Diversity and distribution patterns in high southern latitude sponges

Sponges play a key role in Antarctic marine benthic community structure and dynamics and are often a dominant component of many Southern Ocean benthic communities. Understanding the drivers of sponge distribution in Antarctica enables us to understand many of general benthic biodiversity patterns in the region. The sponges of the Antarctic and neighbouring oceanographic regions were assessed for species richness and biogeographic patterns using over 8,800 distribution records. Species-rich regions include the Antarctic Peninsula, South Shetland Islands, South Georgia, Eastern Weddell Sea, Kerguelen Plateau, Falkland Islands and north New Zealand. Sampling intensity varied greatly within the study area, with sampling hotspots found at the Antarctic Peninsula, South Georgia, north New Zealand and Tierra del Fuego, with limited sampling in the Bellingshausen and Amundsen seas in the Southern Ocean. In contrast to previous studies we found that eurybathy and circumpolar distributions are important but not dominant characteristics in Antarctic sponges. Overall Antarctic sponge species endemism is ～43%, with a higher level for the class Hexactinellida (68%). Endemism levels are lower than previous estimates, but still indicate the importance of the Polar Front in isolating the Southern Ocean fauna. Nineteen distinct sponge distribution patterns were found, ranging from regional endemics to cosmopolitan species. A single, distinct Antarctic demosponge fauna is found to encompass all areas within the Polar Front, and the sub-Antarctic regions of the Kerguelen Plateau and Macquarie Island. Biogeographical analyses indicate stronger faunal links between Antarctica and South America, with little evidence of links between Antarctica and South Africa, Southern Australia or New Zealand. We conclude that the biogeographic and species distribution patterns observed are largely driven by the Antarctic Circumpolar Current and the timing of past continent connectivity.     

The terrestrial Protozoan fauna of South Georgia

Summary 75 taxa of Protozoa (18 flagellates, 9 naked rhizopods, 20 testate rhizopods and 28 ciliates) were found in 14 samples of mineral materials, peats, soils and guano collected from the sub-Antarctic island of South Georgia. The results confirm the existence of distinct natural communities of protozoan species related to the different classes of habitat in the sub-Antarctic and maritime Antarctic, but suggest that species characteristic of the maritime Antarctic also occur in the sub-Antarctic and species characteristic of bryophyte peats also occur in soils with angiosperm vegetation. The diversity of the fauna of different habitats can be related to degree of soil development and successional stage of associated vegetation. Comparison of the data on the testate rhizopod fauna of South Georgia with those from other sub-Antarctic locations shows a clear trend of faunal pauperization with latitude.     

Areas of endemism in the Antarctic – a case study of the benthic hydrozoan genus Oswaldella (Cnidaria,Kirchenpaueriidae)

Aim The aim of this study is to investigate areas of endemism within the distribution of Oswaldella species in the Southern Ocean, thereby testing previous hypotheses and proposing alternative scenarios for Antarctic evolution. Location Southern Ocean, Antarctic and sub-Antarctic waters of southern South America. Methods We prepared a database for the 31 currently known species of the Antarctic genus Oswaldella, which includes geographical locations gathered from published taxonomic studies as well as materials from museums and expeditions. A parsimony analysis of endemicity (PAE) was used to test hypotheses of distribution patterns. Results Four areas of endemism are hypothesized: southern South America, two high Antarctic areas (eastern and western) and a larger area, mainly in western Antarctica at lower latitudes and including insular areas (but not the Balleny Islands). Main conclusions The results support, in part, previous hypotheses for the Southern Ocean region, while providing more detailed resolution. The areas of endemism may reflect both historical and ecological processes that influenced the Antarctic biota. The Magellanic area reflects the well-known affinities of the Antarctic biota with that of South America and may be a consequence of dispersal through deeper (and colder) waters, followed by speciation. The second area, the largest one, encompasses most of the insular faunas and may also be associated with deeper waters formed since 43 Ma. The third area may be explained by the development of seaways in the circum-Antarctic region beginning 50 Ma. Finally, the fourth zone, with a very poor fauna, coincides with the opening of the Tasman Strait and the formation of the Australo-Antarctic Gulf, associated with a minor wind-driven current.     

Biodiversity and biogeography of Southern Ocean pycnogonids

The pycnogonids of the Southern Ocean have been studied for almost two centuries and have played a key role in shaping previous biogeographic regions for the Antarctic benthos. The aim of this study was to assess the biogeographic patterns derived from the most current sample records of pycnogonids from the Southern Ocean and neighbouring areas. 332 species of pycnogonids from 1837 sample locations were analysed using 279 3° by 3° grid cells. We investigated richness patterns and the effect of sampling intensity at both local and regional scales, and used multivariate analysis of distribution patterns and species assemblages to define biogeographic trends. These analyses identified a distinct and isolated Antarctic pycnogonid shelf fauna which was different to that of the deep‐sea around Antarctica, the Sub‐Antarctic islands, South America or New Zealand. Within the Antarctic, we found the South Shetland Islands to be the most speciose region and a probable center of radiation for the pycnogonids. No latitudinal gradients in species richness were detected. We note that the distribution patterns observed are based upon classical taxonomy and discuss the potential for changes to these patterns with new insights from molecular techniques. We conclude that, even with the potential for cryptic species, the large‐scale biogeographic trends observed in the pycnogonids should hold true.     

Benthic ecoregionalization based on echinoid fauna of the Southern Ocean supports current proposals of Antarctic Marine Protected Areas under IPCC scenarios of climate change

The Southern Ocean (SO) is among the regions on Earth that are undergoing regionally the fastest environmental changes. The unique ecological features of its marine life make it particularly vulnerable to the multiple effects of climate change. A network of Marine Protected Areas (MPAs) has started to be implemented in the SO to protect marine ecosystems. However, considering future predictions of the Intergovernmental Panel on Climate Change (IPCC), the relevance of current, static, MPAs may be questioned under future scenarios. In this context, the ecoregionalization approach can prove promising in identifying well‐delimited regions of common species composition and environmental settings. These so‐called ecoregions are expected to show similar biotic responses to environmental changes and can be used to define priority areas for the designation of new MPAs and the update of their current delimitation. In the present work, a benthic ecoregionalization of the entire SO is proposed for the first time based on abiotic environmental parameters and the distribution of echinoid fauna, a diversified and common member of Antarctic benthic ecosystems. A novel two‐step approach was developed combining species distribution modeling with Random Forest and Gaussian Mixture modeling from species probabilities to define current ecoregions and predict future ecoregions under IPCC scenarios RCP 4.5 and 8.5. The ecological representativity of current and proposed MPAs of the SO is discussed with regard to the modeled benthic ecoregions. In all, 12 benthic ecoregions were determined under present conditions, they are representative of major biogeographic patterns already described. Our results show that the most dramatic changes can be expected along the Antarctic Peninsula, in East Antarctica and the sub‐Antarctic islands under both IPCC scenarios. Our results advocate for a dynamic definition of MPAs, they also argue for improving the representativity of Antarctic ecoregions in proposed MPAs and support current proposals of Conservation of Antarctic Marine Living Resources for the creation of Antarctic MPAs.     

Filling biodiversity gaps: benthic hydroids from the Bellingshausen Sea (Antarctica)

The Bellingshausen Sea constitutes the third largest sea in the Southern Ocean, though it is widely recognized as one of the less-studied Antarctic areas. To reduce this lack of knowledge, a survey to study the biodiversity of its marine benthic communities was carried out during the Bentart 2003 and Bentart 2006 Spanish Antarctic expeditions. The study of the hydroid collection has provided 27 species, belonging to ten families and 15 genera. Twenty-one out of the 27 species constitute new records for the Bellingshausen Sea, raising the total number of known species to 37, as also do nine out of the 15 genera. Candelabrum penola, Lafoea annulata, and Staurotheca juncea are recorded for the second time. Most species belong to Leptothecata. Sertulariidae with 13 species (48%) is by far the most speciose family, and Symplectoscyphus with seven species (26%), including S. bellingshauseni sp. nov. and S. hesperides sp. nov., the most diverse genus. Considering the whole benthic hydroid fauna of the Bellingshausen Sea, 18 species (69%) are endemic to Antarctic waters, either with a circum-Antarctic (12 species, 46%) or West Antarctic (6 species, 23%) distribution, 23 (88%) are restricted to Antarctic or Antarctic/sub-Antarctic waters, and only three species have a wider distribution. Bellingshausen Sea hydroid fauna is composed of a relatively high diversity of typical representatives of the Antarctic benthic hydroid fauna, though with a surprisingly low representation of some of the most diverse and widespread Antarctic genera (Oswaldella and Schizotricha), what could be related to the fact that its shelf-inhabiting hydroid fauna remains practically unknown.     

Centers of marine fauna redistribution

The concepts of centers of biota origin and centers of biota accumulation are usually regarded as mutually exclusive. In this paper, they are analyzed within the framework of a unified concept of centers of biota redistribution. Such a center is a biogeographic unit that has three developmental stages: accumulation, diversification, and dispersal. At the accumulation stage, the taxonomic capacity of the corresponding biogeographic district drastically increases and its species richness becomes higher owing to in-migration of species from other regions. At the diversification stage, the species richness continues to increase owing to speciation, and a unified succession system develops. At the dispersal stage, the biotic boundaries of the region act as efficient barriers to species invasion, and the species of the redistribution center gain an advantage over the species of adjacent regions when colonizing new habitats. In the Neogene, the main shallow-water centers of marine fauna redistribution were located in the Indo-Malayan triangle, the western Atlantic, and the northern Pacific. The role of redistribution centers in deep-ocean areas belonged to the Antarctic and the western Pacific. The possibility of using an ecosystem approach to the study of redistribution centers is discussed.     

Diversity and biogeographic patterns of the bryozoan fauna of the Faroe Islands

A total of 228 bryozoan species are recorded within the EEZ of the Faroe Islands, 74 of which are new to the area. Analysis of the distribution of the species among six sectors, each characterized by different environmental conditions, showed three faunal assemblages. Variation of the total Faroese bryozoan fauna and of the bryozoan fauna of most sectors, demonstrated significant negative relationships with depth. In general, analysis of the biogeographic composition showed a strong predominance of boreal over arctic species. However, with respect to faunas of each sector, the Norwegian Basin is characterized by a predominance of arctic species and may be regarded as a part of the Arctic Eurasian sub-region of the Arctic biogeographic region. Comparison of the bryozoan species of each sector with the bryozoan faunas of the other 12 areas in the North Atlantic and the neighbouring Arctic regions showed that only the Faroese shelf fauna has significant similarity with part of them, and thus can be regarded as part of the Scandinavian province of the Norwegian high-boreal sub-region of the Atlantic boreal region. Three sectors, the Faroese–Iceland Ridge, the Faroese–Shetland Channel and Norwegian Basin, belong to a transitional zone between the Atlantic Boreal and the Arctic biogeographic regions. The deep south-western sector forms a separate faunal cluster when compared with both the other sectors within the Faroese area and with the faunas of other large geographic areas, and may be regarded as a separate biogeographic zone of the Boreal Atlantic region due to its high proportion of specific species.     

Detecting glacial refugia in the Southern Ocean

Throughout the Quaternary, the continental-based Antarctic ice sheets expanded and contracted repeatedly. Evidence suggests that during glacial maxima, grounded ice eliminated most benthic (bottom-dwelling) fauna across the Antarctic continental shelf. However, paleontological and molecular evidence indicates most extant Antarctica benthic taxa have persisted in situ throughout the Quaternary. Where and how the Antarctic benthic fauna survived throughout repeated glacial maxima remain mostly hypothesised. If understood, this would provide valuable insights into the ecology and evolution of Southern Ocean biota over geological timescales. Here we synthesised and appraised recent studies and presented an approach to demonstrate how genetic data can be effective in identifying where and how Antarctic benthic fauna survived glacial periods. We first examined the geological and ecological evidence for how glacial periods influenced past species demography in order to provide testable frameworks for future studies. We outlined past ice-free areas from Antarctic ice sheet reconstructions that could serve as glacial refugia and discussed how benthic fauna with pelagic or non-pelagic dispersal strategies moved into and out of glacial refugia. We also reviewed current molecular studies and collated proposed locations of Southern Ocean glacial refugia on the continental shelf around Antarctica, in the deep sea, and around sub-Antarctic islands. Interestingly, the proposed glacial refugia based on molecular data generally do not correspond to the ice-free areas identified by Antarctic ice sheet reconstructions. The potential biases in sampling and in the choice of molecular markers in current literature are discussed, along with the future directions for employing testable frameworks and genomic methods in Southern Ocean molecular studies. Continued data syntheses will elucidate greater understanding of where and how Southern Ocean benthic fauna persisted throughout glacial periods and provide insights into their resilience against climate changes in the future.     

Spatial and temporal variability across life's hierarchies in the terrestrial Antarctic

Antarctica and its surrounding islands lie at one extreme of global variation in diversity. Typically, these regions are characterized as being species poor and having simple food webs. Here, we show that terrestrial systems in the region are nonetheless characterized by substantial spatial and temporal variations at virtually all of the levels of the genealogical and ecological hierarchies which have been thoroughly investigated. Spatial variation at the individual and population levels has been documented in a variety of genetic studies, and in mosses it appears that UV-B radiation might be responsible for within-clump mutagenesis. At the species level, modern molecular methods have revealed considerable endemism of the Antarctic biota, questioning ideas that small organisms are likely to be ubiquitous and the taxa to which they belong species poor. At the biogeographic level, much of the relatively small ice-free area of Antarctica remains unsurveyed making analyses difficult. Nonetheless, it is clear that a major biogeographic discontinuity separates the Antarctic Peninsula and continental Antarctica, here named the 'Gressitt Line'. Across the Southern Ocean islands, patterns are clearer, and energy availability is an important correlate of indigenous and exotic species richness, while human visitor numbers explain much of the variation in the latter too. Temporal variation at the individual level has much to do with phenotypic plasticity, and considerable life-history and physiological plasticity seems to be a characteristic of Antarctic terrestrial species. Environmental unpredictability is an important driver of this trait and has significantly influenced life histories across the region and probably throughout much of the temperate Southern Hemisphere. Rapid climate change-related alterations in the range and abundance of several Antarctic and sub-Antarctic populations have taken place over the past several decades. In many sub-Antarctic locations, these have been exacerbated by direct and indirect effects of invasive alien species. Interactions between climate change and invasion seem set to become one of the most significant conservation problems in the Antarctic. We conclude that despite the substantial body of work on the terrestrial biodiversity of the Antarctic, investigations of interactions between hierarchical levels remain scarce. Moreover, little of the available information is being integrated into terrestrial conservation planning, which lags far behind in this region by comparison with most others.     

Benthic hydroids (Cnidaria: Hydrozoa) from Peter I Island (Southern Ocean, Antarctica)

Twenty-three species of benthic hydroids, belonging to eight families and 13 genera, were found in a hydroid collection from Peter I Island, collected during both the Bentart 2003 and Bentart 2006 Spanish expeditions with BIO Hespérides in 2003 and 2006. Fourteen out of the 23 species constitute new records for Peter I Island, raising the total number of known species in the area to 30, as also do seven out of the 13 genera. The majority of the species are members of the subclass Leptothecata; the subclass Anthoathecata being scarcely represented. Sertulariidae is the family with the greatest number of species in the collection, with eight species (35%), followed by Lafoeidae with five (22%). Symplectoscyphus with four species (17%) and both Antarctoscyphus and Halecium with three (13%), including H. frigidum sp. nov., were the most diverse genera. Twenty species (ca. 77%) are endemic to Antarctic waters, either with a circum-Antarctic (11 species, ca. 42%) or West Antarctic (9 species, ca. 35%) distribution. Twenty-four (ca. 92%) are restricted to Antarctic or Antarctic/sub-Antarctic waters; only two species have a wider distribution. Peter I Island hydroid fauna is composed of typical representatives of the Antarctic benthic hydroid fauna, though it is characterized by the low representation of some of the most diverse and widespread Antarctic genera (Schizotricha and Staurotheca).     

Biogeographical and phylogeographical relationships of the bathyal ophiuroid fauna of the Macquarie Ridge, Southern Ocean

There are relatively few studies examining the latitudinal distribution of polar, subantarctic and temperate faunas on the bathyal seafloor across the Southern Ocean. Here, we investigate the relationship between the subantarctic Macquarie Ridge and adjacent regions of Antarctica (including the Ross Sea) and temperate Australia and New Zealand at depths of 200–2,500 m. We study the fauna at two levels of classification (1) morpho-species (MSPs) accepted by taxonomists and (2) evolutionary significant units defined as reciprocally monophyletic clades derived from phylogenies of mitochondrial DNA. The ophiuroid fauna on the Macquarie Ridge has a predominantly temperate origin, with far more MSPs shared with south-eastern Australia (78 % of species) and southern New Zealand (83 %) than neighbouring Antarctic regions (33 %). However, this asymmetry also reflects the relative species richness of these regions. Many species that are shared between Antarctica and the Macquarie Ridge have diverged into distinct mtDNA lineages indicative of a recent barrier to gene flow.     

Open-ocean barriers to dispersal: a test case with the Antarctic Polar Front and the ribbon worm Parborlasia corrugatus (Nemertea: Lineidae)

Open-ocean environments provide few obvious barriers to the dispersal of marine organisms. Major currents and/or environmental gradients potentially impede gene flow. One system hypothesized to form an open-ocean dispersal barrier is the Antarctic Polar Front, an area characterized by marked temperature change, deep water, and the high-flow Antarctic Circumpolar current. Despite these potential isolating factors, several invertebrate species occur in both regions, including the broadcast-spawning nemertean worm Parborlasia corrugatus. To empirically test for the presence of an open-ocean dispersal barrier, we sampled P. corrugatus and other nemerteans from southern South America, Antarctica, and the sub-Antarctic islands. Diversity was assessed by analyzing mitochondrial 16S rRNA and cytochrome c oxidase subunit I sequence data with Bayesian inference and tcs haplotype network analysis. Appropriate neutrality tests were also employed. Although our results indicate a single well-mixed lineage in Antarctica and the sub-Antarctic, no evidence for recent gene flow was detected between this population and South American P. corrugatus. Thus, even though P. corrugatus can disperse over large geographical distances, physical oceanographic barriers (i.e. Antarctic Polar Front and Antarctic Circumpolar Current) between continents have likely restricted dispersal over evolutionary time. Genetic distances and haplotype network analysis between South American and Antarctic/sub-Antarctic P. corrugatus suggest that these two populations are possibly two cryptic species.     

Shallow-water benthic hydroids from Tethys Bay (Terra Nova Bay, Ross Sea, Antarctica)

The shallow-water hydrozoan Antarctic fauna is still poorly studied, and available knowledge mostly refers to samples gathered by traditional ship-operated gears. By scuba diving in the coastal areas off the Italian Antarctic station “Mario Zucchelli” (Ross Sea, Terra Nova Bay), in the austral summer 2002–2003, a total of 20 hydrozoan species were found, belonging to 10 families and 13 genera. As hypothesized, Anthoathecata (11 species), usually under-represented in collections from indirect sampling gears, are common as also are Leptothecata (9 species). Hydractiniidae and Hydractinia are the dominant family and genus, followed by Haleciidae and Halecium. A new species to science, Halecium exaggeratum sp. nov. is also described. Most species are either endemic to Antarctic waters or restricted to Antarctic/sub-Antarctic areas; only two species have a wider distribution. Material reared in aquaria at the Italian Antarctic Base Mario Zucchelli facilitated knowledge of the life cycle and reproductive biology of several species. In particular, Opercularella belgicae was found to liberate a medusa stage referable to Phialella, and the species is assigned here to that genus, as Phialella belgicae. Also, extraordinary is the complete absence or scant representation of the most typical Antarctic benthic hydroid genera (Antarctoscyphus, Oswaldella, Schizotricha, Staurotheca, and Symplectoscyphus), likely related to the shallow limits of sampling (down to 48 m).     

Benthic hydroids (Cnidaria,Hydrozoa) from the Balleny Islands (Antarctica)

Twenty-two species of benthic hydroids, belonging to ten families and 14 genera, were found in a hydroid collection obtained in the Balleny Islands during the BioRoss expedition with the NIWA research vessel Tangaroa in 2004. Twenty of those species constitute new records for the Balleny Islands, raising the total number of known species in the area to 25. Most are members of the subclass Leptothecata, although the subclass Anthoathecata is also relatively well represented. Kirchenpaueriidae and Sertulariidae constitute families with the greatest numbers of species in the collection, with five species (20%) each. Oswaldella with five species (20%) and Staurotheca with four (16%), were the most diverse genera. Twelve species (63%) are endemic to Antarctic waters, most of them with a circum-Antarctic distribution, and 17 (89%) are restricted to Antarctic or Antarctic/sub-Antarctic waters. Although the Balleny Islands hydroid fauna seems to be a typical Antarctic assemblage, it has some striking peculiarities, namely the absence or low representation of some typical and widespread Antarctic genera (Antarctoscyphus and Schizotricha/Symplectoscyphus, respectively).     

The composition,structure, and distribution of benthic ostracod fauna (Ostracoda,Myodocopa) in Antarctic waters

The benthic fauna of ostracods of the order Myodocopida of Antarctic waters is characterized by high diversity, relative species abundance, and a complicated taxonomic and ecological structure, with a simplified biogeographical structure. This fauna, which is distinguished by a high level of endemicity, although at a low taxonomic rank, includes a great share of deep-sea and subtidal elements. Ostracod populations of High and Low-Antarctic subzones differ qualitatively and quantitatively. A distinct impoverishment of fauna is observed in the region of the Antarctic divergence compared to the more northern areas. The number of species increases with depth to reach its maximum in the lower subtidal zone and on the upper continental slope at depths of 200–500 m. The number of species decreases with increasing depth. Myodocopida have not been yet found in the Antarctic waters deeper than 5000 m.     

Using directed phylogenetic networks to retrace species dispersal history

Methods designed for inferring phylogenetic trees have been widely applied to reconstruct biogeographic history. Because traditional phylogenetic methods used in biogeographic reconstruction are based on trees rather than networks, they follow the strict assumption in which dispersal among geographical units have occurred on the basis of single dispersal routes across regions and are, therefore, incapable of modelling multiple alternative dispersal scenarios. The goal of this study is to describe a new method that allows for retracing species dispersal by means of directed phylogenetic networks obtained using a horizontal gene transfer (HGT) detection method as well as to draw parallels between the processes of HGT and biogeographic reconstruction. In our case study, we reconstructed the biogeographic history of the postglacial dispersal of freshwater fishes in the Ontario province of Canada. This case study demonstrated the utility and robustness of the new method, indicating that the most important events were south-to-north dispersal patterns, as one would expect, with secondary faunal interchange among regions. Finally, we showed how our method can be used to explore additional questions regarding the commonalities in dispersal history patterns and phylogenetic similarities among species.     

The lichen genus Stereocaulon in Antarctica and South Georgia

Summary Six species of Stereocaulon and one unnamed taxon (close to S. glabrum) are reported from South Georgia, the maritime Antarctic islands and Antarctic Peninsula. S. caespitosum is new to the western sub-Antarctic. Variations in morphology and secondary chemistry are provided, and the ecology and geographical distribution in the sub-Antarctic and Antarctic biomes are given for each taxon.