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1.  Emergent Global Patterns of Ecosystem Structure and Function from a Mechanistic General Ecosystem Model 
PLoS Biology  2014;12(4):e1001841.
This paper presents the first mathematical model that attempts to represent the biology and behavior of all individual organisms globally, taking us a step closer to holistic ecological and conservation science founded on mechanistic predictions.
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.
Author Summary
Ecosystems across the world are being rapidly degraded. This threatens their provision of natural goods and services, upon which all life depends. To be able to reduce—and one day reverse—this damage, we need to be able to predict the effects of human actions on ecosystems. Here, we present the first example of a General Ecosystem Model (GEM)—called the Madingley Model—a novel class of computational model that can be applied to any ecosystem, marine or terrestrial, and can be simulated at any spatial scale from local up to global. It covers almost all organisms in ecosystems, from the smallest to the largest, encoding the underlying biology and behaviour of individual organisms to capture the interactions between them and with the environment, to model the fate of each individual organism, and to make predictions about ecosystem structure and function. Predictions made by the Madingley Model broadly resemble what we observe in real-world ecosystems across scales from individuals through to communities, ecosystems, and the world as a whole. Our results show that ecologists can now begin modelling all nonhuman life on earth, and we suggest that this type of approach may hold promise for predicting the ecological implications of different future trajectories of human activity on our shared planet.
PMCID: PMC3995663  PMID: 24756001
2.  Species–energy relationships in deep-sea molluscs 
Biology Letters  2011;7(5):718-722.
Consensus is growing among ecologists that energy and the factors influencing its utilization can play overarching roles in regulating large-scale patterns of biodiversity. The deep sea—the world's largest ecosystem—has simplified energetic inputs and thus provides an excellent opportunity to study how these processes structure spatial diversity patterns. Two factors influencing energy availability and use are chemical (productive) and thermal energy, here represented as seafloor particulate organic carbon (POC) flux and temperature. We related regional patterns of benthic molluscan diversity in the North Atlantic to these factors, to conduct an explicit test of species–energy relationships in the modern day fauna of the deep ocean. Spatial regression analyses in a model-averaging framework indicated that POC flux had a substantially higher relative importance than temperature for both gastropods and protobranch bivalves, although high correlations between variables prevented definitive interpretation. This contrasts with recent research on temporal variation in fossil diversity from deep-sea cores, where temperature is generally a more significant predictor. These differences may reflect the scales of time and space at which productivity and temperature operate, or differences in body size; but both lines of evidence implicate processes influencing energy utilization as major determinants of deep-sea species diversity.
PMCID: PMC3169037  PMID: 21429909
species–energy; productivity; temperature; mollusc; diversity
3.  How Many Species Are There on Earth and in the Ocean? 
PLoS Biology  2011;9(8):e1001127.
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.
Author Summary
Knowing the number of species on Earth is one of the most basic yet elusive questions in science. Unfortunately, obtaining an accurate number is constrained by the fact that most species remain to be described and because indirect attempts to answer this question have been highly controversial. Here, we document that the taxonomic classification of species into higher taxonomic groups (from genera to phyla) follows a consistent pattern from which the total number of species in any taxonomic group can be predicted. Assessment of this pattern for all kingdoms of life on Earth predicts ∼8.7 million (±1.3 million SE) species globally, of which ∼2.2 million (±0.18 million SE) are marine. Our results suggest that some 86% of the species on Earth, and 91% in the ocean, still await description. Closing this knowledge gap will require a renewed interest in exploration and taxonomy, and a continuing effort to catalogue existing biodiversity data in publicly available databases.
PMCID: PMC3160336  PMID: 21886479
4.  Current and Future Patterns of Global Marine Mammal Biodiversity 
PLoS ONE  2011;6(5):e19653.
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.
PMCID: PMC3100303  PMID: 21625431
5.  Global Human Footprint on the Linkage between Biodiversity and Ecosystem Functioning in Reef Fishes 
PLoS Biology  2011;9(4):e1000606.
A global survey of reef fishes shows that the consequences of biodiversity loss are greater than previously anticipated as ecosystem functioning remained unsaturated with the addition of new species. Additionally, reefs worldwide, particularly those most diverse, are highly vulnerable to human impacts that are widespread and likely to worsen due to ongoing coastal overpopulation.
Difficulties in scaling up theoretical and experimental results have raised controversy over the consequences of biodiversity loss for the functioning of natural ecosystems. Using a global survey of reef fish assemblages, we show that in contrast to previous theoretical and experimental studies, ecosystem functioning (as measured by standing biomass) scales in a non-saturating manner with biodiversity (as measured by species and functional richness) in this ecosystem. Our field study also shows a significant and negative interaction between human population density and biodiversity on ecosystem functioning (i.e., for the same human density there were larger reductions in standing biomass at more diverse reefs). Human effects were found to be related to fishing, coastal development, and land use stressors, and currently affect over 75% of the world's coral reefs. Our results indicate that the consequences of biodiversity loss in coral reefs have been considerably underestimated based on existing knowledge and that reef fish assemblages, particularly the most diverse, are greatly vulnerable to the expansion and intensity of anthropogenic stressors in coastal areas.
Author Summary
The increasing intensity of human disturbance worldwide is triggering unprecedented biodiversity losses, which is raising concerns over whether ecosystems will work and continue delivering goods and services to humanity. In contrast to previous experimental studies, which describe saturating relationships between ecosystem functioning and biodiversity, we show that in reef fish systems, functioning (as standing biomass) accelerates with the addition of new species. This non-saturating relationship implies unique contributions of species to the functioning of this ecosystem and indicates that the consequences of losing biodiversity are significantly greater than previously anticipated. We also demonstrate a negative effect of human density on reef fish functioning such that for the same number of people the loss of standing biomass is significantly larger in more diverse ecosystems. Unfortunately, human effects can arise through multiple stressors (such as fishing, coastal development, and land use) and are widespread and likely to worsen, as some 75% of the world's coral reefs are currently nearby human settlements and because almost all countries with coral reefs are expected to double their populations within the next 50 to 100 years. Our results call for both further investigation of the impact of biodiversity on ecosystem functioning and strategies to manage and prevent the increasing intensity and expansion of anthropogenic stressors in coastal areas.
PMCID: PMC3071368  PMID: 21483714
6.  Acute effects of removing large fish from a near-pristine coral reef 
Marine Biology  2010;157(12):2739-2750.
Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems.
PMCID: PMC3873048  PMID: 24391253
7.  The completeness of taxonomic inventories for describing the global diversity and distribution of marine fishes 
Taxonomic inventories (or species censuses) are the most elementary data in biogeography, macroecology and conservation biology. They play fundamental roles in the construction of species richness patterns, delineation of species ranges, quantification of extinction risk and prioritization of conservation efforts in hot spot areas. Given their importance, any issue related to the completeness of taxonomic inventories can have far-reaching consequences. Here, we used the largest publicly available database of georeferenced marine fish records to determine its usefulness in depicting the diversity and distribution of this taxonomic group. All records were grouped at multiple spatial resolutions to generate accumulation curves, from which the expected number of species were extrapolated using a variety of nonlinear models. Comparison of the inventoried number of species with that expected from the models was used to calculate the completeness of the taxonomic inventory at each resolution. In terms of the global number of fish species, we found that approximately 21% of the species remain to be described. In terms of spatial distribution, we found that the completeness of taxonomic data was highly scale dependent, with completeness being lower at finer spatial resolutions. At a 3° (approx. 350 km2) spatial resolution, less than 1.8% of the world's oceans have above 80% of their fish fauna currently described. Censuses of species were particularly incomplete in tropical areas and across the entire range of countries' gross domestic product (GDP), although the few censuses nearing completion were all along the coasts of a few developed countries or territories. Our findings highlight that failure to quantify the completeness of taxonomic inventories can introduce substantial flaws in the description of diversity patterns, and raise concerns over the effectiveness of conservation strategies based upon data that remain largely precarious.
PMCID: PMC2596190  PMID: 17999950
taxonomic census; fishes; completeness; global; spatial scales; species

Results 1-7 (7)