Overcoming the coupled climate and biodiversity crises and their societal impacts.

Earth's biodiversity and human societies face pollution, overconsumption of natural resources, urbanization, demographic shifts, social and economic inequalities, and habitat loss, many of which are exacerbated by climate change. Here, we review links among climate, biodiversity, and society and develop a roadmap toward sustainability. These include limiting warming to 1.5°C and effectively conserving and restoring functional ecosystems on 30 to 50% of land, freshwater, and ocean "scapes." We envision a mosaic of interconnected protected and shared spaces, including intensively used spaces, to strengthen self-sustaining biodiversity, the capacity of people and nature to adapt to and mitigate climate change, and nature's contributions to people. Fostering interlinked human, ecosystem, and planetary health for a livable future urgently requires bold implementation of transformative policy interventions through interconnected institutions, governance, and social systems from local to global levels.

Societal importance of Antarctic negative feedbacks on climate change: blue carbon gains from sea ice, ice shelf and glacier losses.

Diminishing prospects for environmental preservation under climate change are intensifying efforts to boost capture, storage and sequestration (long-term burial) of carbon. However, as Earth's biological carbon sinks also shrink, remediation has become a key part of the narrative for terrestrial ecosystems. In contrast, blue carbon on polar continental shelves have stronger pathways to sequestration and have increased with climate-forced marine ice losses-becoming the largest known natural negative feedback on climate change. Here we explore the size and complex dynamics of blue carbon gains with spatiotemporal changes in sea ice (60-100 MtCyear-1), ice shelves (4-40 MtCyear-1 = giant iceberg generation) and glacier retreat (< 1 MtCyear-1). Estimates suggest that, amongst these, reduced duration of seasonal sea ice is most important. Decreasing sea ice extent drives longer (not necessarily larger biomass) smaller cell-sized phytoplankton blooms, increasing growth of many primary consumers and benthic carbon storage-where sequestration chances are maximal. However, sea ice losses also create positive feedbacks in shallow waters through increased iceberg movement and scouring of benthos. Unlike loss of sea ice, which enhances existing sinks, ice shelf losses generate brand new carbon sinks both where giant icebergs were, and in their wake. These also generate small positive feedbacks from scouring, minimised by repeat scouring at biodiversity hotspots. Blue carbon change from glacier retreat has been least well quantified, and although emerging fjords are small areas, they have high storage-sequestration conversion efficiencies, whilst blue carbon in polar waters faces many diverse and complex stressors. The identity of these are known (e.g. fishing, warming, ocean acidification, non-indigenous species and plastic pollution) but not their magnitude of impact. In order to mediate multiple stressors, research should focus on wider verification of blue carbon gains, projecting future change, and the broader environmental and economic benefits to safeguard blue carbon ecosystems through law.

Intermediate ice scour disturbance is key to maintaining a peak in biodiversity within the shallows of the Western Antarctic Peninsula.

Climate-related disturbance regimes are changing rapidly with profound consequences for ecosystems. Disturbance is often perceived as detrimental to biodiversity; however, the literature is divided on how they influence each other. Disturbance events in nature are diverse, occurring across numerous interacting trophic levels and multiple spatial and temporal scales, leading to divergence between empirical and theoretical studies. The shallow Antarctic seafloor has one of the largest disturbance gradients on earth, due to iceberg scouring. Scour rates are changing rapidly along the Western Antarctic Peninsula because of climate change and with further changes predicted, the Antarctic benthos will likely undergo dramatic shifts in diversity. We investigated benthic macro and megafaunal richness across 10-100 m depth range, much of which, 40-100 m, has rarely been sampled. Macro and megafauna species richness peaked at 50-60 m depth, a depth dominated by a diverse range of sessile suspension feeders, with an intermediate level of iceberg disturbance. Our results show that a broad range of disturbance values are required to detect the predicted peak in biodiversity that is consistent with the Intermediate Disturbance Hypothesis, suggesting ice scour is key to maintaining high biodiversity in Antarctica's shallows.

Variation in zoobenthic blue carbon in the Arctic's Barents Sea shelf sediments.

The flow of carbon from atmosphere to sediment fauna and sediments reduces atmospheric CO2, which in turn reduces warming. Here, during the Changing Arctic Ocean Seafloor programme, we use comparable methods to those used in the Antarctic (vertical, calibrated camera drops and trawl-collected specimens) to calculate the standing stock of zoobenthic carbon throughout the Barents Sea. The highest numbers of morphotypes, functional groups and individuals were found in the northernmost sites (80-81.3° N, 29-30° E). Ordination (non-metric multidimensional scaling) suggested a cline of faunal transition from south to north. The functional group dominance differed across all six sites, despite all being apparently similar muds. Of the environmental variables we measured, only water current speed could significantly explain any of our spatial carbon differences. We found no obvious relationship with sea ice loss and thus no evidence of Arctic blue carbon-climate feedback. Blue carbon in the Barents Sea can be comparable with the highest levels in Antarctic shelf sediments. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Antarctic Futures: An Assessment of Climate-Driven Changes in Ecosystem Structure, Function, and Service Provisioning in the Southern Ocean.

In this article, we analyze the impacts of climate change on Antarctic marine ecosystems. Observations demonstrate large-scale changes in the physical variables and circulation of the Southern Ocean driven by warming, stratospheric ozone depletion, and a positive Southern Annular Mode. Alterations in the physical environment are driving change through all levels of Antarctic marine food webs, which differ regionally. The distributions of key species, such as Antarctic krill, are also changing. Differential responses among predators reflect differences in species ecology. The impacts of climate change on Antarctic biodiversity will likely vary for different communities and depend on species range. Coastal communities and those of sub-Antarctic islands, especially range-restricted endemic communities, will likely suffer the greatest negative consequences of climate change. Simultaneously, ecosystem services in the Southern Ocean will likely increase. Such decoupling of ecosystem services and endemic species will require consideration in the management of human activities such as fishing in Antarctic marine ecosystems.

Marine plastics threaten giant Atlantic Marine Protected Areas.

There has been a recent shift in global perception of plastics in the environment, resulting in a call for greater action. Science and the popular media have highlighted plastic as an increasing stressor [1,2]. Efforts have been made to confer protected status to some remote locations, forming some of the world's largest Marine Protected Areas, including several UK overseas territories. We assessed plastic at these remote Atlantic Marine Protected Areas, surveying the shore, sea surface, water column and seabed, and found drastic changes from 2013-2018. Working from the RRS James Clark Ross at Ascension, St. Helena, Tristan da Cunha, Gough and the Falkland Islands (Figure 1A), we showed that marine debris on beaches has increased more than 10 fold in the past decade. Sea surface plastics have also increased, with in-water plastics occurring at densities of 0.1 items m-3; plastics on seabeds were observed at ≤ 0.01 items m-2. For the first time, beach densities of plastics at remote South Atlantic sites approached those at industrialised North Atlantic sites. This increase even occurs hundreds of meters down on seamounts. We also investigated plastic incidence in 2,243 animals (comprising 26 species) across remote South Atlantic oceanic food webs, ranging from plankton to seabirds. We found that plastics had been ingested by primary consumers (zooplankton) to top predators (seabirds) at high rates. These findings suggest that MPA status will not mitigate the threat of plastic proliferation to this rich, unique and threatened biodiversity.

Severity of seabed spatial competition decreases towards the poles.

For more than a century ecologists have considered that competitive interactions between species are more intense at low latitudes [1,2]. This is frequently invoked as either an explanation or a consequence of higher species richness in the tropics, also suggesting that competition shifts from intra- to inter-specific towards the tropics [1]. Another common assumption is that within a community, intraspecific competition needs to be relatively strong, compared to inter-specific competition, in order to enable stable coexistence of species [3]. However, many analyses have found no consistent large scale geographic patterns in the intensity of intra- or interspecific competition [4]. Here, we show a clear latitudinal trend in contest competition for space in nearshore marine environments, for bryozoans (sessile, colonial, suspension feeding animals). Bryozoans form species-rich assemblages with other encrusting fauna and flora (corraline algae), and are highly abundant across the globe [5]. We find that whilst the intensity of competition (percentage of bryozoan colonies involved in direct physical spatial interactions with bryozoan or other encrusters) differed little with latitude, its severity (percentage of bryozoan colonies involved in contests with a win/loss outcome, leading to death of the loser) was three times lower at the poles than in the tropics. The cause of this change in severity was a strong shift in taxonomic relatedness of competitors, from interactions between species of different families dominating at lower latitudes, to mainly intraspecific competition at the poles.

Antarctic sea ice losses drive gains in benthic carbon drawdown.

Climate forcing of sea-ice losses from the Arctic and West Antarctic are blueing the poles. These losses are accelerating, reducing Earth's albedo and increasing heat absorption. Subarctic forest (area expansion and increased growth) and ice-shelf losses (resulting in new phytoplankton blooms which are eaten by benthos) are the only significant described negative feedbacks acting to counteract the effects of increasing CO2 on a warming planet, together accounting for uptake of ∼10(7) tonnes of carbon per year. Most sea-ice loss to date has occurred over polar continental shelves, which are richly, but patchily, colonised by benthic animals. Most polar benthos feeds on microscopic algae (phytoplankton), which has shown increased blooms coincident with sea-ice losses. Here, growth responses of Antarctic shelf benthos to sea-ice losses and phytoplankton increases were investigated. Analysis of two decades of benthic collections showed strong increases in annual production of shelf seabed carbon in West Antarctic bryozoans. These were calculated to have nearly doubled to >2x10(5) tonnes of carbon per year since the 1980s. Annual production of bryozoans is median within wider Antarctic benthos, so upscaling to include other benthos (combined study species typically constitute ∼3% benthic biomass) suggests an increased drawdown of ∼2.9x10(6) tonnes of carbon per year. This drawdown could become sequestration because polar continental shelves are typically deeper than most modern iceberg scouring, bacterial breakdown rates are slow, and benthos is easily buried. To date, most sea-ice losses have been Arctic, so, if hyperboreal benthos shows a similar increase in drawdown, polar continental shelves would represent Earth's largest negative feedback to climate change.

Effect of disturbance on assemblages: an example using porifera.

Extensive sponge assemblages are found in a number of habitats at Lough Hyne Marine Nature Reserve. These habitats are unusual in experiencing a range of environmental conditions, even though they are only separated by small geographic distances (1-500 m), reducing the possibility of confounding effects between study sites (e.g., silica concentrations and temperature). Sponge assemblages were examined on ephemeral (rocks), stable (cliffs), and artificial (slate panels) hard substrata from high- and low-energy environments that were used to represent two measures of disturbance (flow rate and habitat stability). Sponge assemblages varied considerably between habitat types such that only 26% (25 species) of species reported were common to both rock and cliff habitats. Seven species (of a total of 96 species) were found in the least-developed assemblages (slate panels) and were common to all habitats. Sponge assemblages on rocks and panels varied little between high- and low-energy environments, whereas assemblages inhabiting cliffs varied considerably. Assemblage composition was visualized using Bray-Curtis similarity analysis and Multi-Dimensional Scaling, which enabled differences and similarities between sponge assemblages to be visualized. Cliffs from high- and low-energy sites had different assemblage compositions compared to large rocks, small rocks, and panels, all of which had similar assemblages irrespective of environmental conditions. Differences in assemblages were partially attributed to sponge morphology (shape), as certain morphologies (e.g., arborescent species) were excluded from 2-D rock habitats. Other mechanisms were also considered responsible for the sponge assemblages associated with different habitats.

Resource availability: Ancient homes for hard-up hermit crabs.

Mollusc shells are a vital but sometimes scarce resource for hermit crabs, protecting them from mechanical damage and desiccation, but they require continual replacement as the crab grows. I have discovered that Coenobita rugosus, a large, tropical, semi-terrestrial hermit crab, will resort to using fossil shells when no other suitable casing is available. These unlikely mobile homes fall out of coastal limestone as it is eroded by the sea in southwestern Madagascar, placing the occupants alongside Homo sapiens as resourceful exploiters of prehistoric animal remains.

Overgrowth competition between clades: implications for interpretation of the fossil record and overgrowth indices.

Overgrowth interactions (2693 in total) were observed among three major groups (arguably clades) of bryozoans--cheilostomatids (57 species), ctenostomatids (3 species), and cyclostomatids (14 species). The bryozoans studied here occur in shallow water at high-temperate polar latitudes where they encrust hard substrata such as rock piles. The main study site was the intertidal and infralittoral zones of Kodiak Island, Alaska, but observations were also made in similar zones of South Georgia Island and the Falkland Islands in the South Atlantic Ocean. Cheilostomatids dominated the number of species, individuals, and interactions at all depths. Intraclade interactions formed 73.7% of the encounters for cheilostomatids, 1.6% for ctenostomatids, and 5.7% for cyclostomatids. The competitive ranking of the three clades was broadly ctenostomatids > cyclostomatids > cheilostomatids. Significantly, these results contradict all previous quantitative studies of bryozoan overgrowth, in which cheilostomatids are reported to overgrow cyclostomatids at a higher rate. From these studies and the literature, we calculated win indices to vary from 0 to 0.42 for living cyclostomatids, from 0.08 to 0.9 for living cheilostomatids, and from 0.25 to 0.75 for living ctenostomatids. The win indices of cyclostomatid and cheilostomatid clades show significantly more variation in living assemblages than in fossil assemblages. This disparity may be due to differential preservation (polar and subpolar assemblages last less than 4 years). The diversity was very high in terms of both species richness and interaction types (outcomes between competitor pairs). Comparison with the literature suggests the possibility that nearshore diversity of bryozoans may be bimodal (have two peaks) between high arctic and antarctic latitudes. Indices of success in overgrowth competition have been constructed in various ways. For cheilostomatids, the method of calculation had little influence on the ranking of representatives. In contrast, the apparent success of ctenostomatids and cyclostomatids varied hugely with how the index was calculated. This inconsistency is due to the use of very different strategies in overgrowth competition; among the two latter groups, many interactions involve tied outcomes.

Inheritance of Resistance to Pratylenchus penetrans in Alfalfa.

Fifty-two alfalfa (Medicago sativa L.) clones, randomly selected from the cultivar Baker and the experimental line MNGRN-4, were evaluated for resistance (based on nematode reproduction) to Pratylenchus penetrans in growth chamber tests (25 C). Twenty-five clones, representing the range of nematodes and eggs per plant, were selected and retested. Four moderately resistant and two susceptible alfalfa clones were identified. Inheritance of resistance to P. penetrans was studied in these six clones using a diallel mating design. The S, Fl, and reciprocal progenies differed for numbers of nematodes and eggs per g dry root and for shoot and root weights (P < 0.05). Resistance, measured as numbers of nematodes in roots, was correlated between parental clones and their S families (r = 0.94), parental clones and their half-sib families (r = 0.81), and S and half-sib families (r = 0.88). General combining ability (GCA) effects were significant for nematode resistance traits. Both GCA and specific combining ability (SCA) effects were significant for plant size traits, but SCA was more important than GCA in predicting progeny plant size. Reciprocal effects were significant for both nematode resistance and plant size traits, which may slow selection progress in long-term selection programs. However, the GCA effects are large enough that breeding procedures that capitalize on additive effects should be effective in developing alfalfa cultivars with resistance to P. penetrans.

Molecular and whole-plant responses to selection for enzyme activity in alfalfa root nodules: evidence for molecular compensation of aspartate aminotransferase expression.

The enzyme aspartate aminotransferase (AAT) plays a key role in the assimilation of fixed-N in alfalfa (Medicago sativa L.) root nodules. AAT activity in alfalfa nodules is due to the activity of two dimeric isozymes, AAT-1 and AAT-2, that are products of two distinct genes. Three forms of AAT-2 (AAT-2a, -2b, and-2c) have been identified. It was hypothesized that two alleles occur at the AAT-2 locus, giving rise to the three AAT-2 enzymes. In a prior study bidirectional selection for root nodule AAT and asparagine synthetase (AS) activities on a nodule fresh weight basis in two diverse alfalfa germ plasms resulted in high nodule enzyme activity subpopulations with about 20% more nodule AAT activity than low enzyme activity subpopulations. The objectives of the study presented here were to determine the inheritance of nodule AAT-2 production and to evaluate the effect of bidirectional selection for AAT and AS on AAT-2 allelic frequencies, the relative contributions of AAT-1 and AAT-2 to total nodule activity, nodule enzyme concentration, and correlated traits. Two alleles at the AAT-2 locus were verified by evaluating segregation of isozyme phenotypes among F1 and S1 progeny of crosses or selfs. Characterization of subpopulations for responses associated with selection was conducted using immunoprecipitation of in vitro nodule AAT activity, quantification of AAT enzyme protein by ELISA, and AAT activity staining of native isozymes on PAGE. Results indicate that selection for total AAT activity specifically altered the expression of the nodule AAT-2 isozyme. AAT-2 activity was significantly greater in high compared to low activity subpopulations, and high AAT subpopulations from both germ plasms had about 18% more AAT-2 enzyme (on a nodule fresh weight basis). No significant or consistent changes in AAT-2 genotypic frequencies in subpopulations were caused by selection for AAT activity. Since changes in AAT activity were not associated with changes in AAT-2 genotype, selection must have affected a change(s) at another locus (or loci), which indirectly effects the expression of nodule AAT.

Nitrogen Fixation, Nodule Development, and Vegetative Regrowth of Alfalfa (Medicago sativa L.) following Harvest.

Nitrogenase-dependent acetylene reduction, nodule function, and nodule regrowth were studied during vegetative regrowth of harvested (detopped) alfalfa (Medicago sativa L.) seedlings grown in the glasshouse. Compared with controls, harvesting caused an 88% decline in acetylene reduction capacity of detached root systems within 24 hours. Acetylene reduction in harvested plants remained low for 13 days, then increased to a level comparable to the controls by day 18.Protease activity increased in nodules from harvested plants, reached a maximum at day 7 after harvest, and then declined to a level almost equal to the control by day 22 after harvest. Soluble protein and leghemoglobin decreased in nodules from harvested plants in an inverse relationship to protease activity.Nitrate reductase activity of nodules from harvested plants increased significantly within 24 hours and was inversely associated with acetylene reduction. The difference in nitrate reductase between nodules from harvested plants and control plants became less evident as shoot regrowth occurred and as acetylene reduction increased in the harvested plants.No massive loss of nodules occurred after harvest as evidenced by little net change in nodule fresh weight. There was, however, a rapid localized senescence which occurred in nodules of harvested plants. Histology of nodules from harvested plants showed that they degenerated at the proximal end after harvest. Starch in the nodule was depleted by 10 days after harvest. The meristem and vascular bundles of nodules from harvested plants remained intact. The senescent nodules began to regrow and fix nitrogen after shoot growth resumed.