The role of colonization in establishing patterns of community composition and diversity in shallow-water sedimentary communities

To determine whether pattern and diversity in benthic sedimentary communities are set primarily at colonization or by post-settlement biological interactions, we collected faunal cores and conducted reciprocal sediment transplant experiments at a sandy and a muddy site at 12 m depth, ~3 km apart off New Jersey. Multivariate analyses of cores collected at these sites in September 1994 indicated differences in the taxa determining local pattern, with the bivalve Spisula solidissima and the polychaete Polygordius sp. being dominant at the sandy site, and oligochaetes, several polychaete species and the bivalve Nucula annulata dominant at the muddy site. Individual cores from the sandy site were significantly less diverse than those at the muddy site. Short-term experiments (3-5 d) were deployed by divers at three different times (August-September, 1994). Replicate trays (100 cm2) filled with azoic sand or mud were placed flush with the ambient seafloor at both sites. Multivariate comparisons indicated that sediment treatment in trays played a greater role in determining colonization patterns in the first experiment, site played a greater role in the second, and both variables contributed in the third. This pattern suggests that larval settlement and habitat choice played an important role in the first and third experiments, and that local transport of recently settled juveniles from the surrounding sediments was important in the second and third experiments. Sandy-site trays had significantly lower diversity than muddy-site trays, but there was no effect of sediment type in trays on diversity of colonizers. These experiments focused on small spatial scales and three short time periods, but they demonstrate that species patterns in some environments may be set by habitat selection by larvae and by juvenile colonization from the surrounding community. Post-colonization processes such as predation and competition likely play a major role for some species, but patterns of initial colonization corresponded well with those in the local community

Macrofaunal response to artificial enrichments and depressions in a deep-sea habitat

To test whether colonizing macrofauna specialize on different types of small-scale patches of food and disturbance in the deep sea, sediment tray and artificial depression colonization experiments were conducted on the deep-sea floor at 900-m depth, south of St. Croix, U.S. Virgin Islands. Trays and depressions were unenriched (Unenriched Controls) or enriched with either Thalassiosira sp. or Sargassum sp. Concurrent deployment of different types of enrichment and disturbance made it possible to evaluate whether macrofauna specialize on different patches, and thus avoid species interactions that might lead to competitive exclusion. Depressions create a hydrodynamic regime that traps passive particles, allowing tests of the relative importance of active selection of different patch types versus passive deposition for abundant colonizers. After 23 d, total densities and densities of the four abundant colonizers (Capitella spp., Nereimyra punctata, Cumella sp. and Nebalia sp.) were extremely high in enriched trays, despite relatively low ambient densities. Densities in Unenriched Control Trays were very low, and did not attain ambient densities. After 24 d, total densities in all depression treatments were considerably lower than in enriched tray treatments, and only Sargassum Depression densities exceeded those in the ambient environment. Lower densities of organisms in depression treatments compared with trays and differences in densities among depression treatments suggest that the dominant colonizers were highly active and selective, and were not passively entrained in depressions. Faunal analysis indicated that trays and depressions were very different, and Sargassum Depression fauna was very different from other depression types. A strong difference was not observed between fauna in ambient sediments and Thalassiosira sp. or Unenriched Control Depressions, perhaps because Thalassiosira sp. was dropped in depressions on the sediment surface and may have been more readily available to consumers and more rapidly consumed than in trays. Thalassiosira Trays were colonized by a lower diversity fauna than Sargassum Trays, and Unenriched Control Trays were colonized by very low densities of a fauna that was comparable in diversity to the ambient community. Diversity in Sargassum Depressions was higher than in enriched trays but lower than in other artificial depressions and the ambient fauna. Thalassiosira Depressions and Unenriched Control Depressions were comparable in diversity to ambient fauna and natural depressions, which were highly diverse. These experiments suggest that fauna may respond quickly and selectively to artificial food patches and disturbance, and this fauna is different from that observed in the ambient sediment. Thus, a patch mosaic may be part of the reason for the high species diversity that is observed in deep-sea ecosystems. The different, highly diverse, fauna observed in natural depressions compared with flat ambient sediment suggests that natural analogs of these experiments have unique faunas that may contribute to the species richness of deep-sea habitats

Temporal and spatial variation in infaunal community structure in physically active continental shelf sediments at a long-term ecosystem observatory (LEO-15) off New Jersey, USA

When the LEO-15 (Long-term Ecosystem Observatory at 15-m depth) research area was designated around Beach Haven Ridge (39°27.69′ N, 74°15.81′ W) in the early 1990\u27s several stations were selected to reflect the different physical regimes and habitats present in a highly dynamic shelf environment. This study determined the composition, diversity, and abundance of infaunal communities at three contrasting stations on five dates in 1994–1995. A unique sampling scheme was used where a nested design was combined with targeted sampling by divers of benthic topographic habitat features, namely crests and troughs of sand ripples. The total number of taxa collected was 148. The majority of taxa (80%) were either ubiquitous (67 taxa, 45%) or had an apparent affinity for a single station (52 taxa, 35%). The polychaete Polygordius jouinae was numerically dominant. The three distinctive stations 2–4 km apart, the date of sampling and, somewhat surprisingly, whether or not samples were taken in adjacent crests or troughs, all contributed to the structure of LEO-15 infaunal communities. Together these three stations characterize the sediments and fauna in the LEO-15 research area and form the basis for future studies of short and longer-term changes, and of the processes and mechanisms responsible for the patterns observed

Deep-Sea Nematodes Actively Colonise Sediments, Irrespective of the Presence of a Pulse of Organic Matter: Results from an In-Situ Experiment

A colonisation experiment was performed in situ at 2500 m water depth at the Arctic deep-sea long-term observatory HAUSGARTEN to determine the response of deep-sea nematodes to disturbed, newly available patches, enriched with organic matter. Cylindrical tubes,laterally covered with a 500 µm mesh, were filled with azoic deep-sea sediment and 13C-labelled food sources (diatoms and bacteria). After 10 days of incubation the tubes were analysed for nematode response in terms of colonisation and uptake. Nematodes actively colonised the tubes,however with densities that only accounted for a maximum of 2.13% (51 ind.10 cm−2) of the ambient nematode assemblages. Densities did not differ according to the presence or absence of organic matter, nor according to the type of organic matter added. The fact that the organic matter did not function as an attractant to nematodes was confirmed by the absence of notable 13C assimilation by the colonising nematodes. Overall, colonisationappears to be a process that yields reproducible abundance and diversity patterns, with certain taxa showing more efficiency. Together with the high variability between the colonising nematode assemblages, this lends experimental support to the existence of a spatio-temporal mosaic that emerges from highly localised, partially stochastic community dynamics

Do Larval Supply and Recruitment Vary among Chemosynthetic Environments of the Deep Sea?

BACKGROUND: The biological communities that inhabit chemosynthetic environments exist in an ephemeral and patchily distributed habitat with unique physicochemical properties that lead to high endemicity. Consequently, the maintenance and recovery from perturbation of the populations in these habitats is, arguably, mainly regulated by larval supply and recruitment. METHODOLOGY/PRINCIPAL FINDINGS: WE USE DATA FROM THE PUBLISHED SCIENTIFIC LITERATURE TO: (1) compare the magnitudes of and variability in larval supply and settlement and recruitment at hydrothermal vents, seeps, and whale, wood and kelp falls; (2) explore factors that affect these life history processes, when information is available; and (3) explore taxonomic affinities in the recruit assemblages of the different chemosynthetic habitats, using multivariate statistical techniques. Larval supply at vents can vary across segments by several orders of magnitude for gastropods; for bivalves, supply is similar at vents on different segments, and at cold seeps. The limited information on larval development suggests that dispersal potential may be highest for molluscs from cold seeps, intermediate for siboglinids at vents and lowest for the whale-bone siboglinid Osedax. Settlement is poorly studied and only at vents and seeps, but tends to be highest near an active source of emanating fluid in both habitats. Rate of recruitment at vents is more variable among studies within a segment than among segments. Across different chemosynthetic habitats, recruitment rate of bivalves is much more variable than that of gastropods and polychaetes. Total recruitment rate ranges only between 0.1 and 1 ind dm(-2) d(-1) across all chemosynthetic habitats, falling above rates in the non-reducing deep sea. The recruit assemblages at vents, seeps and kelp falls have lower taxonomic breadth, and include more families and genera that have many species more closely related to each other than those at whale and wood falls. Vents also have the most uneven taxonomic structure, with fewer recruits represented by higher taxonomic levels (phyla, orders, classes) compared to seeps and wood and kelp falls, whereas the opposite is true at whale falls. CONCLUSIONS/SIGNIFICANCE: Based on our evaluation of the literature, the patterns and regulatory factors of the early history processes in chemosynthetic environments in the deep sea remain poorly understood. More research focused on these early life history stages will allow us to make inferences about the ecological and biogeographic linkages among the reducing habitats in the deep sea

How Many Species Are There on Earth and in the Ocean?

The diversity of life is one of the most striking aspects of our planet; hence knowing how many species inhabit Earth is among the most fundamental questions in science. Yet the answer to this question remains enigmatic, as efforts to sample the world's biodiversity to date have been limited and thus have precluded direct quantification of global species richness, and because indirect estimates rely on assumptions that have proven highly controversial. Here we show that the higher taxonomic classification of species (i.e., the assignment of species to phylum, class, order, family, and genus) follows a consistent and predictable pattern from which the total number of species in a taxonomic group can be estimated. This approach was validated against well-known taxa, and when applied to all domains of life, it predicts ∼8.7 million (±1.3 million SE) eukaryotic species globally, of which ∼2.2 million (±0.18 million SE) are marine. In spite of 250 years of taxonomic classification and over 1.2 million species already catalogued in a central database, our results suggest that some 86% of existing species on Earth and 91% of species in the ocean still await description. Renewed interest in further exploration and taxonomy is required if this significant gap in our knowledge of life on Earth is to be closed

Semi-Automated Image Analysis for the Assessment of Megafaunal Densities at the Arctic Deep-Sea Observatory HAUSGARTEN

Megafauna play an important role in benthic ecosystem function and are sensitive indicators of environmental change. Non-invasive monitoring of benthic communities can be accomplished by seafloor imaging. However, manual quantification of megafauna in images is labor-intensive and therefore, this organism size class is often neglected in ecosystem studies. Automated image analysis has been proposed as a possible approach to such analysis, but the heterogeneity of megafaunal communities poses a non-trivial challenge for such automated techniques. Here, the potential of a generalized object detection architecture, referred to as iSIS (intelligent Screening of underwater Image Sequences), for the quantification of a heterogenous group of megafauna taxa is investigated. The iSIS system is tuned for a particular image sequence (i.e. a transect) using a small subset of the images, in which megafauna taxa positions were previously marked by an expert. To investigate the potential of iSIS and compare its results with those obtained from human experts, a group of eight different taxa from one camera transect of seafloor images taken at the Arctic deep-sea observatory HAUSGARTEN is used. The results show that inter- and intra-observer agreements of human experts exhibit considerable variation between the species, with a similar degree of variation apparent in the automatically derived results obtained by iSIS. Whilst some taxa (e. g. Bathycrinus stalks, Kolga hyalina, small white sea anemone) were well detected by iSIS (i. e. overall Sensitivity: 87%, overall Positive Predictive Value: 67%), some taxa such as the small sea cucumber Elpidia heckeri remain challenging, for both human observers and iSIS

Semi-Automated Image Analysis for the Assessment of Megafaunal Densities at the Arctic Deep-Sea Observatory HAUSGARTEN

Megafauna play an important role in benthic ecosystem function and are sensitive indicators of environmental change. Non-invasive monitoring of benthic communities can be accomplished by seafloor imaging. However, manual quantification of megafauna in images is labor-intensive and therefore, this organism size class is often neglected in ecosystem studies. Automated image analysis has been proposed as a possible approach to such analysis, but the heterogeneity of megafaunal communities poses a non-trivial challenge for such automated techniques. Here, the potential of a generalized object detection architecture, referred to as iSIS (intelligent Screening of underwater Image Sequences), for the quantification of a heterogenous group of megafauna taxa is investigated. The iSIS system is tuned for a particular image sequence (i.e. a transect) using a small subset of the images, in which megafauna taxa positions were previously marked by an expert. To investigate the potential of iSIS and compare its results with those obtained from human experts, a group of eight different taxa from one camera transect of seafloor images taken at the Arctic deep-sea observatory HAUSGARTEN is used. The results show that inter- and intra-observer agreements of human experts exhibit considerable variation between the species, with a similar degree of variation apparent in the automatically derived results obtained by iSIS. Whilst some taxa (e. g. Bathycrinus stalks, Kolga hyalina, small white sea anemone) were well detected by iSIS (i. e. overall Sensitivity: 87%, overall Positive Predictive Value: 67%), some taxa such as the small sea cucumber Elpidia heckeri remain challenging, for both human observers and iSIS