The rise of a marine generalist predator and the fall of beta diversity.

Determining the importance of physical and biological drivers in shaping biodiversity in diverse ecosystems remains a global challenge. Advancements have been made towards this end in large marine ecosystems with several studies suggesting environmental forcing as the primary driver. However, both empirical and theoretical studies point to additional drivers of changes in diversity involving trophic interactions and, in particular, predation. Moreover, a more integrated but less common approach to the assessment of biodiversity changes involves analyses of spatial ß diversity, whereas most studies to date assess only changes in species richness (α diversity). Recent research has established that when cod, a dominant generalist predator, was overfished and collapsed in a northwest Atlantic food web, spatial ß diversity increased; that is, the spatial structure of the fish assemblage became increasingly heterogeneous. If cod were to recover, would this situation be reversible, given the inherent complexity and non-linear dynamics that typify such systems? A dramatic increase of cod in an ecologically similar large marine ecosystem may provide an answer. Here we show that spatial ß diversity of fish assemblages in the Barents Sea decreased with increasing cod abundance, while decadal scale changes in temperature did not play a significant role. These findings indicate a reversibility of the fish assemblage structure in response to changing levels of an apex predator and highlight the frequently overlooked importance of trophic interactions in determining large-scale biodiversity patterns. As increased cod abundance was largely driven by changes in fisheries management, our study also shows that management policies and practices, particularly those involving apex predators, can have a strong effect in shaping spatial diversity patterns, and one should not restrict the focus to effects of climate change alone.

Commonness and rarity in the marine biosphere.

Explaining patterns of commonness and rarity is fundamental for understanding and managing biodiversity. Consequently, a key test of biodiversity theory has been how well ecological models reproduce empirical distributions of species abundances. However, ecological models with very different assumptions can predict similar species abundance distributions, whereas models with similar assumptions may generate very different predictions. This complicates inferring processes driving community structure from model fits to data. Here, we use an approximation that captures common features of "neutral" biodiversity models--which assume ecological equivalence of species--to test whether neutrality is consistent with patterns of commonness and rarity in the marine biosphere. We do this by analyzing 1,185 species abundance distributions from 14 marine ecosystems ranging from intertidal habitats to abyssal depths, and from the tropics to polar regions. Neutrality performs substantially worse than a classical nonneutral alternative: empirical data consistently show greater heterogeneity of species abundances than expected under neutrality. Poor performance of neutral theory is driven by its consistent inability to capture the dominance of the communities' most-abundant species. Previous tests showing poor performance of a neutral model for a particular system often have been followed by controversy about whether an alternative formulation of neutral theory could explain the data after all. However, our approach focuses on common features of neutral models, revealing discrepancies with a broad range of empirical abundance distributions. These findings highlight the need for biodiversity theory in which ecological differences among species, such as niche differences and demographic trade-offs, play a central role.

Long-term environmental monitoring for assessment of change: measurement inconsistencies over time and potential solutions.

The importance of long-term environmental monitoring and research for detecting and understanding changes in ecosystems and human impacts on natural systems is widely acknowledged. Over the last decades, a number of critical components for successful long-term monitoring have been identified. One basic component is quality assurance/quality control protocols to ensure consistency and comparability of data. In Norway, the authorities require environmental monitoring of the impacts of the offshore petroleum industry on the Norwegian continental shelf, and in 1996, a large-scale regional environmental monitoring program was established. As a case study, we used a sub-set of data from this monitoring to explore concepts regarding best practices for long-term environmental monitoring. Specifically, we examined data from physical and chemical sediment samples and benthic macroinvertebrate assemblages from 11 stations from six sampling occasions during the period 1996-2011. Despite the established quality assessment and quality control protocols for this monitoring program, we identified several data challenges, such as missing values and outliers, discrepancies in variable and station names, changes in procedures without calibration, and different taxonomic resolution. Furthermore, we show that the use of different laboratories over time makes it difficult to draw conclusions with regard to some of the observed changes. We offer recommendations to facilitate comparison of data over time. We also present a new procedure to handle different taxonomic resolution, so valuable historical data is not discarded. These topics have a broader relevance and application than for our case study.

The role of time and species identities in spatial patterns of species richness and conservation.

Many conservation actions are justified on the basis of managing biodiversity. Biodiversity, in terms of species richness, is largely the product of rare species. This is problematic because the intensity of sampling needed to characterize communities and patterns of rarity or to justify the use of surrogates has biased sampling in favor of space over time. However, environmental fluctuations interacting with community dynamics lead to temporal variations in where and when species occur, potentially affecting conservation planning by generating uncertainty about results of species distribution modeling (including range determinations), selection of surrogates for biodiversity, and the proportion of biodiversity composed of rare species. To have confidence in the evidence base for conservation actions, one must consider whether temporal replication is necessary to produce broad inferences. Using approximately 20 years of macrofaunal data from tidal flats in 2 harbors, we explored variation in the identity of rare, common, restricted range, and widespread species over time and space. Over time, rare taxa were more likely to increase in abundance or occurrence than to remain rare or disappear and to exhibit temporal patterns in their occurrence. Space-time congruency in ranges (i.e., spatially widespread taxa were also temporally widespread) was observed only where samples were collected across an environmental gradient. Fifteen percent of the taxa in both harbors changed over time from having spatially restricted ranges to having widespread ranges. Our findings suggest that rare species can provide stability against environmental change, because the majority of species were not random transients, but that selection of biodiversity surrogates requires temporal validation. Rarity needs to be considered both spatially and temporally, as species that occur randomly over time are likely to play a different role in ecosystem functioning than those exhibiting temporal structure (e.g., seasonality). Moreover, temporal structure offers the opportunity to place management and conservation activities within windows of maximum opportunity.

The role of a dominant predator in shaping biodiversity over space and time in a marine ecosystem.

1. Exploitation of living marine resources has resulted in major changes to populations of targeted species and functional groups of large-bodied species in the ocean. However, the effects of overfishing and collapse of large top predators on the broad-scale biodiversity of oceanic ecosystems remain largely unexplored. 2. Populations of the Atlantic cod (Gadus morhua) were overfished and several collapsed in the early 1990s across Atlantic Canada, providing a unique opportunity to study potential ecosystem-level effects of the reduction of a dominant predator on fish biodiversity, and to identify how such effects might interact with other environmental factors, such as changes in climate, over time. 3. We combined causal modelling with model selection and multimodel inference to analyse 41 years of fishery-independent survey data (1970-2010) and quantify ecosystem-level effects of overfishing and climate variation on the biodiversity of fishes across a broad area (172 000 km(2) ) of the Scotian Shelf. 4. We found that alpha and beta diversity increased with decreases in cod occurrence; fish communities were less homogeneous and more variable in systems where cod no longer dominated. These effects were most pronounced in the colder north-eastern parts of the Scotian Shelf. 5. Our results provide strong evidence that intensive harvesting (and collapse) of marine apex predators can have large impacts on biodiversity, with far-reaching consequences for ecological stability across an entire ecosystem.

Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver-response relationships to develop conceptual models across ecosystems.

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed 'beta diversity') is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.

Resource-driven colonization by cod in a high Arctic food web.

Climate change is commonly associated with many species redistributions and the influence of other factors may be marginalized, especially in the rapidly warming Arctic.The Barents Sea, a high latitude large marine ecosystem in the Northeast Atlantic has experienced above-average temperatures since the mid-2000s with divergent bottom temperature trends at subregional scales.Concurrently, the Barents Sea stock of Atlantic cod Gadus morhua, one of the most important commercial fish stocks in the world, increased following a large reduction in fishing pressure and expanded north of 80°N.We examined the influence of food availability and temperature on cod expansion using a comprehensive data set on cod stomach fullness stratified by subregions characterized by divergent temperature trends. We then tested whether food availability, as indexed by cod stomach fullness, played a role in cod expansion in subregions that were warming, cooling, or showed no trend.The greatest increase in cod occupancy occurred in three northern subregions with contrasting temperature trends. Cod apparently benefited from initial high food availability in these regions that previously had few large-bodied fish predators.The stomach fullness in the northern subregions declined rapidly after a few years of high cod abundance, suggesting that the arrival of cod caused a top-down effect on the prey base. Prolonged cod residency in the northern Barents Sea is, therefore, not a certainty.

Multivariate dispersion as a measure of beta diversity.

Beta diversity can be defined as the variability in species composition among sampling units for a given area. We propose that it can be measured as the average dissimilarity from individual observation units to their group centroid in multivariate space, using an appropriate dissimilarity measure. Differences in beta diversity among different areas or groups of samples can be tested using this approach. The choice of transformation and dissimilarity measure has important consequences for interpreting results. For kelp holdfast assemblages from New Zealand, variation in species composition was greater in smaller holdfasts, while variation in relative abundances was greater in larger holdasts. Variation in community structure of Norwegian continental shelf macrobenthic fauna increased with increases in environmental heterogeneity, regardless of the measure used. We propose a new dissimilarity measure which allows the relative weight placed on changes in composition vs. abundance to be specified explicitly.

Spatio-temporal variability of richness estimators: coastal marine fish communities as examples.

We assessed the performance of two estimators of species richness, the Chao2 and the Coleman 'random placement curve'. Using a dataset of intertidal fish from the Norwegian Skagerrak coast, we found that Chao2 was effective for low sampling intensity, often reaching asymptotic values for few samples, but for higher sampling intensity the performance deteriorated. For large samples, the Coleman random placement curve was more effective than the Chao2 estimates when comparing spatio-temporal patterns of species richness. Spatial patterns were clearly and consistently identified by both methods, whereas the coastal fish communities displayed too much variability in the early summer for any sensible measure of temporal patterns of fish-species richness to be made. To control for spurious results due to systematic differences in mean abundance of the samples the analyses were performed also on data standardised by the number of individuals in the samples, without any significant change in the results. We conclude that modest sampling effort is sufficient to characterise spatial patterns of coastal fish-species richness, while a detailed and high-precision description of seasonal patterns could not be obtained with any reasonable sampling effort.