A niche-based theory of island biogeography.

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.

Ocean climate and hydrodynamics drive decadal shifts in Northeast Atlantic dinoflagellates.

The abundance of large marine dinoflagellates has declined in the North Sea since 1958. Although hypotheses have been proposed to explain this diminution (increasing temperature and wind), the mechanisms behind this pattern have thus far remained elusive. In this article, we study the long-term changes in dinoflagellate biomass and biodiversity in relation to hydro-climatic conditions and circulation within the North Atlantic. Our results show that the decline in biomass has paralleled an increase in biodiversity caused by a temperature-induced northward movement of subtropical taxa along the European shelf-edge, and facilitated by changes in oceanic circulation (subpolar gyre contraction). However, major changes in North Atlantic hydrodynamics in the 2010s (subpolar gyre expansion and low-salinity anomaly) stopped this movement, which triggered a biodiversity collapse in the North Sea. Further, North Sea dinoflagellate biomass remained low because of warming. Our results, therefore, reveal that regional climate warming and changes in oceanic circulation strongly influenced shifts in dinoflagellate biomass and biodiversity.

Phytoplankton life strategies, phenological shifts and climate change in the North Atlantic Ocean from 1850 to 2100.

Significant phenological shifts induced by climate change are projected within the phytoplankton community. However, projections from current Earth System Models (ESMs) understandably rely on simplified community responses that do not consider evolutionary strategies manifested as various phenotypes and trait groups. Here, we use a species-based modelling approach, combined with large-scale plankton observations, to investigate past, contemporary and future phenological shifts in diatoms (grouped by their morphological traits) and dinoflagellates in three key areas of the North Atlantic Ocean (North Sea, North-East Atlantic and Labrador Sea) from 1850 to 2100. Our study reveals that the three phytoplanktonic groups exhibit coherent and different shifts in phenology and abundance throughout the North Atlantic Ocean. The seasonal duration of large flattened (i.e. oblate) diatoms is predicted to shrink and their abundance to decline, whereas the phenology of slow-sinking elongated (i.e. prolate) diatoms and of dinoflagellates is expected to expand and their abundance to rise, which may alter carbon export in this important sink region. The increase in prolates and dinoflagellates, two groups currently not considered in ESMs, may alleviate the negative influence of global climate change on oblates, which are responsible of massive peaks of biomass and carbon export in spring. We suggest that including prolates and dinoflagellates in models may improve our understanding of the influence of global climate change on the biological carbon cycle in the oceans.

Addressing the dichotomy of fishing and climate in fishery management with the FishClim model.

The relative influence of fishing and Climate-Induced Environmental Change (CIEC) on long-term fluctuations in exploited fish stocks has been controversial1-3 because separating their contributions is difficult for two reasons. Firstly, there is in general, no estimation of CIEC for a pre-fishing period and secondly, the assessment of the effects of fishing on stocks has taken place at the same time as CIEC4. Here, we describe a new model we have called FishClim that we apply to North Sea cod from 1963 to 2019 to estimate how fishing and CIEC interact and how they both may affect stocks in the future (2020-2100) using CMIP6 scenarios5. The FishClim model shows that both fishing and CIEC are intertwined and can either act synergistically (e.g. the 2000-2007 collapse) or antagonistically (e.g. second phase of the gadoid outburst). Failure to monitor CIEC, so that fisheries management immediately adjusts fishing effort in response to environmentally-driven shifts in stock productivity, will therefore create a deleterious response lag that may cause the stock to collapse. We found that during 1963-2019, although the effect of fishing and CIEC drivers fluctuated annually, the pooled influence of fishing and CIEC on the North Sea cod stock was nearly equal at ~55 and ~45%, respectively. Consequently, the application of FishClim, which quantifies precisely the respective influence of fishing and climate, will help to develop better strategies for sustainable, long-term, fish stock management.

How do plankton species coexist in an apparently unstructured environment?

In a paper entitled The paradox of the plankton, Hutchinson asked 'how it is possible for a number of species to coexist in a relatively isotropic or unstructured environment all competing for the same sorts of materials' (Hutchinson 1961 Am. Nat. 95, 137-145 (doi:10.1086/282171)). Particularly relevant for phytoplankton, this paradox was based on two implicit, and perhaps naive, postulates, i.e. (i) that all plankton species have similar requirements and (ii) that the marine environment is relatively homogeneous in space and time. A number of hypotheses, based on purely theoretical or experimental studies, have been proposed to solve this conundrum, ranging from spatio-temporal environmental heterogeneity to biotic chaotic variability. Here, we characterize the ecological niche of 117 plankton species belonging to three different taxonomic groups and show that all species have a niche sufficiently distinct to ensure coexistence in a structured marine environment. We also provide evidence that pelagic habitats are, unsurprisingly, more diverse in space and time than Hutchinson imagined, the marine environment being neither unstructured nor stable in space and time. We, therefore, conclude that the niche theory, and its corollary the principle of competitive exclusion, apply as much for the plankton as for other forms of life, be they terrestrial or marine.

The species chromatogram, a new graphical method to represent, characterize, and compare the ecological niches of different species.

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.

Citizens and scientists collect comparable oceanographic data: measurements of ocean transparency from the Secchi Disk study and science programmes.

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.

Plankton biogeography in the North Atlantic Ocean and its adjacent seas: Species assemblages and environmental signatures.

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.