IN REVIEW

Description

State of knowledge 

Autecology

Many (but not all) Antarctic benthic species have characteristics of k-strategists i.e. slow growth, long maturation, long life expectancy, large body size, larger eggs, brooding etc. in comparison to similar temperate or tropical species. These characteristics may be an adaption to living in cold or under a highly seasonal food supply (Clarke, 1979). 

A high proportion of brooding species has been repeatedly recorded within the Antarctic benthos, 50-70% depending on phyla (Perl and Poulin 2010). This trait is particularly dominant in echinoids (Poulin and Feral 1996, Poulin et al. 2002) see Box 1 from Poulin et al. 2002 and crustaceans (Brandt, 2000; Brandt et al. 2007a, 2007b). A review of the evolutionary and environmental factors that may have resulted in different reproductive traits within Antarctic benthic invertebrates are discussed in Pearse et al. (2008). In contrast some of the most successful shallow water species with planktotrophic species includes the sea urchin Sterechinus neumayeri (Bosch et al. 1987, Brey and Gutt 1991), the seastars Odontaster validus, O. meridionalis and Porania antarctica (Pearse, 1994), the ophiuroid Ophionotus victoriae (Pearse, 1994), the Antarctic scallop Adamussium colbecki (Chiantore et al. 2001), and the clam Laternula elliptica (Bosch and Pearse 1988).

The reproduction of shallow water benthic invertebrates has been studied for a select number of species, these are generally collected within the vicinity of research stations. These species include: the brachiopod Liothyrella uva (Meidlinger et al. 1998), the scallop Adamussium colbecki (Chiantore et al. 2002, Tyler et al. 2003), the bivalves Adacnarca nitens (Higgs et al. 2009) and Aequiyoldia eightsiith (Lau et al. 2018), the crabs Lithodes santolla and Paralomis granulosa (Calcagno et al. 2004), the tunicate Cnemidocarpa verrucosa (Sahade et al. 2004, Strathmann et al. 2006), the sea urchin Sterechinus neumayeri (Brockington et al. 2006, Bosch et al. 1987), the crinoid Promachorcinus kerguelensis (McClintock and Pearse, 1987), the seastars Odontaster validus (Grange et al. 2007, Pearse 1965, Stanwell-Smith and Clarke 1998, Pearse and Bosch, 1986), Lophaster gaini, Psilaster charcoti (McClintock, 1989), Porania antarctica, Diplasterias brucei, and Acodontaster conspicuus (Bosch and Pearse, 1990), and the brittle star Ophionotus victoriae (Grange et al. 2004).  Deep sea species studied include the 3 Yoldiella species (Reed et al. 2014).

There is evidence for interannual variation in the reproductive output of benthic species that reflects interannual changes in primary production, this is evident in species whose have seasonal spawning events and such as Odontaster validus and Ophionotus victoriae (Grange et al. 2011, Brasier unpublished data). 

Life history

Mobility

Within the antarctic benthos most species are either sessile or slow moving however some species are motile. The degree of mobility is important for connectivity between populations across local, regional and ocean scales but also for the exposure or avoidance of physical disturbance e.g. ice-seabed interactions and predator avoidance.

The classification of mobility can vary across studies but is important for trait based analysis and assessing the effect of some impact drivers. In Smale (2008a) Low mobility = Taxa are completely or largely immobile. Includes sedentary worms, some bivalves and all sessile taxa and high mobility = Includes mobile worms, all crustaceans, gastropod and some bivalve molluscs. As suggested in Smale (2008a), mobility traits may correlate with disturbance "at one site, the relative abundance of the low mobility group was significantly greater at low disturbance levels, whilst the relative abundance of the high dispersal group (taxa with pelagic larvae) was elevated at high disturbance levels. At the other site, the relative abundance of secondary consumers was greater at high disturbance levels." 

Connectivity between sites and regions may be maintained by larval transport, especially if larvae are pelagic over a long duration (Brasier et al. 2017). However, in highly retentive coastal regions (e.g. fjords) connectivity can still be limited over small spatial scales (100 km) even for larvae with  a long pelagic phase. In these regions, self-recruitment is important for maintaining local populations especially for low dispersal potential organisms with short pelagic phases (e.g. tunicates) and these populations could be more sensitive to future stochastic events (Ziegler et al. in prep).

Diet (foraging and consumption)

Various feeding strategies are used by benthic species here are the main types: 

Filter feeders and Suspension feeders

Many Antarctic benthic communities are dominated by filter and suspension feeders especially in coastal and shelf locations these include sessile sponges, tunicates, cnidarians, bryozoans and some echinoderms (Dayton et al. 1986). A key difference between filter and suspension feeders is their selective abilities. Filter feeders such as sponges filter seawater to obtain particles whereas suspension feeders feed by extending specialised feeding structures into bottom currents to collect organic particles (including detrital material, living plankton, larvae and eggs) from the surrounding water. Therefore, suspension feeders are more susceptible to changes in turbidity, for example, from glacial runoff. Given the seasonal supply of food seasonal variation in suspension feeding activity has been observed, animals can cease feeding for a short period during winter (Barnes and Clarke, 1995, Gili et al. 2001). 

Deposit feeders 

Deposit feeders consume material from the surface or subsurface of the seafloor. Many deposit feeders are motile using specialised body parts, e.g. palps or tentacles, to collect and ingest organic material. Detritus from the summer phytoplankton blooms is the primary source of organic material for most of the Antarctic detritivorous benthos however, biochemical analysis found little evidence of seasonal signals within benthic species suggesting that long-term variability in production processes in the water column are integrated in benthic habitats (Mincks et al. 2008). Nonetheless, during periods of high phytodetrital accumulation at the seafloor, deposit feeders can consume organic matter at rates comparable to shallow, warmer water deposit feeders and affect rates of organic carbon burial and cycling (Ziegler et al. in prep). In this way, pelagic-benthic coupling is seasonally decoupled in some regions such as the WAP. 

Deposit feeding species including many holothurians species, including the surface deposit feeders Peniagone vignoni, Protelpidia murrayi and Pseudostichopus mollis and subsurface deposit feeders Molpadia musculus and Ypsilocucumis turricata, as well as polychaetes, including many surface feeding spinoidae and subsurface feeding scalibregmatidae (Mincks et al. 2008, Brasier et al. in prep).  In coastal fjords (e.g. Andvord Bay), the deposit-feeding community can be dominated by smaller deposit feeders such as Amythas membranifera, an ampharetid polychaete (Grange and Smith, 2013).

Omnivores

Omnivores have a broad diet and can switch between feeding on organic material derived from primary producers to scavenging on other animals. Omnivory can be opportunistic or temporal omnivores depending on conditions and may be more common in food limited environments such as the deep-sea. Omnivorous species include some of the most abundant echinoderms including the brittle star Ophionotus victoriae, the echinoid Sterechinus neumayeri, the seastar Odontaster validus (McClintock, 1994 and references there in) as well as many polychaetes and crustaceans. 

Predators

Predators prey on other animals by active hunting or luring individuals, they often have specialised sensory organs or jaws. Predators can have a signficant structuring role in marine communities, in the Antarctic "modern" predators such as fast moving, skeleton-crushing (durophagos) fish, sharks and crabs are rare or absent and asteroids and nemerteans are generally the top predators (Aronson et al. 2007 and references there in). 

Scavenger

Scavengers feed on carrion or dead material that they do not kill themselves thus scavengers are sometimes referred to as necrophages. Necorphages are opportunistic feeders exploiting energy rich food falls (e.g. marine mammals) from surface waters or dead benthic animals. This trait may be widespread in the Antarctic benthos among amphipods, gastropods, ophiuroids, echinoids and nemerteans (Presler, 1986). Baited experiments have shown variation in levels of necrophagy between Antarctic sites and within species across seasons (Smale et al. 2007a). 

A note on benthic dietary analyses

Historically the trophic traits and dietary information of benthic marine species has been assigned based on morphological traits i.e. if the species has large jaws it is likely to be a predator/scavenger, or where possible gut content analysis and observations. More recently there has been an increase in the number of species level investigations using biochemical methods such as fatty acid biomarkers and stable isotopes to determine the trophic level and trophic traits of different species. 

For example Würzberg et al. (2011a-c) has used fatty acid biomarkers to examine the dietery components of polychaetes, demersal fish and peracarid crustaceans in the deep-sea and Antarctic shelf. These studies highlight selective feeding between higher taxonomic classifications and within the same family at different depths for polychaetes and ontogenetic changes in demersal fish.  

Stable isotope analysis has also been used to estimate trophic position of Antarctic benthic species e.g. Nyssen et al. (2005) and Norkko et al. (2007). A limitation of many stable isotope studies or the use of collection material for analysis is obtaining a signature or value for the "source" material such as filtered seawater or phytodetrius. A way of overcoming this problem is using methods like compound specific stable isotope analysis where a source signature is contained within each individual. This method has been trialed in preserved polychaetes from the West Antarctic Peninsula (Brasier et al. in prep) and has the potential to obtain information about the trophic level of collection material. 

At present the SO-Diet database can not filter for benthic species. 

Energetics

Habitat

Benthic habitats can vary in many ways and can be "classified" by geomorphic features that can relate to topographic, sedimentary and oceanographic conditions such as substrate, elevation and current regime (e.g. in Post et al. 2014). The table below shows how depth and sediment can be used to describe benthic habitats and influence the faunal communities found there. 

Benthic habitats can also vary by the sediment type, this can impact the fauna that inhabits them. Benthic habitats are "patchy" in that a flat sandy area may have infrequent drop stones or rocks that give rise to different fauna, a few examples are shown in the table below from Brasier et al. (2018): 

For some regions of the continental shelf, geomorphic features can be used to map the distribution benthic biota and communities e.g. Barry et al. 2003, Beaman and Harris 2005, Thrush et al. 2006, Gutt 2007, Post et al. 2011. However, benthic environments are patchy at all scales measured to date and fine scale variation can override any larger scale geomorphic community patterns, for example, the presence of soft or hard substrate across different geomorphological areas (Post et al. 2010, Brasier et al. 2018).

Relationships, thresholds and limits

Species within the Antarctic benthos are generally considered to be stenothermal having evolved in a relatively stable cold environment (Peck et al. 2004) this may make then vulnerable to increasing temperatures and ocean acidificiation especially the developmental stage of calcifying organisms. Juveniles are also considered to be vulnerable however when warmed at 1°C 3 days⁻¹, adults of Laternula elliptica, Cucumaria georgiana, Sterechinus neumayeri, and Odontaster validus juveniles survived to higher temperatures than adults in all species studied (Peck et al. 2013). 

Functional responses of taxon (y-axis) to habitat variables (x-axis) are described here (with citations to the evidence). Parameters for a response type are to be given with their attendant uncertainties/errors/range, with references.

As temperature changes on the Antarctic shelf species may "lose" or "gain" in potential available habitat (area within their temperature tolerance). This has been investigated using occurrence records and projected climate models (RCP8.5) in Griffiths et al.(2017). Figure shows the percentage of 963 shelf-dwelling species that are expected to experience a change in suitable available habitat. 

Population

Measuring the abundance or density of benthic fauna is incredibly difficult especially over large spatial scales. Most sampling methods are qualitative or quantitative and the patchiness of benthic environments can lead to high uncertainty when up-scaling. In general species abundance is similar to that of temperate and tropical regions and diversity is generally high (Clarke, 1996). 

Range and Structure

The Antarctic benthos is a complex ecosystem and the benthic communities exhibit high patchiness at all spatial scales. This patchiness is a result of interacting biological and physical factors including ice scour, species-specific habitat requirements (e.g. attachment for sessile species) and other biological characteristics of single species such as predominance of vegetative reproduction (e.g. sponges and soft corals), slow dispersion and slow growth (Barnes et al. 2007 and refs there in). Fine scale patchiness can prevent the detection of large scale ecological patterns within the benthos (e.g. Brasier et al. 2018). 

Changes in species abundance and diversity with depth have been recorded. For example Neal et al. (2018) found depth to be the main structuring factor of polychaete diversity in the Atlantic and East Pacific sectors of the Southern Ocean between 500 and 3500 m, similar results were found for Isopoda by Brandt et al. (2004). Additionally, some cryptic species have been found to be isolated by depth (Schüller, 2011). 

Dynamics

In general the Antarctic benthos is considered to be relatively stable an exceptional to this are shallow water communities at depths <10 m which may be impacted by ice more than once a year (Brown et al. 2004). The frequency of ice disturbance decreases with depth (Smale et al. 2007b) and below 30 m less likely to be structured by ice disturbance but other physical and biological factors. Including, for biological, primary production input, competition and predation, and physical, substratum, contamination and current flow (Smale 2008b and refs therein). However, ice scour can impact the deep benthos up to 550 m from grounding of tabular icebergs in coastal regions (Barnes and Conlan, 2007 and refs therein).

Responses to climate change have already been detected in some benthic species on the West Antarctic Peninsula, community changes including a sudden shift from filter feeding-ascidian dominated communities to a mixed assemblage at King George Island might be associated with sediment run-off triggered by glacier retreat (Sahade et al. 2015). 

In the Ross Sea where the phytoplankton bloom period is lengthening, driven by winds that have expanded the extent of polynyas, has resulted in the growth of phytoplankton-consuming benthos communities (Barnes et al. 2011). 

In McMurdo sound, the growth rates and recruitment of suspension feeding sponges have dramatically increased in response to a shift in the pelagic community to a dominance by bacterioplankton which the sponges are particularly effective at exploiting (Dayton et al. 2019).

Species movements and range shifts are predicted under changing water temperatures at the seafloor. These changes could remove physiological barriers to some species and facilitate their movement/re-invasion into the shallow Antarctic benthos including shell-crushing predatory crabs which may have significant impacts on the benthic communities (Aronson et al. 2007).

Synecology

Modern predators (e.g. shell crushing species such as crabs, fish, sharks and rays) are rare or absent from the Antarctic benthos. Among Antarctic marine mammals Weddell seals will occasionally take benthic prey but primarily feed on fish and squid. The key predators to benthic species are slow moving invertebrates within the benthos including asteroids and nemerteans (Aronson et al. 2007). 

Along the West Antarctic Peninsula demersal fish are major consumers of benthos at inshore habitats. At offshore locations demersal fish are less dependent on benthos where they feed on more nutritional prey such as krill and nekton. Benthic feeders include Gobionotothen gibberifrons and Notothenia rossii juveniles (Barrera-O 2002). Gut content and fatty acid analyses by Würzberg et al. (2011c) found amphipods, polychaetes, gastropods and other crustaceans were the main dietary components of demersal fish from the Nototheniidae, Macrouridae, Channichtyidae, Bathydraconidae and Artedidraconidae families. 

Competitors

Benthic species can be in direct competition for space and food. 

Analysis of bryozoa from Alaska to Antarctica has suggested that competition within the benthos is more hierarchical towards the poles (Barnes, 2002).

In the Antarctic benthos in the absence of disturbance (ice scour), the combination of slow, temporally restricted growth with consistent annual recruitment of larvae (characteristic of Antarctic fauna) may result in successional processes during early colonisation being largely deterministic, with the occupation of free space being achieved primarily by high density of recruitment rather than by competitive interaction or differential growth rates (Bowden et al. 2006). 

Sub-lethal mortality (e.g. some zooids are killed but the colony survives) by competitive interaction is common along the Antarctic Peninsula in coastal rock communities (Barnes and Arnold, 2001). 

However species may have evolved to exist symmetrically by reducing competition. A recent study of octopods off South Georgia shows how functionally similar co-existing species (Adelieledone polymorpha and Pareledone turqueti) can have different trophic niches that may reduce competition. Such differences may be of benefit under current changing conditions in this area (Matias et al. 2019).  

Cryo-pelagic-benthic coupling

The degree of coupling between the cryosphere, pelagic process and benthic communities is an area of current and future research. Understanding how changes in the cryopshere and/or the pelagic system might effect the benthos could identify drivers of change. 

On the West Antarctic Peninsula, Smith et al. (2006) found poor coupling between the summer phytoplankton bloom and benthic processes on the shelf, with limited seasonality in the trophic signatures of benthic fauna. Also along the West Antarctic Peninsula weak hints of cryo-pelagic-benthic coupling were recorded in Gutt et al. (2019) which may indicate the complexity of ecological interactions within benthic habitats. 

In a shallow water study in the Ross Sea Cummings et al. (2018) found that 95% of variation within macrofaunal communities was explained by 6 factors including ice duration, ice thickness, snow cover as well as % cover of sediment detritus and sand, % organic content.  

Some pelagic species including krill and salps interact with the benthos. Salps can migrate to a depth of 700 m and are a significant prey species (almost 100% of prey items) consumed by the octocorallian Anthomastus bathyproctus (Gili et al. 2006). For krill, net and acoustic data show that 2-20% of the Scotia Sea summer population can swamp at depths between 200 and 2000 m just above the seabed. Deep migrations and foraging above the seafloor may be important for the coupling of benthic and pelagic environments and food webs (Schmidt et al. 2011). 

Studies generally investigate how the cryo or pelagic environment might influence the benthos, whilst how the benthos may influence the pelagic environment is often ignored Arntz et al. (1999). 

Role in biogeochemical cycles 

Benthic activity has a significant role in biogeochemical cycles. Feeding and burrowing activity reworks sediment that effects the rate of diagenic rations (Kirtensen, 1988). 

Benthic production also has a significant role in carbon storage known as "blue carbon" see Barnes (2017a, 2017b) and Barnes et al. (2018). 

Other interactions e.g. disease

Symbioses

• Abundance of symbiotic associations within the Antarctic benthos have been noted within the Weddell and Ross Sea (see Alvaro and Braco, 2013 and Schiaparelli, 2014).
• Most of the symbioses have shifted towards parasitism with polychaetes and molluscs being the most common symbionts (Schiaparelli et al. 2007, Schiaparelli, 2014)
• Very few of these relationships have been studied in detail with the exception of inquilistic commensalism and parasitic species on echinoderm hosts (Schiapelli et al. 2010, 2011) 
• Current work investigating symbioses between polynoid polychaetes and coral hosts within the South Orkney Islands Southern Shelf MPA (Brasier et al. in prep). 

Human Impacts

Bottom trawling, long-lines

• Kock (1990) first to examine the impacts of bottom trawling in the South Atlantic, indicating that over a 20 year period fishing activities have had a considerable effect on benthic community structure. 
• In Prydz Bay the biomass of benthic invertebrates caught from bottom trawls was greater (up to 10x) than the biomas of finfish. The most abundant species (by biomass) were sponges, ascidians, holothurians and crinoids (Constable, 1991). 
• CCAMLR has a management framework designed for the protection of benthic habitats through the vulnerbale marine ecosystem (VME) indicator taxa. VME taxa are deemed vulnerable to the impacts of destructive fishing, these taxa include many reef building and slow growning species (e.g. corals, sponges) (for details see Parker and Bowden, 2010). A VME risk area is contains a threshold of 10 kg biomass of VME taxa from a single longline segment (1200 m section of longline gear), if this threshold is reached during fishing activities the area is closed to fishing until managment actions are determined by CCAMLR (CCAMLR, 2009)  
• Impacts of longline gear are harder to assess, species may be damaged but not recovered to the surface, in this case it may underestimate the impacts of fishing gear and the presence of VME taxa (Brasier et al. 2018). 
• Regionally high abundance of octopus around Elephant Island which may be due to overfishing of fin fish in that area in the 1980s (Vecchione et al. 2009). 

Marine debris

• Circumpolar recorded of micro and macroplastics in the Southern Ocean (Waller et al. 2017)
• Microplastics have been found in sediments in the Ross Sea (Munari et al. 2017), Admiralty Bay (South Shetland Islands) (Absher et al. 2019), near Rothera Station (Reed et al. 2018), the vicinity of South Georgia (Barnes et al. 2009), the deep Weddell Sea (Van Cauwenberghe et al. 2013). 
• Suspension and deposit feeding animals at risk of microplastic ingestion that can effect food intake, energy reserves and survival rates (Waller et al. 2017). 

Outflow from research stations

• Evidence of sewage enrichment near Casey, Davis and McMurdo Station (Thompson et al. 2003, Conlan et al. 2004, Stark et al. 2015, 2016).
• Implementation of sewage treatment might have reduced this impact.
• Other sources of chemical contamination e.g. oil spills discussed in Tin et al. (2009). 

• High metal concentrations were identified in the Antarctic marine benthos near Rothera research station, but bioaccumulation varied between species. "Anthropogenic activities at Rothera Research Station appeared to have some impact on metal concentrations in the sampled invertebrates, with concentrations of several metals higher in Laternula elliptica near the runway and aircraft activities, but this was not a trend that was detected in the other species." Evidence of trace metal pollution from local anthropogenic sources was limited and sedimentary analysis of bedrock suggest that metals were released into the marine environment by glacial activity (Webb et al. 2019). 

Assessments of Status

IUCN Red List

'None of these species has been assessed for the Red List'

Other

Parker, S.J. & Bowden, D.A. (2010). Identifying taxonomic groups vulnerable to bottom longline fishing gear in the Ross Sea region. CCAMLR Science, 17: 105-127.

References

A list of references referred to on this page.

Please use Ecology style, for more information and examples see: 

https://besjournals.onlinelibrary.wiley.com/hub/journal/13652745/author-guidelines

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