Five new species and two new genera of xenophyophores (Foraminifera: Rhizaria) from part of the abyssal equatorial Pacific licensed for polymetallic nodule exploration

Based on a combination of morphological and molecular data, we describe five new species and two new genera of xenophyophores from the Clarion–Clipperton Zone (abyssal eastern Pacific), an area with commercially valuable seafloor deposits of polymetallic nodules. Bizarria bryiformis gen. et sp. nov. displays unusual features, notably an organic-walled test, largely devoid of agglutinated particles, comprising interconnected branches growing upwards from the nodule substrate; the bases of the branches contain dark masses of waste material (stercomare) and pale strands of cytoplasm (granellare), the whitish, tuft-like extremities contain sediment particles. Tendalia reteformis gen. et sp. nov. forms a delicate network of agglutinated tubes. Shinkaiya contorta sp. nov. is characterized by a contorted, partly reticulated plate-like test while the simpler plate-like test of Galatheammina interstincta sp. nov. combines characters typical of Galatheammina and Psammina. In Semipsammina mattaeformis sp. nov., a thin, delicate test with one or more tubular extensions forms a flat canopy over the mat-like stercomare encrusting the nodule substrate. Tendalia reteformis and S. contorta are free-living; the other species are sessile on nodules. Together, they illustrate the considerable morphological diversity of xenophyophores in a region where they dominate the megafauna, and highlight some major taxonomic challenges posed by these giant monothalamous foraminifera

The genetic diversity, phylogeography and morphology of Elphidiidae (Foraminifera) in the Northeast Atlantic

Genetic characterisation (SSU rRNA genotyping) and Scanning ElectronMicroscope (SEM) imaging of individualtests were used in tandem to determine the modern species richness of the foraminiferal family Elphidiidae(Elphidium, Haynesina and related genera) across the Northeast Atlantic shelf biomes. Specimens were collectedat 25 locations fromthe High Arctic to Iberia, and a total of 1013 individual specimenswere successfully SEMimagedand genotyped. Phylogenetic analyses were carried out in combination with 28 other elphidiid sequencesfrom GenBank and seventeen distinct elphidiid genetic types were identified within the sample set, sevenbeing sequenced for the first time. Genetic types cluster into sevenmain cladeswhich largely represent their generalmorphologicalcharacter. Differences between genetic types at the genetic, morphological and biogeographiclevels are indicative of species level distinction. Their biogeographic distributions, in combination with elphidiidSSU sequences from GenBank and high resolution images from the literature show that each of them exhibitsspecies-specific rather than clade-specific biogeographies. Due to taxonomic uncertainty and divergent taxonomicconcepts between schools, we believe that morphospecies names should not be placed onto molecularphylogenies unless both the morphology and genetic type have been linked to the formally named holotype,or equivalent. Based on strictmorphological criteria,we advocate using only a three-stage approach to taxonomyfor practical application in micropalaeontological studies. It comprises genotyping, the production of a formalmorphological description of the SEM images associated with the genetic type and then the allocation of themost appropriate taxonomic name by comparison with the formal type description. Using this approach, wewere able to apply taxonomic names to fifteen genetic types. One of the remaining two may be potentially cryptic,and one is undescribed in the literature. In general, the phylogeographic distribution is in agreement with ourknowledge of the ecology and biogeographical distribution of the corresponding morphospecies, highlighting thegenerally robust taxonomic framework of the Elphidiidae in time and space

FIGURE 1 in Flexammina islandica gen. nov. sp. nov. and some new phylotypes of monothalamous foraminifera from the coast of Iceland

FIGURE 1. Phylogenetic tree of monothalamous foraminifera based on 3' partial SSU rDNA sequences from our database and some GenBank sequences derived from environmental clone libraries (marked by the 'ENV' prefix and highlighted in bold). New sequences obtained in this study are marked by frames with the names of corresponding phylotypes / species given in bold; the name of every new sequence consists of two parts separated by dashes (foraminiferal DNA collection number and clone number). Foraminiferal DNA collection numbers and GenBank accession numbers (if available) are also shown for other sequences. Major monothalamid clades are indicated in accordance with Pawlowski et al. (2002, 2011)

Eyes of the Deep-sea Floor: The Integrative Taxonomy of the Foraminiferal Genus Vanhoeffenella

Vanhoeffenella is a common deep-sea monothalamous foraminifer, some species of which have a unique eye-like test morphology. Owing to its world-wide distribution, it has been recorded numerous times since the “heroic age” of the deep-sea exploration in the early 20th century. So far, only 4 species have been described, and no attempts have been made to estimate the real diversity of this peculiar genus. Over the last fifteen years, we have collected specimens of Vanhoeffenella from various deep-sea areas, providing the basis for an integrative taxonomy and biogeography of this genus. Here, we clarify the phylogenetic position of Vanhoeffenella and give an account of its diversity in the Atlantic, Arctic and Southern Oceans (the Weddell Sea) as revealed by genetic marker (SSU rDNA) and morphology. Our study shows that Vanhoeffenella branches within Clade F of monothalamids and incorporates at least 10 putative species. Some could be distinguished by either morphological or molecular features, but only the integrative taxonomic approach provides a robust way to assess their diversity. We examine the new material of the type species (V. gaussi Rhumbler), redescribe the poorly-known V. oculus Earland and describe formally a fifth species, V. dilatata sp. nov

Toxisarcon taimyr sp. nov., a new large monothalamous foraminifer from the Kara Sea inner shelf

A large monothalamous foraminiferan, Toxisarcon taimyr sp. nov., has been isolated from the benthic samples from the Kara Sea inner shelf near the mouth of Yenisey river estuary, at a depth of 50–100 m. In its overall morphology, the new species closely resembles T. synsuicidica, one of the two species of Toxisarcon described to date. It possesses a large irregularly shaped cell body, covered by a thin layer of a fibrous organic coating. Numerous reticulopodia typically extend from all over the cell surface; the species is very motile and rapidly changes cell shape. Long and thick reticulopodial bundles form in the direction of movement. In the phylogenetic tree based on partial small-subunit ribosomal DNA (SSU rDNA) sequences, T. taimyr branches together with the two other known species of Toxisarcon within the clade C of monothalamous foraminifera

Abyssal foraminifera attached to polymetallic nodules from the eastern Clarion Clipperton Fracture Zone: a preliminary description and comparison with North Atlantic dropstone assemblages

We examined encrusting organisms on seven polymetallic nodules from the eastern Clarion Clipperton Fracture Zone (CCFZ, ?4070 m water depth, eastern equatorial Pacific). Apart from occasional sponges and a single bryozoan, all the organisms were foraminifera or foraminifera-like protists. A total of 75 morphotypes (presumed to be morphospecies) was recognised, with between 9 and 19 being present on individual nodules. Additional species were observed during shipboard examination of the nodules, bringing the total number of species to 86. The assemblage was dominated by a variety of mat-like formations, clusters of patches, isolated domes, broad trails, anastomosing networks and branched or unbranched tubular structures that either lay flat against the nodule surface or projected away from the surface. These forms were interpreted as monothalamous foraminifera (monothalamids). Most have mainly agglutinated walls but a few are predominately organic. Some can be assigned to the Komokiacea (notably the genus Chondrodapis) or families such as the Hemisphaeramminidae ('domes'), while others (e.g. many of the mats and patch-like forms) are difficult to place into existing monothalamid groupings. Some of the branching and anastomosing tubes resemble the genus Rhizammina. The most easily recognisable morphotypes include Telammina, in which tiny chambers are linked by extremely thin tubes to form a network, and sinuous orange tubes that incorporate sponge spicules and can be assigned to the genus Saccorhiza based on the occasional presence of a proloculus. Polythalamous foraminifera are also fairly common. They include various calcareous species (mainly Cibicides spp.), as well as agglutinated forms such as Hormosina, ?Placopsilina and trochamminaceans. Similar assemblages, including some morphospecies that are clearly identical to ours, have been described previously from somewhat deeper sites (4500–5000 m) in the CCFZ. In order to explore distributions at a global scale, we compared our Pacific nodule assemblages to foraminifera attached to ice-rafted dropstones from several deep seamounts in the abyssal northeast Atlantic Ocean, mainly on the Porcupine Abyssal Plain (4630–4680 m depth) with additional material from the BIOTRANS area (3796–4351 m). These hosted superficially similar assemblages of mats, tubular forms, komokiaceans and polythalamous calcareous and agglutinated foraminifera. However, apart from the sometimes extensive development of Telammina networks, there were no morphospecies in common between the Pacific and Atlantic assemblages. These preliminary observations, based on limited material, suggest that most of the foraminiferal morphospecies encrusting hard substrates are widely distributed at regional scales in the abyssal Pacific but not necessarily at global scales. The study of abyssal encrusting assemblages poses considerable challenges. Priorities for the future include the development of reliable methods for distinguishing living and dead individuals, and molecular approaches to clarifying the taxonomic affinities of novel morphotypes