Catalog to families, genera, and species of orders Actiniaria and Corallimorpharia (Cnidaria: Anthozoa).

This book inventories all available (and some unavailable) names in the family, genus, and species groups of extant members of orders Actiniaria and Corallimorpharia [cnidarian subclass Hexacorallia (Zoantharia) of class Anthozoa], providing a benchmark of names, their status, and taxon membership. I have attempted to make the compilation complete as of 2010; some names created after 2010 are included. The book is derived from a database I compiled that was available through a website. Most of the book is from the literature that defines taxa and documents their geographic distribution-primarily publications on nomenclature, taxonomy, and biogeography, but also some on ecology, pharmacology, reproductive biology, physiology, etc. of anemones (the common name for these groups); the reference section comprises 845 entries. As for previous anemone catalogs, this contains taxonomic as well as nomenclatural information,  the former based on subjective opinion of working biologists, the latter objectively verifiable and unchanging (except by action of the International Commission on Zoological Nomenclature).        Each family-group name, genus-group name, and original combination for species-group names has an entry. The entry contains the bibliographic reference to the publication in which each name was made available. This book contains for Corallimorpharia seven family names (four considered valid [57%]), 20 generic names (10 considered valid [50%] and one unavailable), and 65 species names (46 considered valid [70%]). It contains for Actiniaria 86 family names (50 considered valid [58%] and three unavailable), 447 generic names (264 considered valid [59%] and two unavailable), and 1427 species names (1101 considered valid [77%] and nine unavailable). Type specimens are inventoried from more than 50 natural history museums in Africa, Australia, Europe, New Zealand, and North America, including those with the largest collections of anemones; the geographic sources of specimens that were the bases of new names are identified. I resolve some nomenclatural issues, acting as First Reviser. A few taxonomic opinions are published for the first time. I have been unable to resolve a small number of problematic names having both nomenclatural and taxonomic problems. Molecular phylogenetic analyses are changing assignment of genera to families and species to genera. Systematics may change, but the basics of nomenclature remain unchanged in face of such alterations.        All actions are in accord with the principles of nomenclature enunciated in the International Code of Zoological Nomenclature. These include the type concept, the Principle of Coordination, and the Principle of Priority. Nomenclatural acts include the creation of new replacement names; seven actiniarian generic names and one species name that are junior homonyms but have been treated as valid are replaced and an eighth new genus name is created. I designate type species for two genera. Except for published misspellings, names are rendered correctly according to the International Code of Zoological Nomenclature; I have altered spelling of some species names to conform to orthographic regulations. I place several species that had been assigned to genera now considered junior synonyms in the genus to which the type species was moved; experts on these anemones should determine whether those generic placements, which follow the nomenclatural rules, are taxonomically appropriate.        This inventory can be a useful starting point in assembling the literature and trying to understand the rationale for the creation and use of names for the taxonomic matters yet to be resolved.  Some nomenclatural conundra will not be resolved until taxonomic uncertainties are. A taxonomist familiar with the animals needs to ascertain whether the published synonymies are justified. If so, the senior synonym should be used, which, in many instances, will involve determining the proper generic assignment of the species and the correct rendering of the name; if changing the name would be disruptive, retaining the junior name would require an appeal to the Commission (Code Article 23.11).

The List of Available Names (LAN): A new generation for stable taxonomic names in zoology?

The List of Available Names in Zoology (LAN) is an inventory of names with specific scope in time and content, presented and approved in parts, and constituted as a cumulative index of names available for use in zoological nomenclature. It was defined in Article 79 in the fourth edition of the International Code of Zoological Nomenclature. The LAN is likely to gain importance with the development of the online Official Registry for Zoological Nomenclature (ZooBank) as it is potentially a source of many nomenclaturally certified names. Article 79 describes the deliberative process for adding large numbers of names to the LAN simultaneously, detailing steps and chronology for submission of a candidate Part to the LAN and consideration of a candidate Part by the public and Commission, but it is largely mute about the contents of a candidate Part. It does make clear that a name within the scope of a Part but not on the LAN has no nomenclatural standing, even if it had previously been considered available, thereby preventing long-forgotten names from displacing accepted ones and the accumulation of nomina dubia. Thus, for taxa on the LAN, nomenclatural archaeology - the resurrecting of old unused names to replace by priority names in current usage - will not be worthwhile. Beyond that, it has been unclear if Article 79 is intended to document every available name known within the scope of the Part, or if its intention is to pare the inventory of available names within the scope of the Part. Consideration by the Commission and two committees to deal with the LAN have defined steps to implement Article 79 with the latter intent. Procedures for consideration of a candidate Part are defined in a manual, published as an appendix in this volume.

Latitudinal diversity of sea anemones (Cnidaria: Actiniaria).

We sought to determine if the global distribution of sea anemones (cnidarian order Actiniaria) conforms to the classic pattern of biogeography--taxon richness at the equator with attenuation toward the poles--a pattern that is derived almost entirely from data on terrestrial plants and animals. We plotted the empirical distribution of species occurrences in 10° bands of latitude based on published information, then, using the Chao2 statistic, inferred the completeness of that inventory. We found the greatest species richness of sea anemones at 30-40° N and S, with lower numbers at tropical latitudes and the fewest species in polar areas. The Chao2 statistic allowed us to infer that the richness pattern we found is not due to particularly poor knowledge of tropical sea anemones. No 10° band of latitude has less than 60% of the theoretical number of species known, but for only about half of them could we reject the null hypothesis (P = 0.05) that information is complete; anemone diversity is best documented at high latitudes. We infer that the 1089 valid species currently known constitute about 70% of the theoretical total of about 1500 species of Actiniaria. The distribution pattern of sea anemone species resembles that of planktonic foraminiferans and benthic marine algae, although planktonic bacteria, marine bivalves, and shallow and deep scleractinian corals show the terrestrial pattern of equatorial richness attenuating with latitude. Sea anemone species richness is complementary to that of scleractinian corals at many scales; our findings affirm it at the global scale.

The magnitude of global marine species diversity.

BACKGROUND: The question of how many marine species exist is important because it provides a metric for how much we do and do not know about life in the oceans. We have compiled the first register of the marine species of the world and used this baseline to estimate how many more species, partitioned among all major eukaryotic groups, may be discovered. RESULTS: There are ∼226,000 eukaryotic marine species described. More species were described in the past decade (∼20,000) than in any previous one. The number of authors describing new species has been increasing at a faster rate than the number of new species described in the past six decades. We report that there are ∼170,000 synonyms, that 58,000-72,000 species are collected but not yet described, and that 482,000-741,000 more species have yet to be sampled. Molecular methods may add tens of thousands of cryptic species. Thus, there may be 0.7-1.0 million marine species. Past rates of description of new species indicate there may be 0.5 ± 0.2 million marine species. On average 37% (median 31%) of species in over 100 recent field studies around the world might be new to science. CONCLUSIONS: Currently, between one-third and two-thirds of marine species may be undescribed, and previous estimates of there being well over one million marine species appear highly unlikely. More species than ever before are being described annually by an increasing number of authors. If the current trend continues, most species will be discovered this century.

Stylobates birtlesi sp. n., a new species of carcinoecium-forming sea anemone (Cnidaria, Actiniaria, Actiniidae) from eastern Australia.

We describe a new species of carcinoecium-forming sea anemone, Stylobates birtlesisp. n., from sites 590-964 m deep in the Coral Sea, off the coast of Queensland, Australia. An anemone of this genus settles on a gastropod shell inhabited by a hermit crab, then covers and extends the shell to produce a chitinous structure termed a carcinoecium. Stylobates birtlesisp. n. is symbiotic with the hermit crab Sympagurus trispinosus (Balss, 1911). The nature of marginal sphincter muscle and nematocyst size and distribution distinguish Stylobates birtlesi sp. n. from other species in the genus. The four known species of Stylobates are allopatric, each inhabiting a separate ocean basin of the Indo-West Pacific. We also extend the known range of Stylobates loisetteae in the Indian Ocean off the coast of Western Australia.

An overview of marine biodiversity in United States waters.

Marine biodiversity of the United States (U.S.) is extensively documented, but data assembled by the United States National Committee for the Census of Marine Life demonstrate that even the most complete taxonomic inventories are based on records scattered in space and time. The best-known taxa are those of commercial importance. Body size is directly correlated with knowledge of a species, and knowledge also diminishes with distance from shore and depth. Measures of biodiversity other than species diversity, such as ecosystem and genetic diversity, are poorly documented. Threats to marine biodiversity in the U.S. are the same as those for most of the world: overexploitation of living resources; reduced water quality; coastal development; shipping; invasive species; rising temperature and concentrations of carbon dioxide in the surface ocean, and other changes that may be consequences of global change, including shifting currents; increased number and size of hypoxic or anoxic areas; and increased number and duration of harmful algal blooms. More information must be obtained through field and laboratory research and monitoring that involve innovative sampling techniques (such as genetics and acoustics), but data that already exist must be made accessible. And all data must have a temporal component so trends can be identified. As data are compiled, techniques must be developed to make certain that scales are compatible, to combine and reconcile data collected for various purposes with disparate gear, and to automate taxonomic changes. Information on biotic and abiotic elements of the environment must be interactively linked. Impediments to assembling existing data and collecting new data on marine biodiversity include logistical problems as well as shortages in finances and taxonomic expertise.

Formation of the apical flaps in nematocysts of sea anemones (cnidaria: actiniaria).

Using scanning and transmission electron microscopy, we studied formation of the structure at the apical end of sea anemone nematocysts through which the tubule everts at discharge. In anemones of the genus Metridium, we found that each of the three solid triangular apical flaps comprises two layers that are continuous with those of the capsule wall: the electron-lucent inner layer is bound to the electron-dense outer layer. The two-layer structure is obvious in some discharged capsules in which, perhaps due to fixation, the layers part at the flap's periphery. Before the nematocyst discharges, a channel leads from a pore at the tip of the joined flaps into the lumen of the inverted tubule. The thin laminate layer that coats each flap lines the channel. The base of the nematocyst tubule adheres to the capsule wall near the capsule's apical end, and a branch of the tubule underlies part of the laminate layer that coats the flaps. Thus the tubule is not continuous with the capsule wall but structurally separate from it. This helps reconcile differences in understanding of the number of layers constituting the capsule wall, and makes clear that the tubule should be considered part of the capsule contents.

Structural diversity, systematics, and evolution of cnidae.

Cnidae are secreted by the Golgi apparatus of all cnidarians and only cnidarians. Of the three categories of cnidae (also called cnidocysts), nematocysts occur in all cnidarians, and are the means by which cnidarians defend themselves and obtain prey; spirocysts and ptychocysts are restricted to a minority of major taxa. A cnida discharges by eversion of its tubule; venom may be associated with the tubule of a nematocyst. About 30 major morphological types of nematocysts are recognized, but no single nomenclature for them is accepted. Function seems not to correlate tightly with morphology--nematocysts of at least some types are used both offensively and defensively. Similarly, it is not clear if morphology correlates with toxicity. Some types of nematocysts are taxonomically diagnostic whereas others are widespread. Nonetheless, an inventory of types of cnidae (the cnidom), with their distribution and size, is an essential component of most taxonomic descriptions. Complicating the taxonomic value of cnidae are the facts that not all members of a species may have the same types of cnidae, even at the same life-cycle stage, and size of nematocysts of a species may vary geographically and with size of individual. The diversity of nematocysts is so great and the features within each major type are so variable that homologies have not been determined. Nematocyst complement, morphology, and size likely reflect both phylogeny and biology; the feedback between the two may confound analysis. Although cnidae are valuable in taxonomy of at least some groups, more understanding of the forces that affect them is needed for their systematic and phylogenetic value to be understood and their potential as indicators of evolution to be realized.

Finding NEMO: nestedness engendered by mutualistic organization in anemonefish and their hosts.

The interaction structure of mutualistic relationships, in terms of relative specialization of the partners, is important to understanding their ecology and evolution. Analyses of the mutualistic interaction between anemonefish and their host sea anemones show that the relationship is highly nested in structure, generalist species interacting with one another and specialist species interacting mainly with generalists. This supports the hypothesis that the configuration of mutualistic interactions will tend towards nestedness. In this case, the structure of the interaction is at a much larger scale than previously hypothesized, across more than 180 degrees of longitude and some 60 degrees of latitude, probably owing to the pelagic dispersal capabilities of these species in a marine environment. Additionally, we found weak support for the hypothesis that geographically widespread species should be more generalized in their interactions than species with small ranges. This study extends understanding of the structure of mutualistic relationships into previously unexplored taxonomic and physical realms, and suggests how nestedness analysis can be applied to the conservation of obligate species interactions.