Puerto Rico that was long considered Carybdea marsupialis but should be referred to as C. xaymacana. Although C. marsupialis is currently considered the only species of Cubozoa to occur in the Mediterranean, specimens collected in Algeria and Tunisia suggest that species of Alatinidae may also be present in the Mediterranean. Our investigations indicate that Carybdea spp. are more restricted in their geographical distribution than has been recognized historically. These findings confirm that Carybdea arborifera Maas, 1897 from Hawaii, Carybdea branchi, Gershwin Gibbons, 2009 from South Africa, Carybdea brevipedalia Kishinouye, 1891 from Japan, Carybdea confusa Straehler-Pohl, Matsumoto Acevedo, 2017 from California, Carybdea marsupialis Linnaeus, 1758 from the European Mediterranean Sea, Carybdea rastonii Haacke, 1886 from South Australia, and Carybdea xaymacana, Conant, 1897 from the Caribbean Sea are valid names representing distinct species, rather than synonyms. A taxonomic key for all valid species is provided, and a neotype for C. marsupialis is designated.

Regenerative Capacity of the Upside-down Jellyfish Cassiopea xamachana.

This study provides the first observation that umbrellar tissue can lead to the formation of virtually all body structures in jellyfish of the order Rhizostomeae. The regeneration process was observed in two specimens of the upside-down jellyfish Cassiopea xamachana Bigelow, 1892, one housed at the Vienna Zoo, Austria and the other in a laboratory at the University of São Paulo, Brazil. The process was triggered by an injury and ended with the formation of two new sets of body structures. Our observation offers evidence that C. xamachana has a hidden regenerative capacity exceeding that previously recorded.

A new species of Arachnanthus from the Red Sea (Cnidaria, Ceriantharia).

A new species of the genus Arachnanthus (Cnidaria: Ceriantharia), Arachnanthus lilith Stampar & El Didi, sp. n., is described. This species is widely distributed in the Red Sea, and recorded from 2-30 m depths. Arachnanthus lilith Stampar & El Didi, sp. n. is the fifth species of the genus and the first recorded from the Red Sea. The number of labial tentacle pseudocycles, arrangement of mesenteries, and distribution of acontioids allow the differentiation of the new species from other species of the genus.

Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters.

We have compiled available records in the literature for medusozoan cnidarians and ctenophores of South America. New records of species are also included. Each entry (i.e., identified species or still as yet not determined species referred to as "sp." in the literature) includes a synonymy list for South America, taxonomical remarks, notes on habit, and information on geographical occurrence. We have listed 800 unique determined species, in 958 morphotype entries: 5 cubozoans, 905 hydrozoans, 25 scyphozoans, 3 staurozoans, and 20 ctenophores. Concerning nomenclatural and taxonomical decisions, two authors of this census (Miranda, T.P. & Marques, A.C.) propose Podocoryna quitus as a nomen novum for the junior homonym Hydractinia reticulata (Fraser, 1938a); Euphysa monotentaculata Zamponi, 1983b as a new junior synonym of Euphysa aurata Forbes, 1848; and Plumularia spiralis Milstein, 1976 as a new junior synonym of Plumularia setacea (Linnaeus, 1758). Finally, we also reassign Plumularia oligopyxis Kirchenpauer, 1876 as Kirchenpaueria oligopyxis (Kirchenpauer, 1876) and Sertularella margaritacea Allman, 1885 as Symplectoscyphus margaritaceus (Allman, 1885).

A new species of Diadumene (Actiniaria: Diadumenidae) from the subtropical coast of Brazil.

Diadumene paranaensis n. sp., collected from the Yacht Club of Paranaguá (Paranaguá Bay, Paraná State, southern Brazil), is described as a new species of sea anemone, based on external and internal morphology, cnidome, and molecular data for 16S/CO3 mitochondrial DNA. This species is partially similar to D. cincta due to the presence of macrobasic p-amastigophores in the tentacles, but is distinguished by the cinclides arranged in longitudinal rows and microbasic p-amastigophores in the acontia.

A new species of Pachycerianthus (Cnidaria, Anthozoa, Ceriantharia) from Tropical Southwestern Atlantic.

A new species of Pachycerianthus (Cnidaria: Ceriantharia) is described from the Brazilian coast of the southwestern Atlantic Ocean. Pachycerianthus schlenzae sp. nov. is found in fine sand or mud in shallow waters of Abrolhos and Royal Charlotte Bank. The new species is only known from this area and is most notably different from other species of the genus Pachycerianthus in mesentery arrangement and cnidome. In addition to the description, we provide some biological data collected from individuals cultivated under laboratory conditions.

A new Fenestrulina (Bryozoa, Cheilostomata) commensal with tube-dwelling anemones (Cnidaria, Ceriantharia) in the tropical southwestern Atlantic.

A new species of cheilostome bryozoan, Fenestrulina commensalis n. sp., was collected in December 2008 by scuba at 5-10 meters depth at Guaibura Beach, Guarapari, Espírito Santo state, southeastern Brazil. The specimen was found associated with tubes of the cerianthid Pachycerianthus sp., representing the first commensal association between a bryozoan and a tube-dwelling anemone. Fenestrulina commensalis n. sp. is the third species of the genus found in Brazilian waters; it is distinguished from other Atlantic species of Fenestrulina by its small angular orificial condyles, a single oral spine and basal anchoring rhizoids arising from abfrontal pore chambers. Morphological adaptations to encrust the tubes of cerianthids include anchoring rootlets and weakly contiguous zooids. These morphological features allow the colony the flexibility to grow around the tube and feed relatively undisturbed by silt and detritus, being raised well above the soft-sediment substratum in which the tube-anemone grows.

Fast-evolving mitochondrial DNA in Ceriantharia: a reflection of hexacorallia paraphyly?

The low evolutionary rate of mitochondrial genes in Anthozoa has challenged their utility for phylogenetic and systematic purposes, especially for DNA barcoding. However, the evolutionary rate of Ceriantharia, one of the most enigmatic "orders" within Anthozoa, has never been specifically examined. In this study, the divergence of mitochondrial DNA of Ceriantharia was compared to members of other Anthozoa and Medusozoa groups. In addition, nuclear markers were used to check the relative phylogenetic position of Ceriantharia in relation to other Cnidaria members. The results demonstrated a pattern of divergence of mitochondrial DNA completely different from those estimated for other anthozoans, and phylogenetic analyses indicate that Ceriantharia is not included within hexacorallians in most performed analyses. Thus, we propose that the Ceriantharia should be addressed as a separate clade.

Evolutionary diversification of banded tube-dwelling anemones (Cnidaria; Ceriantharia; Isarachnanthus) in the Atlantic Ocean.

The use of molecular data for species delimitation in Anthozoa is still a very delicate issue. This is probably due to the low genetic variation found among the molecular markers (primarily mitochondrial) commonly used for Anthozoa. Ceriantharia is an anthozoan group that has not been tested for genetic divergence at the species level. Recently, all three Atlantic species described for the genus Isarachnanthus of Atlantic Ocean, were deemed synonyms based on morphological simmilarities of only one species: Isarachnanthus maderensis. Here, we aimed to verify whether genetic relationships (using COI, 16S, ITS1 and ITS2 molecular markers) confirmed morphological affinities among members of Isarachnanthus from different regions across the Atlantic Ocean. Results from four DNA markers were completely congruent and revealed that two different species exist in the Atlantic Ocean. The low identification success and substantial overlap between intra and interspecific COI distances render the Anthozoa unsuitable for DNA barcoding, which is not true for Ceriantharia. In addition, genetic divergence within and between Ceriantharia species is more similar to that found in Medusozoa (Hydrozoa and Scyphozoa) than Anthozoa and Porifera that have divergence rates similar to typical metazoans. The two genetic species could also be separated based on micromorphological characteristics of their cnidomes. Using a specimen of Isarachnanthus bandanensis from Pacific Ocean as an outgroup, it was possible to estimate the minimum date of divergence between the clades. The cladogenesis event that formed the species of the Atlantic Ocean is estimated to have occured around 8.5 million years ago (Miocene) and several possible speciation scenarios are discussed.