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"Tittensor, Derek P."
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How Many Species Are There on Earth and in the Ocean?
The diversity of life is one of the most striking aspects of our planet; hence knowing how many species inhabit Earth is among the most fundamental questions in science. Yet the answer to this question remains enigmatic, as efforts to sample the world's biodiversity to date have been limited and thus have precluded direct quantification of global species richness, and because indirect estimates rely on assumptions that have proven highly controversial. Here we show that the higher taxonomic classification of species (i.e., the assignment of species to phylum, class, order, family, and genus) follows a consistent and predictable pattern from which the total number of species in a taxonomic group can be estimated. This approach was validated against well-known taxa, and when applied to all domains of life, it predicts ~8.7 million (± 1.3 million SE) eukaryotic species globally, of which ~2.2 million (± 0.18 million SE) are marine. In spite of 250 years of taxonomic classification and over 1.2 million species already catalogued in a central database, our results suggest that some 86% of existing species on Earth and 91% of species in the ocean still await description. Renewed interest in further exploration and taxonomy is required if this significant gap in our knowledge of life on Earth is to be closed.
Journal Article
Current and Future Patterns of Global Marine Mammal Biodiversity
Quantifying the spatial distribution of taxa is an important prerequisite for the preservation of biodiversity, and can provide a baseline against which to measure the impacts of climate change. Here we analyse patterns of marine mammal species richness based on predictions of global distributional ranges for 115 species, including all extant pinnipeds and cetaceans. We used an environmental suitability model specifically designed to address the paucity of distributional data for many marine mammal species. We generated richness patterns by overlaying predicted distributions for all species; these were then validated against sightings data from dedicated long-term surveys in the Eastern Tropical Pacific, the Northeast Atlantic and the Southern Ocean. Model outputs correlated well with empirically observed patterns of biodiversity in all three survey regions. Marine mammal richness was predicted to be highest in temperate waters of both hemispheres with distinct hotspots around New Zealand, Japan, Baja California, the Galapagos Islands, the Southeast Pacific, and the Southern Ocean. We then applied our model to explore potential changes in biodiversity under future perturbations of environmental conditions. Forward projections of biodiversity using an intermediate Intergovernmental Panel for Climate Change (IPCC) temperature scenario predicted that projected ocean warming and changes in sea ice cover until 2050 may have moderate effects on the spatial patterns of marine mammal richness. Increases in cetacean richness were predicted above 40° latitude in both hemispheres, while decreases in both pinniped and cetacean richness were expected at lower latitudes. Our results show how species distribution models can be applied to explore broad patterns of marine biodiversity worldwide for taxa for which limited distributional data are available.
Journal Article
Assessing changing baleen whale distributions and reported incidents relative to vessel activity in the Northwest Atlantic
Baleen whales are among the largest marine megafauna, and while mostly well-protected from direct exploitation, they are increasingly affected by vessel traffic, interactions with fisheries, and climate change. Adverse interactions, notably vessel strikes and fishing gear entanglement, often result in distress, injury, or death for these animals. In Atlantic Canadian waters, such negative interactions or ‘incidents’ are consistently reported to marine animal response organizations but have not yet been analyzed relative to the spatial distribution of whales and vessels. Using a database of 483,003 whale sightings, 1,110 incident reports, and 82 million hours of maritime vessel activity, we conducted a spatiotemporal vulnerability analysis for all six baleen whale species occurring in the Northwest Atlantic Ocean by developing an ensemble of habitat-suitability models. The relative spatial risk of vessel-induced incidents was assessed for present (1985–2015) and projected near-future (2035–2055) distributions of baleen whales. Areas of high habitat suitability for multiple baleen whale species were intrinsically linked to sea surface temperature and salinity, with multispecies hotspots identified in the Bay of Fundy, Scotian Shelf, Laurentian Channel, Flemish Cap, and Gulf of St. Lawrence. Present-day model projections were independently evaluated using a separate database of acoustic detections and found to align well. Regions of high relative incident risk were projected close to densely inhabited regions, principal maritime routes, and major fishing grounds, in general coinciding with reported incident hotspots. While some high-risk regions already benefit from mitigation strategies aimed at protecting North Atlantic Right Whales, our analysis highlights the importance of considering risks to multiple species, both in the present day and under continued environmental change.
Journal Article
Emergent Global Patterns of Ecosystem Structure and Function from a Mechanistic General Ecosystem Model
Anthropogenic activities are causing widespread degradation of ecosystems worldwide, threatening the ecosystem services upon which all human life depends. Improved understanding of this degradation is urgently needed to improve avoidance and mitigation measures. One tool to assist these efforts is predictive models of ecosystem structure and function that are mechanistic: based on fundamental ecological principles. Here we present the first mechanistic General Ecosystem Model (GEM) of ecosystem structure and function that is both global and applies in all terrestrial and marine environments. Functional forms and parameter values were derived from the theoretical and empirical literature where possible. Simulations of the fate of all organisms with body masses between 10 µg and 150,000 kg (a range of 14 orders of magnitude) across the globe led to emergent properties at individual (e.g., growth rate), community (e.g., biomass turnover rates), ecosystem (e.g., trophic pyramids), and macroecological scales (e.g., global patterns of trophic structure) that are in general agreement with current data and theory. These properties emerged from our encoding of the biology of, and interactions among, individual organisms without any direct constraints on the properties themselves. Our results indicate that ecologists have gathered sufficient information to begin to build realistic, global, and mechanistic models of ecosystems, capable of predicting a diverse range of ecosystem properties and their response to human pressures.
Journal Article
Range contraction in large pelagic predators
Large reductions in the abundance of exploited land predators have led to significant range contractions for those species. This pattern can be formalized as the range–abundance relationship, a general macroecological pattern that has important implications for the conservation of threatened species. Here we ask whether similar responses may have occurred in highly mobile pelagic predators, specifically 13 species of tuna and billfish. We analyzed two multidecadal global data sets on the spatial distribution of catches and fishing effort targeting these species and compared these with available abundance time series from stock assessments. We calculated the effort needed to reliably detect the presence of a species and then computed observed range sizes in each decade from 1960 to 2000. Results suggest significant range contractions in 9 of the 13 species considered here (between 2% and 46% loss of observed range) and significant range expansions in two species (11–29% increase). Species that have undergone the largest declines in abundance and are of particular conservation concern tended to show the largest range contractions. These include all three species of bluefin tuna and several marlin species. In contrast, skipjack tuna, which may have increased its abundance in the Pacific, has also expanded its range size. These results mirror patterns described for many land predators, despite considerable differences in habitat, mobility, and dispersal, and imply ecological extirpation of heavily exploited species across parts of their range.
Journal Article
Future ocean biomass losses may widen socioeconomic equity gaps
Future climate impacts and their consequences are increasingly being explored using multi-model ensembles that average across individual model projections. Here we develop a statistical framework that integrates projections from coupled ecosystem and earth-system models to evaluate significance and uncertainty in marine animal biomass changes over the 21
st
century in relation to socioeconomic indicators at national to global scales. Significant biomass changes are projected in 40%–57% of the global ocean, with 68%–84% of these areas exhibiting declining trends under low and high emission scenarios, respectively. Given unabated emissions, maritime nations with poor socioeconomic statuses such as low nutrition, wealth, and ocean health will experience the greatest projected losses. These findings suggest that climate-driven biomass changes will widen existing equity gaps and disproportionally affect populations that contributed least to global CO
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emissions. However, our analysis also suggests that such deleterious outcomes are largely preventable by achieving negative emissions (RCP 2.6).
Numerous marine ecosystem models are used to project animal biomass over time but integrating them can be challenging. Here the authors develop a test for statistical significance in multi-model ensemble trends, and thus relate future biomass trends to current patterns of ecological and socioeconomic status.
Journal Article
Global patterns and predictors of marine biodiversity across taxa
Marine biodiversity survey
Building on the decade-long Census of Marine Life project, a new global analysis of data on the distribution of 11,567 marine species from 13 different taxonomic groups, including zooplankton, plants, invertebrates, fishes and mammals, reveals temperature as a main correlate of biodiversity across the various taxa, with generally higher diversity in warmer waters. Two further distinct trends emerge: coastal species show maximum diversity in the Western Pacific, whereas ocean-dwelling species peak in the mid-latitudes. This contrasts with the situation for terrestrial species, where biodiversity peaks in the tropics.
Using large-scale data sets, these authors present a new assessment of global marine species diversity and its correlation with environmental and spatial parameters.
Global patterns of species richness and their structuring forces have fascinated biologists since Darwin
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and provide critical context for contemporary studies in ecology, evolution and conservation. Anthropogenic impacts and the need for systematic conservation planning have further motivated the analysis of diversity patterns and processes at regional to global scales
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. Whereas land diversity patterns and their predictors are known for numerous taxa
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, our understanding of global marine diversity has been more limited, with recent findings revealing some striking contrasts to widely held terrestrial paradigms
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. Here we examine global patterns and predictors of species richness across 13 major species groups ranging from zooplankton to marine mammals. Two major patterns emerged: coastal species showed maximum diversity in the Western Pacific, whereas oceanic groups consistently peaked across broad mid-latitudinal bands in all oceans. Spatial regression analyses revealed sea surface temperature as the only environmental predictor highly related to diversity across all 13 taxa. Habitat availability and historical factors were also important for coastal species, whereas other predictors had less significance. Areas of high species richness were disproportionately concentrated in regions with medium or higher human impacts. Our findings indicate a fundamental role of temperature or kinetic energy in structuring cross-taxon marine biodiversity, and indicate that changes in ocean temperature, in conjunction with other human impacts, may ultimately rearrange the global distribution of life in the ocean.
Journal Article
Past and future decline of tropical pelagic biodiversity
A major research question concerning global pelagic biodiversity remains unanswered: when did the apparent tropical biodiversity depression (i.e., bimodality of latitudinal diversity gradient [LDG]) begin? The bimodal LDG may be a consequence of recent ocean warming or of deep-time evolutionary speciation and extinction processes. Using rich fossil datasets of planktonic foraminifers, we show here that a unimodal (or only weakly bimodal) diversity gradient, with a plateau in the tropics, occurred during the last ice age and has since then developed into a bimodal gradient through species distribution shifts driven by postglacial ocean warming. The bimodal LDG likely emerged before the Anthropocene and industrialization, and perhaps ∼15,000 y ago, indicating a strong environmental control of tropical diversity even before the start of anthropogenic warming. However, our model projections suggest that future anthropogenic warming further diminishes tropical pelagic diversity to a level not seen in millions of years.
Journal Article
AquaX: An enhanced and revised AquaMaps framework to model marine species distributions and biodiversity
Marine biodiversity underpins ecosystem health and is critical for the provision of essential ecological services. Global efforts to mitigate biodiversity loss are underway but require comprehensive knowledge on the biogeography of species to be effective. However, key challenges limit comprehensive mapping of species distributions, including the ecosystem complexity and difficulty of sampling the marine realm. Global initiatives such as AquaMaps pioneered large-scale marine species mapping using species distribution models or ecological niche models and provided the knowledge base for effective marine conservation and management. Recently, methodological and data advances have enabled a more modern and robust approach that enables higher resolution outputs more suited to conservation applications at all scales. Building on AquaMaps, we developed a next-generation marine species habitat suitability modelling platform called AquaX, providing a suite of advances that include an ensemble of ten machine learning algorithms, enabling spatial uncertainty assessments, validation indices, and ecological niche representation at a ten-fold improved spatial resolution of 0.05°. Furthermore, AquaX integrates (i) accepted taxonomy from the World Register of Marine Species, (ii) species-specific ecological, physiological, and biogeographical information (D3-Ocean system), (iii) updated occurrence records validated through expert input, and (iv) refined species range maps using expert knowledge and biogeographical divisions. AquaX also projects species’ habitat suitability for both present and future conditions based on two time periods and three climate scenarios. This work provides species range maps for numerous species compared to previously available datasets and improves the accurate use of observational data. The approaches described here improve predictive accuracy at scales more relevant to marine biodiversity conservation and offer an openly accessible tool to support marine biodiversity research and conservation planning under accelerating environmental change. AquaX represents an important step forward in species distribution modeling, enabling researchers and policymakers to better understand marine biodiversity patterns and develop more effective conservation strategies.
Journal Article