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Abstract The equilibrium theory of island biogeography (ETIB) is a widely applied dynamic theory proposed in the 1960s to explain why islands have coherent differences in species richness. The development of the ETIB was temporarily challenged in the 1970s by the alternative static theory of ecological impoverishment (TEI). The TEI suggests that the number of species on an island is determined by its number of habitats or niches but, with no clear evidence relating species richness to the number of niches however, the TEI has been almost dismissed as a theory in favour of the original ETIB. Here, we show that the number of climatic niches on islands is an important predictor of the species richness of plants, herpetofauna and land birds. We therefore propose a model called the niche‐based theory of island biogeography (NTIB), based on the MacroEcological Theory on the Arrangement of Life (METAL), which successfully integrates the number of niches sensu Hutchinson into ETIB. To account for greater species turnover at the beginning of colonisation, we include higher initial extinction rates. When we test our NTIB for resident land birds in the Krakatau Islands, it reveals a good correspondence with observed species richness, immigration and extinction rates. Provided the environmental regime remains unchanged, we estimate that the current species richness at equilibrium is ~45 species (range between 38.39 and 61.51). Our NTIB provides better prediction because it counts for changes in species richness with latitude, which is not considered in any theory of island biogeography.

Abstract Global cooling has been proposed as a driver of the Great Ordovician Biodiversification Event, the largest radiation of Phanerozoic marine animal Life. Yet, mechanistic understanding of the underlying pathways is lacking and other possible causes are debated. Here we couple a global climate model with a macroecological model to reconstruct global biodiversity patterns during the Ordovician. In our simulations, an inverted latitudinal biodiversity gradient characterizes the late Cambrian and Early Ordovician when climate was much warmer than today. During the Mid-Late Ordovician, climate cooling simultaneously permits the development of a modern latitudinal biodiversity gradient and an increase in global biodiversity. This increase is a consequence of the ecophysiological limitations to marine Life and is robust to uncertainties in both proxy-derived temperature reconstructions and organism physiology. First-order model-data agreement suggests that the most conspicuous rise in biodiversity over Earth’s history – the Great Ordovician Biodiversification Event – was primarily driven by global cooling.

Abstract Planktonic Foraminifera are unique paleo-environmental indicators through their excellent fossil record in ocean sediments. Their distribution and diversity are affected by different environmental factors including anthropogenically forced ocean and climate change. Until now, historical changes in their distribution have not been fully assessed at the global scale. Here we present the FORCIS (Foraminifera Response to Climatic Stress) database on foraminiferal species diversity and distribution in the global ocean from 1910 until 2018 including published and unpublished data. The FORCIS database includes data collected using plankton tows, continuous plankton recorder, sediment traps and plankton pump, and contains ~22,000, ~157,000, ~9,000, ~400 subsamples, respectively (one single plankton aliquot collected within a depth range, time interval, size fraction range, at a single location) from each category. Our database provides a perspective of the distribution patterns of planktonic Foraminifera in the global ocean on large spatial (regional to basin scale, and at the vertical scale), and temporal (seasonal to interdecadal) scales over the past century.

Long term variability in temperature and wind patterns impacts diatom populations in the Northeast Atlantic, with warming leading to increased abundance in cold oceanic regions and decreased abundance in warm regions, according to an analysis of 60-years of continuous plankton recorder observations.

Abstract Fisheries and aquaculture are facing many challenges worldwide, especially adaptation to climate change. Investigating future distributional changes of largely harvested species has become an extensive research topic, aiming at providing realistic ecological scenarios on which to build management measures, to help fisheries and aquaculture adapt to future climate-driven changes. Here, we use an ensemble modelling approach to estimate the contemporary and future distributional range of eight demersal fish species of high economic value in the Mediterranean Sea. We identify a cardinal influence of (i) temperature on fish species distributions, all being shaped by yearly mean and seasonality in sea bottom temperature, and (ii) the primary production. By assessing the effects of changes in future climate conditions under three Representative Concentration Pathway (RCP2.6, RCP4.5 and RCP8.5) scenarios over three periods of the twenty-first century, we project a contraction of the distributional range of the eight species in the Mediterranean Sea, with a general biogeographical displacement towards the North European coasts. This will help anticipating changes in future catch potential in a warmer world, which is expected to have substantial economic consequences for Mediterranean fisheries.

The effect of environmental change in structuring the phytoplankton communities of the coastal waters of the Eastern English Channel was investigated by applying a trait-based approach on two decades (1996-2019) of monitoring on diatoms and Phaeocystis. We show that phytoplankton species richness in an unbalanced nutrient supply context was influenced by wind-driven processes, ecological specialization for dissolved inorganic phosphorous, temporal niche differentiation, and a competition-defense and/or a growth-defense trade-off, a coexistence mechanism where weak competitors (i.e., slower growing) are better protected against predation. Under the influence of both environmental perturbations (e.g., wind-driven processes, freshwater influence, unbalanced nutrient levels) and biotic interactions (e.g., competition, predation, facilitation), phytoplankton species exhibited specific survival strategies such as investment on growth, adaptation and tolerance of species to environmental stresses, silicification and resource specialization. These strategies have led to more speciose communities, higher productivity, functional redundancy and stability in the last decade. Our results revealed that the unbalanced nutrient reduction facilitated Phaeocystis blooms and that anthropogenic climate warming and nitrate reduction may threaten the diatom communities of the eastern English Channel in a near future. Our results provide strong support for biogeographical historical and niche-based processes in structuring the phytoplankton community in this temperate region. The variety of species responses that we characterized in this region may help to better understand future changes in pelagic ecosystems, and can serve as a basis to consider functional approaches for future ecosystem management.

Abstract Marine phytoplankton accounts for approximately 50% of all photosynthesis on Earth, underpins the marine food chain and plays a central role in the Earth’s biogeochemical cycles and climate. In situ measurements of ocean transparency can be used to estimate phytoplankton biomass. The scale and challenging conditions of the ocean make it a difficult environment for in situ studies, however. Here, we show that citizen scientists (seafarers) using a simple white Secchi Disk can collect ocean transparency data to complement formal scientific efforts using similar equipment. Citizen scientist data can therefore help understand current climate-driven changes in phytoplankton biomass at a global scale.

Planktonic Foraminifera are ubiquitous marine protozoa inhabiting the upper ocean. During life, they secrete calcareous shells, which accumulate in marine sediments, providing a geological record of past spatial and temporal changes in their community structure. As a result, they provide the opportunity to analyze both current and historical patterns of species distribution and community turnover in this plankton group on a global scale. The FORCIS project aims to unlock this potential by synthesizing a comprehensive global database of abundance and diversity observations of living planktonic Foraminifera in the upper ocean over more than 100 years starting from 1910. The database will allow for unravelling the impact of multiple global-change stressors acting on planktonic Foraminifera in historical times, using an approach that combines statistical analysis of temporal diversity changes in response to environmental changes with numerical modeling of species response based on their ecological traits.

Abstract Although we understand how species evolve, we do not appreciate how this process has filled an empty world to create current patterns of biodiversity. Here, we conduct a numerical experiment to determine why biodiversity varies spatially on our planet. We show that spatial patterns of biodiversity are mathematically constrained and arise from the interaction between the species’ ecological niches and environmental variability that propagates to the community level. Our results allow us to explain key biological observations such as (a) latitudinal biodiversity gradients (LBGs) and especially why oceanic LBGs primarily peak at midlatitudes while terrestrial LBGs generally exhibit a maximum at the equator, (b) the greater biodiversity on land even though life first evolved in the sea, (c) the greater species richness at the seabed than at the sea surface, and (d) the higher neritic (i.e., species occurring in areas with a bathymetry lower than 200 m) than oceanic (i.e., species occurring in areas with a bathymetry higher than 200 m) biodiversity. Our results suggest that a mathematical constraint originating from a fundamental ecological interaction, that is, the niche–environment interaction, fixes the number of species that can establish regionally by speciation or migration.

Although eutrophication induced by anthropogenic nutrient enrichment is a driver of shifts in community composition and eventually a threat to marine biodiversity, the causes and consequences on ecosystem functioning remain greatly unknown. In this study, by applying a trait-based approach and measuring niche breadth of diatoms and copepods, the drivers and underlying mechanisms of the seasonal species succession of these ecological communities in a coastal system dominated in spring by Phaeocystis blooms were explored. It is suggested that the seasonal succession of diatoms and copepods is the result of several trade-offs among functional traits that are controlled by the seasonal abiotic and biotic pressure encountered by the plankton communities. The results of this study highlight that a trade-off between competition and predator, i.e., weak competitors are better protected against predation, plays an important role in promoting plankton species richness and triggers the Phaeocystis bloom. As often observed in eutrophicated ecosystems, only the biotic homogenization of the copepod community and the shift in the diet of copepods toward Phaeocystis detrital materials have been detected during the Phaeocystis bloom. The diatom and copepod communities respond synchronously to fluctuating resources and biotic conditions by successively selecting species with specific traits. This study confirms the key role of competition and predation in controlling annual plankton succession.

The oceans' phytoplankton that underpin the marine food chain appear to be changing in abundance due to global climate change. Here, we compare the first four years of data from a citizen science ocean transparency study, conducted by seafarers using home-made Secchi Disks and a free Smartphone application called Secchi, with contemporaneous satellite ocean colour measurements. Our results show seafarers collect useful Secchi Disk measurements of ocean transparency that could help future assessments of climate-induced changes in the phytoplankton when used to extend historical Secchi Disk data.

Against the backdrop of warming of the Northern Hemisphere it has recently been acknowledged that North Atlantic temperature changes undergo considerable variability over multidecadal periods. The leading component of natural low-frequency temperature variability has been termed the Atlantic Multidecadal Oscillation (AMO). Presently, correlative studies on the biological impact of the AMO on marine ecosystems over the duration of a whole AMO cycle (∼60 years) is largely unknown due to the rarity of continuously sustained biological observations at the same time period. To test whether there is multidecadal cyclic behaviour in biological time-series in the North Atlantic we used one of the world's longest continuously sustained marine biological time-series in oceanic waters, long-term fisheries data and historical records over the last century and beyond. Our findings suggest that the AMO is far from a trivial presence against the backdrop of continued temperature warming in the North Atlantic and accounts for the second most important macro-trend in North Atlantic plankton records; responsible for habitat switching (abrupt ecosystem/regime shifts) over multidecadal scales and influences the fortunes of various fisheries over many centuries.

Abstract Many methods have been proposed to model the spatial distribution of a species. While some methods have been specifically designed for this purpose, others are well‐known statistical tools that can be used in many scientific fields. In this paper, I propose a new ecological niche model, called the Environmental String Model (ESM), that is based on the concept of environmental string, which is defined as being a combination of environmental variables, with as many nodes as environmental variables. There are two types of environmental strings: (1) the abundance‐known string and (2) the abundance‐unknown string (or target string) for which an estimation of abundance is searched. The novelty of the model is that it assesses the abundance associated with a target string from nearby abundance‐known strings, which preserve the local multidimensional relationships with the target string. The model does not provide an abundance estimate in the absence of data from a similar environment and it can therefore deal with truncated spatial distributions or niches. It is tested in the North Atlantic Ocean on two key copepod species, Calanus finmarchicus and Calanus helgolandicus, which have been monitored by the Continuous Plankton Recorder (CPR) survey for decades. I investigate the influence of variables on model performance. I show that the model reconstructs the mean spatial distribution and seasonal fluctuations in both Calanus well. When compared with generalized linear models (GLMs), generalized additive models (GAMs) and generalized regression neural network (GRNN), the ESM gives the best performance. I propose a number of indicators to evaluate the robustness of estimated abundance in space and time and show how the model may be extended to presence/absence or presence‐only data. I think that the ESM could be used to fill gaps in any sampling program such as the CPR survey and many satellite databases (e.g., ocean color and photosynthetically active radiation).

The FishClim model is presented, which quantifies the influence of fishing and climate on fisheries in order to develop strategies for sustainable, long-term, fish stock management.

Abstract Plankton biodiversity is a key component of marine pelagic ecosystems. They are at the base of the food web, control the productivity of marine ecosystems, and provide many provisioning and regulating ecological services. It is therefore important to understand how plankton are organized in both space and time. Here, we use data of varying taxonomic resolution, collected by the Continuous Plankton Recorder (CPR) survey, to map phytoplankton and zooplankton biodiversity in the North Atlantic and its adjacent seas. We then decompose biodiversity into 24 species assemblages and investigate their spatial distribution using ecological units and ecoregions recently proposed. Finally, we propose a descriptive method, which we call the environmental chromatogram, to characterize the environmental signature of each plankton assemblage. The method is based on a graphic that identifies where species of an assemblage aggregate along an environmental gradient composed of multiple ecological dimensions. The decomposition of the biodiversity into species assemblages allows us to show (a) that most marine regions of the North Atlantic are composed of coenoclines (i.e., gradients of biocoenoses or communities) and (b) that the overlapping spatial distribution of assemblages is the result of their environmental signatures. It follows that neither the ecoregions nor the ecological units identified in the North Atlantic are characterized by a unique assemblage but instead by a mosaic of assemblages that overlap in many places.

Abstract Heatwaves have increased in intensity, duration and frequency over the last decades due to climate change. Intertidal species, living in a highly variable environment, are likely to be exposed to such heatwaves since they can be emerged for more than 6 h during a tidal cycle. Little is known, however, on how temperature affects species traits (e.g. locomotion and behaviour) of slow-moving organisms such as benthic foraminifera (single-celled protists), which abound in marine sediments. Here, we examine how temperature influences motion-behaviour and metabolic traits of the dominant temperate foraminifera Haynesina germanica by exposing individuals to usual (6, 12, 18, 24, 30 °C) and extreme (high; i.e. 32, 34, 36 °C) temperature regimes. Our results show that individuals reduced their activity by up to 80% under high temperature regimes whereas they remained active under the temperatures they usually experience in the field. When exposed to a hyper-thermic stress (i.e. 36 °C), all individuals remained burrowed and the photosynthetic activity of their sequestered chloroplasts significantly decreased. Recovery experiments subsequently revealed that individuals initially exposed to a high thermal regime partially recovered when the hyper-thermic stress ceased. H. germanica contribution to surface sediment reworking substantially diminished from 10 mm3 indiv−1 day−1 (usual temperature) to 0 mm3 indiv−1 day−1 when individuals were exposed to high temperature regimes (i.e. above 32 °C). Given their role in sediment reworking and organic matter remineralisation, our results suggest that heatwaves may have profound long-lasting effects on the functioning of intertidal muddy ecosystems and some key biogeochemical cycles.

Abstract The ecological niche sensu Hutchinson is defined as the set of environmental conditions allowing a species to grow, maintain, and reproduce. This conception of the niche, which is assimilated to a p‐dimensional hypervolume, with p representing all environmental variables, has been widely applied in ecology. However, displaying the niche hypervolume has proved challenging when more than three environmental dimensions are considered simultaneously. We propose a simple method (implemented in the specieschrom R package) that displays the full multidimensionality of the ecological niche of a species into a two‐dimensional space by means of a graphic we call species chromatogram. This method gives a graphical summary of the niche by representing together abundance gradients with respect to all environmental variables. A chromatogram enables niche optimums and breaths to be rapidly quantified, and when several chromatograms are examined (one per species), rapid comparisons can be made. From our chromatograms, we proposed a procedure that quantifies niche optimum and breadth as well as niche overlapping (index D) and the identification of the most discriminant combination of environmental variables. We apply these analyses on eight planktonic species collected by the Continuous Plankton Recorder (CPR) survey in the North Atlantic Ocean using 10 environmental variables. We display their full multidimensional niches and quantify their niche optimums and breadths along each dimension. We also compare our index D with other indices by means of hypervolume and dynRB R packages. By catching the full complexity of the niche, species chromatograms allow many different niche properties to be rapidly assessed and compared among species from niche optimums and breadths to the identification of the most relevant environmental parameters and the degree of niche overlapping among species. Species chromatograms may be seen as species’ fingerprint and may also allow a better identification of the mechanisms involved in species assembly.

Species richness is greater in places where the number of potential niches is high. Consequently, the niche may be fundamental for understanding the arrangement of life and especially, the establishment and maintenance of the well-known Latitudinal Biodiversity Gradient (LBG). However, not all potential niches may be occupied fully in a habitat, as measured by niche vacancy/saturation. Here, we theoretically reconstruct oceanic biodiversity and analyse modeled and observed data together to examine patterns in niche saturation (i.e. the ratio between observed and theoretical biodiversity of a given taxon) for several taxonomic groups. Our results led us to hypothesize that the arrangement of marine life is constrained by the distribution of the maximal number of species' niches available, which represents a fundamental mathematical limit to the number of species that can co-exist locally. We liken this arrangement to a type of chessboard where each square on the board is a geographic area, itself comprising a distinct number of sub-squares (species' niches). Each sub-square on the chessboard can accept a unique species of a given ecological guild, whose occurrence is determined by speciation/extinction. Because of the interaction between the thermal niche and changes in temperature, our study shows that the chessboard has more sub-squares at mid-latitudes and we suggest that many clades should exhibit a LBG because their probability of emergence should be higher in the tropics where more niches are available. Our work reveals that each taxonomic group has its own unique chessboard and that global niche saturation increases when organismal complexity decreases. As a result, the mathematical influence of the chessboard is likely to be more prominent for taxonomic groups with low (e.g. plankton) than great (e.g. mammals) biocomplexity. Our study therefore reveals the complex interplay between a fundamental mathematical constraint on biodiversity resulting from the interaction between the species' ecological niche and fluctuations in the environmental regime (here, temperature), which has a predictable component and a stochastic-like biological influence (diversification rates, origination and clade age) that may alter or blur the former.

Over the past years, a large number of new approaches in the domain of ocean-color have been developed, leading to a variety of innovative descriptors for phytoplankton communities. One of these methods, named PHYSAT, currently allows for the qualitative detection of five main phytoplankton groups from ocean-color measurements. Even though PHYSAT products are widely used in various applications and projects, the approach is limited by the fact it identifies only dominant phytoplankton groups. This current limitation is due to the use of biomarker pigment ratios for establishing empirical relationships between in-situ information and specific ocean-color radiance anomalies in open ocean waters. However, theoretical explanations of PHYSAT suggests that it could be possible to detect more than dominance cases but move more toward phytoplanktonic assemblage detection. Thus, to evaluate the potential of PHYSAT for the detection of phytoplankton assemblages, we took advantage of the Continuous Plankton Recorder (CPR) survey, collected in both the English Channel and the North Sea. The available CPR dataset contains information on diatom abundance in two large areas of the North Sea for the period 1998-2010. Using this unique dataset, recurrent diatom assemblages were retrieved based on classification of CPR samples. Six diatom assemblages were identified in-situ, each having indicators taxa or species. Once this first step was completed, the in-situ analysis was used to empirically associate the diatom assemblages with specific PHYSAT spectral anomalies. This step was facilitated by the use of previous classifications of regional radiance anomalies in terms of shape and amplitude, coupled with phenological tools. Through a matchup exercise, three CPR assemblages were associated with specific radiance anomalies. The maps of detection of these specific radiances anomalies are in close agreement with current in-situ ecological knowledge.

Global change has become a major driving force of both terrestrial and marine systems. Located at the interface between these two realms, estuarine ecosystems are probably the place where both direct and indirect effects of human activities conspire together to affect biodiversity from phytoplankton to top predators. Among European estuarine systems, the Gironde is the largest estuary of Western Europe and many studies have provided evidence that it has been affected by a variety of anthropogenic stressors such as thermal and chemical pollution, physical alterations and exploitation, especially for maritime traffic. In such a context, species introduction is also a current major issue with the establishment of strong competitive species that could lead to ecosystem reorganization with potential decrease or even disappearance of native species. In the Gironde estuary, this hypothesis was proposed for the invasive shrimp species Palaemon macrodactylus as a decrease in the native species abundance was observed at the same time. Although species introduction often takes place via ballast water, the influence of climate-driven changes on the establishment of new species remains a key issue. The calanoid copepod Acartia tonsa, observed in the Gironde estuary for the first time in 1983, have since colonized most part of the estuary, reaching a level of abundance comparable to the dominant native species Eurytemora affinis. In this study, using both the concept of the ecological niche sensu Hutchinson (fundamental and realized niches) and statistical models, we reveal that the dynamics of the colonization of A. tonsa was facilitated by environmental conditions that have become closer to its environmental optimum with respect to temperature and salinity.

Kelp ecosystems form widespread underwater forests playing a major role in structuring the biodiversity at a regional scale. Some seaweeds such as Laminaria digitata are also economically important, being exploited for their alginate and iodine content. Although some studies have shown that kelp ecosystems are regressing and that multiple causes are likely to be at the origin of the disappearance of certain populations, the extent to which global climate change may play a role remains speculative. Here we show that many populations of L. digitata along European coasts are on the verge of local extinction due to a climate-caused increase in sea temperature. By modeling the spatial distribution of the seaweed, we evaluate the possible implications of global climate change for the geographical patterns of the species using temperature data from the Coupled Model Intercomparison Project phase 5 (CMIP5). Projections of the future range of L. digitata throughout the 21st century show large shifts in the suitable habitat of the kelp and a northward retreat of the southern limit of its current geographic distribution from France to Danish coasts and the southern regions of the United Kingdom. However, these projections depend on the intensity of warming. A medium to high warming is expected to lead to the extirpation of the species as early as the first half of the 21st century and there is high confidence that regional extinction will spread northwards by the end of this century. These changes are likely to cause the decline of species whose life cycle is closely dependent upon L. digitata and lead to the establishment of new ecosystems with lower ecological and economic values.

One of the major challenges to understanding population changes in ecology for assessment purposes is the difficulty in evaluating the suitability of an area for a given species. Here we used a new simple approach able to faithfully predict through time the abundance of two key zooplanktonic species by focusing on the relationship between the species' environmental preferences and their observed abundances. The approach is applied to the marine copepods Calanus finmarchicus and C. helgolandicus as a case study characterising the multidecadal dynamics of the North Sea ecosystem. We removed all North Sea data from the Continuous Plankton Recorder (CPR) dataset and described for both species a simplified ecological niche using Sea Surface Temperature (SST) and CPR Phytoplankton Colour Index (PCI). We then modelled the dynamics of each species by associating the North Sea's environmental parameters to the species' ecological niches, thus creating a method to assess the suitability of this area. By using both C. finmarchicus and C. helgolandicus as indicators, the procedure reproduces the documented switches from cold to warm temperate states observed in the North Sea.

Climate change is having a discernible effect on many biological and ecological processes. Among observed changes, modifications in bird phenology have been widely documented. However, most studies have interpreted phenological shifts as gradual biological adjustments in response to the alteration of the thermal regime. Here we analysed a long-term dataset (1980-2010) of short-distance migratory raptors in five European regions. We revealed that the responses of these birds to climate-induced changes in autumn temperatures are abrupt and synchronous at a continental scale. We found that when the temperatures increased, birds delayed their mean passage date of autumn migration. Such delay, in addition to an earlier spring migration, suggests that a significant warming may induce an extension of the breeding-area residence time of migratory raptors, which may eventually lead to residency.

Marine environments are greatly affected by climate change, and understanding how this perturbation affects marine vertebrates is a major issue. In this context, it is essential to identify the environmental drivers of animal distribution. Here, we focused on the little auk (Alle alle), one of the world's most numerous seabirds and a major component in Arctic food webs. Using a multidisciplinary approach, we show how little auks adopt specific migratory strategies and balance environmental constraints to optimize their energy budgets. Miniature electronic loggers indicate that after breeding, birds from East Greenland migrate >2000 km to overwinter in a restricted area off Newfoundland. Synoptic data available from the Continuous Plankton Recorder (CPR) indicate that this region harbours some of the highest densities of the copepod Calanus finmarchicus found in the North Atlantic during winter. Examination of large-scale climatic and oceanographic data suggests that little auks favour patches of high copepod abundance in areas where air temperature ranges from 0°C to 5°C. These results greatly advance our understanding of animal responses to extreme environmental constraints, and highlight that information on habitat preference is key to identifying critical areas for marine conservation.

Climate change is unambiguous and its effects are clearly detected in all functional units of the Earth System. This paper presents analyses of sea surface temperature and show that climate change is affecting both biological and ecological systems of the world and most specifically the North Atlantic and its adjacent seas. Changes are seen from phytoplankton to zooplankton to fish and are modifying the dominance of species and the structure, the diversity and the functioning of marine ecosystems. Changes also range from biogeographical to phenological shifts and have involved in some regions of the Atlantic abrupt ecosystem shifts also called regime shifts. These alterations reflect the response of ecosystems to a warmer temperature regime. Mechanisms are complex because they are nonlinear exhibiting tipping points and varying in space and time. Sensitivity of organisms to temperature changes is high, implicating that a small modification in the temperature regime can have sustained ecosystem effects. It is urgent to develop monitoring systems, indicators and both statistical and mathematical tools to detect, better understand and anticipate the alterations of both biological and ecological systems that may be triggered by global climate change.