Diversity and Disturbances in the Antarctic Megabenthos: Feasible versus Theoretical Disturbance Ranges

The intermediate disturbance hypothesis (IDH) predicts a hump-shaped relationship between regional diversity and the disturbance rate. We tested the IDH for the megabenthos inhabiting the Antarctic sea floor, which is disturbed by iceberg scouring. We used models based on the empirical knowledge of succession to calculate the IDH curve for this system and to extrapolate the presently observable range of the IDH curve to higher and lower disturbance rates. Although the hump-shaped relationship has been found for a purely theoretical (extremely large) disturbance range, within the feasible disturbance range (assumed as realistic in the Antarctic region under climate change), the regional diversity of successional stages due to iceberg scouring strongly decreases with lower disturbance rates but levels off only slowly with higher disturbance rates. The reason is the unevenness in the lifetimes of the successional stages, in that early stages are short-lived whereas late stages are long-lived. With such unevenness, increasing disturbances support the early stages without jeopardizing the later ones. Additionally, we converted this regional diversity of stages to the regional diversity of taxa using a transformation formula based on empirical knowledge of the number and mean abundance of taxa in the particular stages. Our results suggest that a decrease in iceberg scouring due to climate change would be more detrimental to the diversity of the Antarctic megabenthos than an increase.     

Polychaete assemblages of the northwestern Ross Sea shelf: worming out the environmental drivers of Antarctic macrobenthic assemblage composition

We have limited understanding of which environmental factors structure the distribution patterns and composition of Antarctic macrobenthos assemblages, and the spatial scales on which such factors operate. In 2004, the “BioRoss Survey” was conducted on the northwestern Ross Sea shelf between Cape Adare and Cape Hallett in depths of 50–750 m to describe and quantify the assemblage patterns of benthic macroinvertebrates. In order to determine the influence of primary productivity, disturbance and habitat heterogeneity on the distribution and composition of the macrofaunal assemblages, polychaete data derived from 52 grab samples were analysed. Although differences in the composition of polychaete assemblages among different sampling transects and depth strata were not particularly pronounced (yet statistically significant), the results suggested that large-scale differences in both primary productivity and iceberg disturbance influence distribution patterns. The combination of sediment chl a content, sorting coefficient, sponge spicule content and distance to the nearest iceberg scour best explained polychaete assemblage patterns. This finding supports previous contentions that multiple environmental drivers working at varying scales influence Antarctic shelf assemblages. The results do not supply support for a pronounced decoupling of pelagic and benthic systems, as has been suggested by another study of deeper water benthic assemblages on the Ross Sea shelf.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

Recent research on Arctic benthos: common notions need to be revised

Increased public awareness of the global significance of polar regions and opening of the Russian Arctic to foreign researchers have led to a pronounced intensification of benthic research in Arctic seas. The wealth of information gathered in these efforts has markedly enhanced our knowledge on the Arctic benthos. While some scientific concepts have been corroborated by the novel findings (e.g., low endemism and high faunistic affinity to northern Atlantic assemblages), other common notions need to be revised, particularly with regard to the often-cited differences between Arctic seas and the Southern Ocean. It has been demonstrated that benthos assemblages vary broadly in diversity between Arctic regions and that, hence, the idea of a consistently poor Arctic benthos—being in stark contrast to the rich Antarctic bottom fauna—is an undue overgeneralization. In terms of biogeographic diversity, both Arctic and Antarctic waters seem to be characterized by intermediate species richness. Levels of disturbance—a major ecological agent known to heavily affect benthic diversity and community structure—have been assumed to be relatively high in the Arctic but exceptionally low in the Southern Ocean. The discovery of the great role of iceberg scouring in Antarctic shelf ecosystems, which has largely been overlooked in the past, calls for a reconsideration of this notion. The novel data clearly demonstrate that there are marked differences in geographical and environmental setting, impact of fluvial run-off, pelagic production regime, strength of pelago–benthic coupling and, hence, food supply to the benthos among the various Arctic seas, impeding the large-scale generalization of local and regional findings. Field evidence points to the great significance of meso-scale features in hydrography and ice cover (marginal ice zones, polynyas, and gyres) as ‘hot spots’ of tight pelago–benthic coupling and, hence, high benthic biomass. In contrast, the importance of terrigenic organic matter discharged to the Arctic seas through fluvial run-off as an additional food source for the benthos is still under debate. Studies on the partitioning of energy flow through benthic communities strongly suggest that megafauna has to be adequately considered in overall benthic energy budgets and models of carbon cycling, particularly in Arctic shelf systems dominated by abundant echinoderm populations. Much progress has been made in the scientific exploration of the deep ice-covered Arctic Ocean. There is now evidence that it is one order of magnitude more productive than previously thought. Therefore, the significance of shelf–basin interactions, i.e., the importance of excess organic carbon exported from productive shelves to the deep ocean, is still debated and, hence, a major topic of on-going research. Another high-priority theme of current/future projects are the ecological consequences of the rapid warming in the Arctic. Higher water temperatures, increased fluvial run-off and reduced ice cover will give rise to severe ecosystem changes, propagating through all trophic levels. It is hypothesized that there would be a shift in the relative importance of marine biota in the overall carbon and energy flux, ultimately resulting in a switch from a ‘sea-ice algae–benthos’ to a ‘phytoplankton–zooplankton’ dominance.     

Disturbance, colonization and development of Antarctic benthic communities

A decade has yielded much progress in understanding polar disturbance and community recovery-mainly through quantifying ice scour rates, other disturbance levels, larval abundance and diversity, colonization rates and response of benthos to predicted climate change. The continental shelf around Antarctica is clearly subject to massive disturbance, but remarkably across so many scales. In summer, millions of icebergs from sizes smaller than cars to larger than countries ground out and gouge the sea floor and crush the benthic communities there, while the highest wind speeds create the highest waves to pound the coast. In winter, the calm associated with the sea surface freezing creates the clearest marine water in the world. But in winter, an ice foot encases coastal life and anchor ice rips benthos from the sea floor. Over tens and hundreds of thousands of years, glaciations have done the same on continental scales-ice sheets have bulldozed the seabed and the zoobenthos to edge of shelves. We detail and rank modern disturbance levels (from most to least): ice; asteroid impacts; sediment instability; wind/wave action; pollution; UV irradiation; volcanism; trawling; non-indigenous species; freshwater inundation; and temperature stress. Benthic organisms have had to recolonize local scourings and continental shelves repeatedly, yet a decade of studies have demonstrated that they have (compared with lower latitudes) slow tempos of reproduction, colonization and growth. Despite massive disturbance levels and slow recolonization potential, the Antarctic shelf has a much richer fauna than would be expected for its area. Now, West Antarctica is among the fastest warming regions and its organisms face new rapid changes. In the next century, temperature stress and non-indigenous species will drastically rise to become dominant disturbances to the Antarctic life. Here, we describe the potential for benthic organisms to respond to disturbance, focusing particularly on what we know now that we did not a decade ago.     

森林景观恢复过程中景观要素斑块规模的动态分析

以１９５９年以来的４期航片为基础资料,通过对研究地区各类景观要素的斑块平均规模和粒级结构及其动态变化过程进行分析,揭示了研究地区森林景观恢复演替过程中景观要素斑块动态特点,阐明了植被演替和干扰格局控制下森林景观要素斑块恢复演替的过程和规律。研究地区森林景观自５０年代以来在整体上始终属于细景观,但景观要素斑志规模和粒级结构发生了显著变化,在三个不同时期表现出不同的变化特点,而且不同景观要素的斑块规模     

Response of Antarctic benthic communities to disturbance: first results from the artificial Benthic Disturbance Experiment on the eastern Weddell Sea Shelf, Antarctica

The long-term benthic disturbance experiment (BENDEX) was started on the eastern Weddell Sea shelf off Austasen (Antarctica) during ‘Polarstern’ cruise ANT XXI/2 in December 2003 to simulate the impact of grounding icebergs on the seabed and follow the steps and timescales of recovery of disturbed benthos and demersal fish communities. Here, we report the basic approach and first results for this experimental field study. By means of 11 densely-placed hauls with a modified bottom trawl, a seabed area of approximately 100 × 1000 m was artificially scoured to inflict a similar damage to the benthic habitats as a grounding iceberg. Before the disturbance event and 11 days after it, the seafloor communities were sampled (invertebrate assemblages by multibox corers, the fish fauna by trawl hauls) and comparatively analyzed. Sediment texture and chemistry was not significantly altered by the heavy disturbance inflicted by repeated trawling, whereas the fauna was negatively affected. Invertebrate benthic biomass was drastically reduced by a factor of 10, while mean abundances were only slightly reduced. Demersal fish biomass and abundance were slightly but not significantly smaller after the disturbance. Effects of disturbance became more evident in the composition of the fish fauna, with Trematomus pennelli and T. hansoni being dominant at disturbed sites, whereas Chionodraco myersi was the dominant species in trawl catches from undisturbed stations.     

Structural changes of vegetation and its association with microclimate in a successional gradient of low thorn forest in northeastern Mexico

Understanding how microclimate and vegetation are associated during secondary succession is of primary importance for plant conservation in the face of the increasing land cover modification. However, these patterns are still unstudied for many plant communities. This study aimed to evaluate the structure (species richness, Shannon's diversity index, Simpson´s dominance index, abundance of each species, average height of species, species cover (%), species composition, and indicator values) of a low thorn forest fragment and to analyze its relation with microclimate along a successional gradient. Four stages of succession were delimited by the analysis of Landsat images, in the state of Tamaulipas, northeast Mexico. Statistical models incorporated species richness, diversity indices, abundance, height, and cover, as variables for searching differences between stages, or to evaluate microclimate associations. A total of 70 species, 54 genera, and 27 families were determined. Height of tree layer was the most important variable for discrimination of the successional stages. Conserved areas differed floristically from other stages, associated mainly with the lowest values of wind speed originated by tree layer characteristics. A significant association between species and microclimate was found, being wind speed and relative humidity the most important variables. Some species, due to their high importance values and their patterns of association with microclimate, may be considered as key taxa for low thorn forest, which is a threatened semitropical community in northeast Mexico. Conserved and late successional areas account for climatic regulation of this plant community, and the importance of these forest patches may be considered when establishing biodiversity protection areas.

     

Variation of scavenger richness and abundance between sites of high and low iceberg scour frequency in Ryder Bay,west Antarctic Peninsula

Physical disturbance, particularly from iceberg scour, is a major structuring force in polar benthic communities at shelf depths. Scouring kills and damages benthic organisms providing food for the abundant scavenging fauna of coastal Antarctic waters. This trophic group is likely to be strongly affected by changes in iceberg scouring. A baited underwater camera system was used to examine the distribution of scavenging fauna in relation to the spatial variation in exposure to iceberg impacts experienced at different iceberg scouring conditions and depths within Ryder Bay. The results indicate that the relationships between depth and scavenger abundance and assemblage composition differed between high and low scour sites. Scavenger abundance increased with depth at high scour sites and fell with depth a low scour sites. There was also significant difference in community composition between sites within each scouring condition. Scavenger species richness also exhibited an increase with depth at most sites consistent with the established pattern of declining iceberg scouring frequency with depth. Shannon–Wiener diversity increased with depth but significantly more steeply in highly scoured sites. Our results suggest that depth and exposure to icebergs interact to shape the scavenger community. The significant differences within the high and low scour groups suggest that other factors remain to be investigated and that there is probably a nonlinear relationship between scouring intensity and the favourability of a site for scavengers.     

Disturbance frequency influences patch dynamics in stream benthic algal communities

Disturbance is integral to the organisation of riverine ecosystems. Fluctuating low flows caused by supra-seasonal drought and water management periodically dewater habitat patches, potentially creating heterogeneity in the taxonomic composition and successional dynamics of benthic communities. The frequency of disturbance induced by low flows is contingent upon the topography of the river bed and thus varies among patches. We investigated whether the frequency of patch dewatering influenced the structure and temporal dynamics of benthic algal communities attached to the upper surfaces of stones in stream mesocosms (4 m²). In a 693-day disturbance experiment, we applied short dewatering disturbances (6 days) at high (33-day cycles) and low frequencies (99-day cycles) and compared algal assemblages with undisturbed controls at 21 endpoints. In the absence of disturbance, epilithic space was dominated by the green encrusting alga Gongrosira incrustans. However, drying disturbances consistently reduced the dominance of the green alga, and crust abundance decreased with increasing disturbance frequency, thereby opening space for a diversity of mat-forming diatoms. The response of mat diatoms to disturbance varied markedly during the experiment, from strong reductions in the abundance of loosely attached mats in mid-late 2000 to the exploitation of open space by closely adhering mats in 2001. Contrary responses were attributed to changes in the species composition of mat diatoms, which influenced the physiognomy and hence stress-resistance and resilience of the assemblage. Our results indicate that patchy dewatering of habitat patches during periods of low flow influences the successional dynamics of algae, thereby creating distinctive mosaics on the stream bed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

On the origin of Antarctic marine benthic community structure

Environmental conditions fostering marine communities around Antarctica differ fundamentally from those in the rest of the world's oceans, particularly in terms of pronounced climatic fluctuations and extreme cold. Here, we argue that the rarity of pelagic larval stages in Antarctic marine benthic invertebrate species is a consequence of evolutionary temperature adaptation and that this has greatly contributed to the current structure of the Antarctic benthic community. In arguing this position, we challenge the likelihood of previously suggested survival strategies of benthic communities on the Antarctic continental shelf and slope during Cenozoic glacial periods. By integrating evidence from marine geology and geophysics, we suggest that the Antarctic continental shelf and slope were both unfavourable environments for benthic communities during glacial periods and that community survival was only possible in the deep sea or in shelters on the continental shelf as a result of the diachronism in maximum ice extent.     

Response of nematode communities after large‐scale ice‐shelf collapse events in the Antarctic Larsen area

Owing to large‐scale ice‐shelf disintegration events, the Antarctic Larsen A and B areas recently became ice‐free. During the ANT‐XXIII/8 Polarstern campaign, this region was sampled for the first time. Our study is the first to investigate benthic communities in this area and their response to the collapse of ice shelves in the Antarctic. The nematofauna appears to be strongly influenced by the sudden ice‐cover removal, although its response differs from that of the macro‐ and megabenthos. Our results indicate that precollapse, sub‐ice communities were impoverished and characterized by low densities, low diversity and high dominance of a few taxa. This might still be visible at a station located deep inside the Larsen B embayment, where Halomonhystera was dominant. Post‐collapse recolonization of the ‘inner’ stations, i.e. those located furthermost from the former ice‐shelf edge, is believed to be a long‐time process. At the time of sampling, community structure at the inner stations was not or only slightly influenced by colonization, and might be structured by local environmental conditions. Our results indicate that a locally increased food supply after ice‐cover removal could provoke a faster, local response of the nematode assemblages compared with the response due to recolonization. Thalassomonhystera is recognized as an opportunist, taking advantage of increased food supply at inner stations A_South and B_North. Communities living close to the former ice‐shelf edge are believed to be at an intermediate or late stage of succession, with a dominance of Microlaimus, a common Antarctic genus and quick colonizer. Densities here were comparable with those at other Antarctic stations, whereas they were considerably decreased at the inner stations. In general, the collapse of the Larsen ice shelves initially has a positive effect on the shelf nematode fauna in the area, both in terms of abundance and diversity.     

甘肃兴隆山自然保护区森林演替对鸟类群落结构的影响

运用固定面积法调查了青林三个主要演替阶段森林(阔叶林、混交林和针叶林)的鸟类群落结构。结果表明,混交林中鸟类物种数、鸟类物种多样性和种团多样性最高,阔叶林最低。方差分析和聚类分析均表明,三种演替阶段森林的鸟类群落组成差异显著。所记录到的35种鸟类中有11种差异显著。其中,山雀科和科在阔叶林中占优势,而柳莺亚科在混交林和针叶林中占优势。食性种团中,阔叶林以食虫鸟和食果食虫鸟为主;混交林和针叶林均以食虫鸟、食果食虫鸟、食谷鸟和食果鸟为主。鉴于阔叶林和混交林有向顶级群落(针叶林)演替的趋势,建议采取人为干扰来维持三种演替阶段森林的面积,以便从景观水平上保护鸟类物种多样性。     

Successional theories

Succession is a fundamental concept in ecology because it indicates how species populations, communities, and ecosystems change over time on new substrate or after a disturbance. A mechanistic understanding of succession is needed to predict how ecosystems will respond to land-use change and to design effective ecosystem restoration strategies. Yet, despite a century of conceptual advances a comprehensive successional theory is lacking. Here we provide an overview of 19 successional theories (‘models’) and their key points, group them based on conceptual similarity, explain conceptual development in successional ideas and provide suggestions how to move forward. Four groups of models can be recognised. The first group (patch & plants) focuses on plants at the patch level and consists of three subgroups that originated in the early 20th century. One subgroup focuses on the processes (dispersal, establishment, and performance) that operate sequentially during succession. Another subgroup emphasises individualistic species responses during succession, and how this is driven by species traits. A last subgroup focuses on how vegetation structure and underlying demographic processes change during succession. A second group of models (ecosystems) provides a more holistic view of succession by considering the ecosystem, its biota, interactions, diversity, and ecosystem structure and processes. The third group (landscape) considers a larger spatial scale and includes the effect of the surrounding landscape matrix on succession as the distance to neighbouring vegetation patches determines the potential for seed dispersal, and the quality of the neighbouring patches determines the abundance and composition of seed sources and biotic dispersal vectors. A fourth group (socio-ecological systems) includes the human component by focusing on socio-ecological systems where management practices have long-lasting legacies on successional pathways and where regrowing vegetations deliver a range of ecosystem services to local and global stakeholders. The four groups of models differ in spatial scale (patch, landscape) or organisational level (plant species, ecosystem, socio-ecological system), increase in scale and scope, and reflect the increasingly broader perspective on succession over time. They coincide approximately with four periods that reflect the prevailing view of succession of that time, although all views still coexist. The four successional views are: succession of plants (from 1910 onwards) where succession was seen through the lens of species replacement; succession of communities and ecosystems (from 1965 onwards) when there was a more holistic view of succession; succession in landscapes (from 2000 onwards) when it was realised that the structure and composition of landscapes strongly impact successional pathways, and increased remote-sensing technology allowed for a better quantification of the landscape context; and succession with people (from 2015 onwards) when it was realised that people and societal drivers have strong effects on successional pathways, that ecosystem processes and services are important for human well-being, and that restoration is most successful when it is done by and for local people. Our review suggests that the hierarchical successional framework of Pickett is the best starting point to move forward as this framework already includes several factors, and because it is flexible, enabling application to different systems. The framework focuses mainly on species replacement and could be improved by focusing on succession occurring at different hierarchical scales (population, community, ecosystem, socio-ecological system), and by integrating it with more recent developments and other successional models: by considering different spatial scales (landscape, region), temporal scales (ecosystem processes occurring over centuries, and evolution), and by taking the effects of the surrounding landscape (landscape integrity and composition, the disperser community) and societal factors (previous and current land-use intensity) into account. Such a new, comprehensive framework could be tested using a combination of empirical research, experiments, process-based modelling and novel tools. Applying the framework to seres across broadscale environmental and disturbance gradients allows a better insight into what successional processes matter and under what conditions.     

Scale- and intensity-dependent disturbance determines the magnitude of opportunistic response

Opportunistic, fugitive or pioneer species are species that posses life-history characteristics that allow them to respond quickly to disturbances. Their abundance during the early stages of succession is central to ecological models of benthic soft-sediment succession and these species can play important roles in affecting subsequent successional trajectories. Nevertheless, numerous studies have demonstrated seemingly random patterns of opportunistic responses following disturbance, questioning the generality of currently accepted successional models. In this paper we provide examples from two case studies and argue that the spatial scale or magnitude of disturbance is key to the development of opportunistic responses, and that the scale of disturbance may be particularly important in determining (a) the levels of resources made available and (b) the magnitude of release from competitive interactions, which permit opportunists to flourish.     

Effects of Late-Cenozoic Glaciation on Habitat Availability in Antarctic Benthic Shrimps (Crustacea: Decapoda: Caridea)

Marine invertebrates inhabiting the high Antarctic continental shelves are challenged by disturbance of the seafloor by grounded ice, low but stable water temperatures and variable food availability in response to seasonal sea-ice cover. Though a high diversity of life has successfully adapted to such conditions, it is generally agreed that during the Last Glacial Maximum (LGM) the large-scale cover of the Southern Ocean by multi-annual sea ice and the advance of the continental ice sheets across the shelf faced life with conditions, exceeding those seen today by an order of magnitude. Conditions prevailing at the LGM may have therefore acted as a bottleneck event to both the ecology as well as genetic diversity of today''s fauna. Here, we use for the first time specific Species Distribution Models (SDMs) for marine arthropods of the Southern Ocean to assess effects of habitat contraction during the LGM on the three most common benthic caridean shrimp species that exhibit a strong depth zonation on the Antarctic continental shelf. While the shallow-water species Chorismus antarcticus and Notocrangon antarcticus were limited to a drastically reduced habitat during the LGM, the deep-water shrimp Nematocarcinus lanceopes found refuge in the Southern Ocean deep sea. The modeling results are in accordance with genetic diversity patterns available for C. antarcticus and N. lanceopes and support the hypothesis that habitat contraction at the LGM resulted in a loss of genetic diversity in shallow water benthos.     

Old-field successional dynamics on the Inland Pampa,Argentina

Abstract. The first 10 yr of old-field successional dynamics on the Argentine Inland Pampa were studied on a series of adjacent plots established consecutively between 1978 and 1989. We examined differences in species abundance patterns among plots in order to detect the spatial and temporal variability of succession. Perennial grasses steadily increased in cover and replaced the dominant annual species after 5 yr. Pioneer dicots persisted in older seral stages with 20 — 23 species/plot. Overall, exotic species (mostly the grasses Lolium multiflorum and Cynodon dactylon) contributed much to the plant cover in these communities. Native grasses comprised 45 % of total cover at years 7 — 10 of succession, but occurred with less than 7 species/plot. Substantial variation was found in the successional pathway, which reflected the particular sequence from annual forbs to short-lived and perennial grasses in the various plots. The course of succession was apparently influenced by a 2-yr period of unusually high rainfall. Deyeuxia viridiflavescens, a native perennial grass virtually absent before the wet period, spread over the study area and dominated seral communities for 3 yr, irrespective of plot age. Climatic conditions thus affected the successional turnover of life forms by increasing the rate of colonization by perennial grasses. We further point out the constraints imposed on secondary succession by the life histories of ‘available’ species.     

Perspective: Increasing blue carbon around Antarctica is an ecosystem service of considerable societal and economic value worth protecting

Precautionary conservation and cooperative global governance are needed to protect Antarctic blue carbon: the world's largest increasing natural form of carbon storage with high sequestration potential. As patterns of ice loss around Antarctica become more uniform, there is an underlying increase in carbon capture‐to‐storage‐to‐sequestration on the seafloor. The amount of carbon captured per unit area is increasing and the area available to blue carbon is also increasing. Carbon sequestration could further increase under moderate (+1°C) ocean warming, contrary to decreasing global blue carbon stocks elsewhere. For example, in warmer waters, mangroves and seagrasses are in decline and benthic organisms are close to their physiological limits, so a 1°C increase in water temperature could push them above their thermal tolerance (e.g. bleaching of coral reefs). In contrast, on the basis of past change and current research, we expect that Antarctic blue carbon could increase by orders of magnitude. The Antarctic seafloor is biophysically unique and the site of carbon sequestration, the benthos, faces less anthropogenic disturbance than any other ocean continental shelf environment. This isolation imparts both vulnerability to change, and an avenue to conserve one of the world's last biodiversity refuges. In economic terms, the value of Antarctic blue carbon is estimated at between £0.65 and £1.76 billion (~2.27 billion USD) for sequestered carbon in the benthos around the continental shelf. To balance biodiversity protection against society's economic objectives, this paper builds on a proposal incentivising protection by building a ‘non‐market framework’ via the 2015 Paris Agreement to the United Nations Framework Convention on Climate Change. This could be connected and coordinated through the Antarctic Treaty System to promote and motivate member states to value Antarctic blue carbon and maintain scientific integrity and conservation for the positive societal values ingrained in the Antarctic Treaty System.     

Iceberg killing fields limit huge potential for benthic blue carbon in Antarctic shallows

Climate‐forced ice losses are increasing potential for iceberg‐seabed collisions, termed ice scour. At Ryder Bay, West Antarctic Peninsula (WAP) sea ice, oceanography, phytoplankton and encrusting zoobenthos have been monitored since 1998. In 2003, grids of seabed markers, covering 225 m², were established, surveyed and replaced annually to measure ice scour frequency. Disturbance history has been recorded for each m² of seabed monitored at 5–25 m for ~13 years. Encrusting fauna, collected from impacted and nonimpacted metres each year, show coincident benthos responses in growth, mortality and mass of benthic immobilized carbon. Encrusting benthic growth was mainly determined by microalgal bloom duration; each day, nanophytoplankton exceeded 200 μg L^?1 produced ~0.05 mm radial growth of bryozoans, and sea temperature >0 °C added 0.002 mm day^?1. Mortality and persistence of growth, as benthic carbon immobilization, were mainly influenced by ice scour. Nearly 30% of monitored seabed was hit each year, and just 7% of shallows were not hit. Hits in deeper water were more deadly, but less frequent, so mortality decreased with depth. Five‐year recovery time doubled benthic carbon stocks. Scour‐driven mortality varied annually, with two‐thirds of all monitored fauna killed in a single year (2009). Reduced fast ice after 2006 ramped iceberg scouring, killing half the encrusting benthos each year in following years. Ice scour coupled with low phytoplankton biomass drove a phase shift to high mortality and depressed zoobenthic immobilized carbon stocks, which has persevered for 10 years since. Stocks of immobilized benthic carbon averaged nearly 15 g m^?2. WAP ice scouring may be recycling 80 000 tonnes of carbon yr^?1. Without scouring, such carbon would remain immobilized and the 2.3% of shelf which are shallows could be as productive as all the remaining continental shelf. The region's future, when glaciers reach grounding lines and iceberg production diminishes, is as a major global sink of carbon storage.