Cryptic species in time and space: an assessment of cryptic diversity within eight nominal species of Hydrozoa (Cnidaria).

Sampling in multiple localities, coupled with molecular barcoding, has shown that nominal species with wide geographical distribution often harbour local cryptic species in allopatry. Cryptic species in sympatry, however, can be easily missed if they have different seasonality, because they can be identified only through long-term frequent sampling (i.e. sampling through time of the same species in the same location). This is especially true in planktonic invertebrates that exhibit strong seasonality. By integrating mitochondrial 16S sequences of eight species of Hydrozoa (Cnidaria) collected weekly for a year in one Gulf of Mexico region, with sequences gathered globally, we investigate the presence of cryptic species within a temporal gradient (regionally) and on a spatial (worldwide) scale. We find that eight species of Hydrozoa are composed of 28 cryptic species, with 16 of them appearing in sympatry but with non-overlapping seasonality. The high number of sympatric cryptic species could only be discovered through extensive and prolonged regional sampling efforts. The bi-dimensional cryptic diversity (in time and space) highlighted in this study is essential for understanding processes of evolution, biogeography dispersal in the sea, and for more realistic biodiversity assessments.

Demystifying Cassiopea species identity in the Florida Keys: Cassiopea xamachana and Cassiopea andromeda coexist in shallow waters.

The phylogeny of the Upside-Down Jellyfish (Cassiopea spp.) has been revised multiple times in its history. This is especially true in the Florida Keys, where much of the Cassiopea stock for research and aquarium trade in the United States are collected. In August 2021, we collected 55 Cassiopea medusae at eight shallow water sites throughout the Florida Keys and sequenced COI, 16S, and 28S genes. Mitochondrial genes demonstrate that the shallow waters in Florida are inhabited by both Cassiopea xamachana and a non-native Cassiopea andromeda lineage, identified in multispecies assemblages at least thrice. While C. xamachana were present at all sites, the C. andromeda-mitotype individuals were present at only a minority of sites. While we cannot confirm hybridization or lack thereof between the C. xamanchana and C. andromeda lineages, these previously unknown multispecies assemblages are a likely root cause for the confusing and disputed COI-based species identities of Cassiopea in the Florida Keys. This also serves as a cautionary note to all Cassiopea researchers to barcode their individuals regardless of the location in which they were collected.

An integrative re-evaluation of Typhlatya shrimp within the karst aquifer of the Yucatán Peninsula, Mexico.

The Yucatán Peninsula, Mexico is a carbonate platform well-known for extensive karst networks of densely stratified aquifer ecosystems. This aquifer supports diverse anchialine fauna, including species of the globally distributed anchialine shrimp genus Typhlatya (Atyidae). Four species (T. campecheae, T. pearsei, T. dzilamensis and T. mitchelli) are endemic to the Peninsula, of which three are federally listed in Mexico. This first integrative evaluation (i.e., molecular, morphological, broad geographic and type locality sampling, and environmental data) of Yucatán Typhlatya reveals considerable species identity conflict in prior phylogenetic assessments, broad species ranges, syntopy within cave systems and five genetic lineages (of which two are new to science). Despite sampling from the type locality of endangered T. campecheae, specimens (and molecular data) were indistinguishable from vulnerable T. pearsei. Ancestral/divergence reconstructions support convergent evolution of a low-salinity ancestor for a post-Paleogene arc Yucatán + Cuba Typhlatya clade within the anchialine Atyidae clade. A secondary adaptation for the coastal-restricted euryhaline (2-37 psu), Typhlatya dzilamensis (unknown conservation status) was identified, while remaining species lineages were low-salinity (< 5 psu) adapted and found within the meteoric lens of inland and coastal caves. This study demonstrates the need for integrative/interdisciplinary approaches when conducting biodiversity assessments in complex and poorly studied aquifers.

Cellular Reprogramming and Immortality: Expression Profiling Reveals Putative Genes Involved in Turritopsis dohrnii's Life Cycle Reversal.

To gather insight on the genetic network of cell reprogramming and reverse development in a nonmodel cnidarian system, we produced and annotated a transcriptome of the hydrozoan Turritopsis dohrnii, whose medusae respond to damage or senescence by metamorphosing into a juvenile stage (the polyp), briefly passing through an intermediate and uncharacterized stage (the cyst), where cellular transdifferentiation occurs. We conducted sequential and pairwise differential gene expression (DGE) analyses of the major life cycle stages involved in the ontogenetic reversal of T. dohrnii. Our DGE analyses of sequential stages of T. dohrnii's life cycle stages show that novel and characterized genes associated with aging/lifespan, regulation of transposable elements, DNA repair, and damage response, and Ubiquitin-related processes, among others, were enriched in the cyst stage. Our pairwise DGE analyses show that, when compared with the colonial polyp, the medusa is enriched with genes involved in membrane transport, the nervous system, components of the mesoglea, and muscle contraction, whereas genes involved in chitin metabolism and the formation of the primary germ layers are suppressed. The colonial polyp and reversed polyp (from cyst) show significant differences in gene expression. The reversed polyp is enriched with genes involved in processes such as chromatin remodeling and organization, matrix metalloproteinases, and embryonic development whereas suppressing genes involved in RAC G-protein signaling pathways. In summary, we identify genetic networks potentially involved in the reverse development of T. dohrnii and produce a transcriptome profile of all its life cycle stages, and paving the way for its use as a system for research on cell reprogramming.

Planktonic associations between medusae (classes Scyphozoa and Hydrozoa) and epifaunal crustaceans.

Jellyfish are known to carry various epibionts, including many of the subphylum Crustacea. However, the associations between gelatinous zooplankton and other invertebrates have been chronically overlooked. Crustacea, a massive clade of economically, ecologically, and culturally important species, includes many taxa that utilize gelatinous zooplankton for food, transport, and protection as both adults and juveniles. Here we compile 211 instances of epifaunal crustaceans recorded on Hydromedusae and Scyphomedusae from a century of literature. These include 78 identified crustacean species in 65 genera across nine orders found upon 37 Hydromedusa species and 48 Scyphomedusae. The crustacean life stage, location, nature of the association with the medusa, years, months, and depths are compiled to form a comprehensive view of the current state of the literature. Additionally, this review highlights areas where the current literature is lacking, particularly noting our poor understanding of the relationships between juvenile crabs of commercially valuable species and medusae.

Fluctuation and diversity of Hydromedusae (Hydrozoa, Cnidaria) in a highly productive region of the Gulf of Mexico inferred from high frequency plankton sampling.

Hydrozoa medusae undergo blooms and seasonal fluctuations; however the drivers of such fluctuations are unknown. To understand how medusa populations fluctuate in response to seasonal factors such as temperature, salinity, dissolved oxygen, and chlorophyll a, and to enhance our taxonomic knowledge of Hydrozoa in Galveston Bay (TX), we performed frequent plankton sampling from September 2015 to September 2016. We collected 1,321 medusae in 190 sampling days. Using molecular barcoding and morphological analyses we identified 25 species, of which 21 are a first record for Galveston Bay and eight for the Gulf of Mexico. Daily medusa abundance is non-linearly related to temperature, with peak abundance estimated with multivariate regression analysis at approximately 21C. The role that temperature plays in driving medusa abundance has implications for future climate change scenarios, given that temperature in the Gulf of Mexico is expected to rise 4 °C by the end of the century. We also show that the biodiversity of the Galveston Bay and the Gulf of Mexico is underestimated and that molecular barcoding is an important and efficient tool to identify large number of medusae. We conclude that dense plankton sampling is necessary to capture both diversity and abundance of planktonic medusae.

Transcriptome Characterization of Reverse Development in Turritopsis dohrnii (Hydrozoa, Cnidaria).

Medusae of Turritopsis dohrnii undergo reverse development in response to physical damage, adverse environmental conditions, or aging. Senescent, weakened or damaged medusae transform into a cluster of poorly differentiated cells (known as the cyst stage), which metamorphose back into a preceding life cycle stage, the polyp. During the metamorphosis, cell transdifferentiation occurs. The cyst represents the intermediate stage between a reverting medusa and a healthy polyp, during which cell transdifferentiation and tissue reorganization take place. Here we characterize and compare the transcriptomes of the polyp and newborn medusa stages of T. dohrnii with that of the cyst, to identify biological networks potentially involved in the reverse development and transdifferentiation processes. The polyp, medusa and cyst of T. dohrnii were sequenced through Illumina RNA-sequencing and assembled using a de novo approach, resulting in 92,569, 74,639 and 86,373 contigs, respectively. The transcriptomes were annotated and comparative analyses among the stages identified biological networks that were significantly over-and under-expressed in the cyst as compared to the polyp and medusa stages. Biological processes that occur at the cyst stage such as telomerase activity, regulation of transposable elements and DNA repair systems, and suppression of cell signaling pathways, mitotic cell division and cellular differentiation and development may be involved in T. dohrnii's reverse development and transdifferentiation. Our results are the first attempt to understand T. dohrnii's life-cycle reversal at the genetic level, and indicate possible avenues of future research on developmental strategies, cell transdifferentiation, and aging using T. dohrnii as a non-traditional in vivo system.

Turritopsis fascicularis Fraser, 1943 (Cnidaria: Hydrozoa): redescription and discussion of its phylogenetic position within the genus.

Turritopsis fascicularis Fraser, 1943 was first described off Alligator Reef, Florida, USA, at a depth of 216 m. Presumably a deep-sea species, its validity has often been questioned due to the scarcity of available records. In this paper, T. fascicularis is re-described from some mature colonies from the upper slope of the Gulf of Mexico. Furthermore, new pictures of the colony, polyps, and medusa buds, are provided. A ~600bp sequence of the large ribosomal subunit of the mitochondrial RNA (lsu-rRNA, 16S), also known as the Hydrozoan barcoding molecule, is used for the first time to confirm the validity of T. fascicularis as a species, and analyze its phylogenetic position within the genus Turritopsis.

TaxaGloss - A Glossary and Translation Tool for Biodiversity Studies.

BACKGROUND: Correctly identifying organisms is key to most biological research, and is especially critical in areas of biodiversity and conservation. Yet it remains one of the greatest challenges when studying all but the few well-established model systems. The challenge is in part due to the fact that most species have yet to be described, vanishing taxonomic expertise and the relative inaccessibility of taxonomic information. Furthermore, identification keys and other taxonomic resources are based on complex, taxon-specific vocabularies used to describe important morphological characters. Using these resources is made difficult by the fact that taxonomic documentation of the world's biodiversity is an international endeavour, and keys and field guides are not always available in the practitioner's native language. NEW INFORMATION: To address this challenge, we have developed a publicly available on-line illustrated multilingual glossary and translation tool for technical taxonomic terms using the Symbiota Software Project biodiversity platform. Illustrations, photographs and translations have been sourced from the global community of taxonomists working with marine invertebrates and seaweeds. These can be used as single-language illustrated glossaries or to make customized translation tables. The glossary has been launched with terms and illustrations of seaweeds, tunicates, sponges, hydrozoans, sea anemones, and nemerteans, and already includes translations into seven languages for some groups. Additional translations and development of terms for more taxa are underway, but the ultimate utility of this tool depends on active participation of the international taxonomic community.

Barcoding Techniques Help Tracking the Evolutionary History of the Introduced Species Pennaria disticha (Hydrozoa, Cnidaria).

The Christmas tree hydroid Pennaria disticha is listed as one of the most common introduced species in Hawaii. Firstly reported in Kaneohe Bay (Oahu) in 1928, it is now established throughout the entire archipelago, including the Northwestern Hawaiian Islands, a U.S. National Monument and World Heritage site. The Hawaiian population of P. disticha has also been reported as being the source of further introductions to Palmyra Atoll in the U.S. Line Islands. Using a phylogenetic hypothesis based on a 611 base pair fragment of the mitochondrial 16S barcoding gene, we demonstrate that P. disticha is a complex of cryptic species, rather than one species with cosmopolitan distribution. We also show that in Hawaii there are three species of Pennaria, rather than one introduced species. Two of these species share haplotypes with specimens from distant locations such as Florida and Panama and may have been introduced, possibly from the Atlantic Ocean. A third species could either represent a lineage with nearly cosmopolitan distribution, or another introduced species. Our dataset refutes the widely accepted idea that only one lineage of P. disticha is present in Hawaii. On the contrary, P. disticha in Hawaii may be the outcome of multiple independent introductions of several morphologically undistinguishable cryptic lineages. Our results uncover an unsuspected complexity within the very common hydroid P. disticha, and highlight the need for routine use of molecular tools, such as DNA barcoding, to improve the identification and recognition of non-indigenous species.

Podocoryna loyola, n. sp. (Hydrozoa, Hydractiniidae): a probably introduced species on artificial substrate from southern Brazil.

Podocoryna loyola, a new hydractiniid species, has been found on artificial substrates in Baía de Paranaguá, southern coast of Brazil, since April 2007. Its main morphological characteristics are: (1) polymorphic colonies with reticular stolons or encrusting hydrorhiza not covered by periderm and smooth chitinous spines; (2) newly-released medusae with eight tentacles and small interradial gonads; (3) mature medusae with eight tentacles and unbranched oral lips; gastric peduncle absent. Molecular data show that P. loyola is distinct from all other examined species of Podocoryna, and from P. hayamaensis Hirohito (1988), its sister species from Japan. As the polyps having been noted only quite recently, and in having been found only on man-made objects in port areas and estuaries, the species is most likely exotic to the region.

Evolution of life cycle, colony morphology, and host specificity in the family Hydractiniidae (Hydrozoa, Cnidaria).

Biased transitions are common throughout the tree of life. The class hydrozoa is no exception, having lost the feeding medusa stage at least 70 times. The family hydractiniidae includes one lineage with pelagic medusae (Podocoryna) and several without (e.g., Hydractinia). The benthic colony stage also varies widely in host specificity and in colony form. The five-gene phylogeny presented here requires multiple transitions between character states for medusae, host specificity, and colony phenotype. Significant phylogenetic correlations exist between medusoid form, colony morphology, and host specificity. Species with nonfeeding medusae are usually specialized on a single host type, and reticulate colonies are correlated with nonmotile hosts. The history of feeding medusae is less certain. Podocoryna is nested within five lineages lacking medusae. This requires either repeated losses of medusae, or the remarkable re-evolution of a feeding medusa after at least 150 million years. Traditional ancestral reconstruction favors medusa regain, but a likelihood framework testing biased transitions cannot distinguish between multiple losses versus regain. A hypothesis of multiple losses of feeding medusae requires transient selection pressure favoring such a loss. Populations of species with feeding medusae are always locally rare and lack of feeding medusae does not result in restricted species distribution around the world.

Reversing opinions on Dollo's Law.

Dollo's Law, the idea that the loss of complex features in evolution is irreversible, is a popular concept in evolutionary biology. Here we review how application of recent phylogenetic methods, genomics and evo-devo approaches is changing our view of Dollo's Law and its underlying mechanisms. Phylogenetic studies have recently demonstrated cases where seemingly complex features such as digits and wings have been reacquired. Meanwhile, large genomics databases and evo-devo studies are showing how the underlying developmental pathways and genetic architecture can be retained after the loss of a character. With dwindling evidence for the law-like nature of Dollo's Law, we anticipate a return to Dollo's original focus on irreversibility of all kinds of changes, not exclusively losses.