Walter Jetz

The global diversity of birds in space and time

Current global patterns of biodiversity result from processes that operate over both space and time and thus require an integrated macroecological and macroevolutionary perspective. Molecular time trees have advanced our understanding of the tempo and mode of diversification and have identified remarkable adaptive radiations across the tree of life. However, incomplete joint phylogenetic and geographic sampling has limited broad-scale inference. Thus, the relative prevalence of rapid radiations and the importance of their geographic settings in shaping global biodiversity patterns remain unclear. Here we present, analyse and map the first complete dated phylogeny of all 9,993 extant species of birds, a widely studied group showing many unique adaptations. We find that birds have undergone a strong increase in diversification rate from about 50 million years ago to the near present. This acceleration is due to a number of significant rate increases, both within songbirds and within other young and mostly temperate radiations including the waterfowl, gulls and woodpeckers. Importantly, species characterized with very high past diversification rates are interspersed throughout the avian tree and across geographic space. Geographically, the major differences in diversification rates are hemispheric rather than latitudinal, with bird assemblages in Asia, North America and southern South America containing a disproportionate number of species from recent rapid radiations. The contribution of rapidly radiating lineages to both temporal diversification dynamics and spatial distributions of species diversity illustrates the benefits of an inclusive geographical and taxonomical perspective. Overall, whereas constituent clades may exhibit slowdowns, the adaptive zone into which modern birds have diversified since the Cretaceous may still offer opportunities for diversification.

Global patterns and predictors of marine biodiversity across taxa

Global patterns of species richness and their structuring forces have fascinated biologists since Darwin 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. Whereas land diversity patterns and their predictors are known for numerous taxa, our understanding of global marine diversity has been more limited, with recent findings revealing some striking contrasts to widely held terrestrial paradigms. 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.

Global patterns and determinants of vascular plant diversity

Plants, with an estimated 300,000 species, provide crucial primary production and ecosystem structure. To date, our quantitative understanding of diversity gradients of megadiverse clades such as plants has been hampered by the paucity of distribution data. Here, we investigate the global-scale species-richness pattern of vascular plants and examine its environmental and potential historical determinants. Across 1,032 geographic regions worldwide, potential evapotranspiration, the number of wet days per year, and measurements of topographical and habitat heterogeneity emerge as core predictors of species richness. After accounting for environmental effects, the residual differences across the major floristic kingdoms are minor, with the exception of the uniquely diverse Cape Region, highlighting the important role of historical contingencies. Notably, the South African Cape region contains more than twice as many species as expected by the global environmental model, confirming its uniquely evolved flora. A combined multipredictor model explains ≈70% of the global variation in species richness and fully accounts for the enigmatic latitudinal gradient in species richness. The models illustrate the geographic interplay of different environmental predictors of species richness. Our findings highlight that different hypotheses about the causes of diversity gradients are not mutually exclusive, but likely act synergistically with water–energy dynamics playing a dominant role. The presented geostatistical approach is likely to prove instrumental for identifying richness patterns of the many other taxa without single-species distribution data that still escape our understanding.

Essential Biodiversity Variables

A global system of harmonized observations is needed to inform scientists and policy-makers. Reducing the rate of biodiversity loss and averting dangerous biodiversity change are international goals, reasserted by the Aichi Targets for 2020 by Parties to the United Nations (UN) Convention on Biological Diversity (CBD) after failure to meet the 2010 target (1, 2). However, there is no global, harmonized observation system for delivering regular, timely data on biodiversity change (3). With the first plenary meeting of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) soon under way, partners from the Group on Earth Observations Biodiversity Observation Network (GEO BON) (4) are developing—and seeking consensus around—Essential Biodiversity Variables (EBVs) that could form the basis of monitoring programs worldwide.

A global assessment of endemism and species richness across island and mainland regions

Endemism and species richness are highly relevant to the global prioritization of conservation efforts in which oceanic islands have remained relatively neglected. When compared to mainland areas, oceanic islands in general are known for their high percentage of endemic species but only moderate levels of species richness, prompting the question of their relative conservation value. Here we quantify geographic patterns of endemism-scaled richness (“endemism richness”) of vascular plants across 90 terrestrial biogeographic regions, including islands, worldwide and evaluate their congruence with terrestrial vertebrates. Endemism richness of plants and vertebrates is strongly related, and values on islands exceed those of mainland regions by a factor of 9.5 and 8.1 for plants and vertebrates, respectively. Comparisons of different measures of past and future human impact and land cover change further reveal marked differences between mainland and island regions. While island and mainland regions suffered equally from past habitat loss, we find the human impact index, a measure of current threat, to be significantly higher on islands. Projected land-cover changes for the year 2100 indicate that land-use-driven changes on islands might strongly increase in the future. Given their conservation risks, smaller land areas, and high levels of endemism richness, islands may offer particularly high returns for species conservation efforts and therefore warrant a high priority in global biodiversity conservation in this century.

Species richness, hotspots, and the scale dependence of range maps in ecology and conservation.

Most studies examining continental-to-global patterns of species richness rely on the overlaying of extent-of-occurrence range maps. Because a species does not occur at all locations within its geographic range, range-map-derived data represent actual distributional patterns only at some relatively coarse and undefined resolution. With the increasing availability of high-resolution climate and land-cover data, broad-scale studies are increasingly likely to estimate richness at high resolutions. Because of the scale dependence of most ecological phenomena, a significant mismatch between the presumed and actual scale of ecological data may arise. This may affect conclusions regarding basic drivers of diversity and may lead to errors in the identification of diversity hotspots. Here, we examine avian range maps of 834 bird species in conjunction with geographically extensive survey data sets on two continents to determine the spatial resolutions at which range-map data actually characterize species occurrences and patterns of species richness. At resolutions less than 2° (≈200 km), range maps overestimate the area of occupancy of individual species and mischaracterize spatial patterns of species richness, resulting in up to two-thirds of biodiversity hotspots being misidentified. The scale dependence of range-map accuracy poses clear limitations on broad-scale ecological analyses and conservation assessments. We suggest that range-map data contain less information than is generally assumed and provide guidance about the appropriate scale of their use.

Homage to Linnaeus: How many parasites? How many hosts?

Estimates of the total number of species that inhabit the Earth have increased significantly since Linnaeuss initial catalog of 20,000 species. The best recent estimates suggest that there are ≈6 million species. More emphasis has been placed on counts of free-living species than on parasitic species. We rectify this by quantifying the numbers and proportion of parasitic species. We estimate that there are between 75,000 and 300,000 helminth species parasitizing the vertebrates. We have no credible way of estimating how many parasitic protozoa, fungi, bacteria, and viruses exist. We estimate that between 3% and 5% of parasitic helminths are threatened with extinction in the next 50 to 100 years. Because patterns of parasite diversity do not clearly map onto patterns of host diversity, we can make very little prediction about geographical patterns of threat to parasites. If the threats reflect those experienced by avian hosts, then we expect climate change to be a major threat to the relatively small proportion of parasite diversity that lives in the polar and temperate regions, whereas habitat destruction will be the major threat to tropical parasite diversity. Recent studies of food webs suggest that ≈75% of the links in food webs involve a parasitic species; these links are vital for regulation of host abundance and potentially for reducing the impact of toxic pollutants. This implies that parasite extinctions may have unforeseen costs that impact the health and abundance of a large number of free-living species.

Additive threats from pathogens, climate and land-use change for global amphibian diversity

Amphibian population declines far exceed those of other vertebrate groups, with 30% of all species listed as threatened by the International Union for Conservation of Nature. The causes of these declines are a matter of continued research, but probably include climate change, land-use change and spread of the pathogenic fungal disease chytridiomycosis. Here we assess the spatial distribution and interactions of these primary threats in relation to the global distribution of amphibian species. We show that the greatest proportions of species negatively affected by climate change are projected to be found in Africa, parts of northern South America and the Andes. Regions with the highest projected impact of land-use and climate change coincide, but there is little spatial overlap with regions highly threatened by the fungal disease. Overall, the areas harbouring the richest amphibian faunas are disproportionately more affected by one or multiple threat factors than areas with low richness. Amphibian declines are likely to accelerate in the twenty-first century, because multiple drivers of extinction could jeopardize their populations more than previous, mono-causal, assessments have suggested.

Terrestrial animal tracking as an eye on life and planet

A brave new world with a wider view Researchers have long attempted to follow animals as they move through their environment. Until relatively recently, however, such efforts were limited to short distances and times in species large enough to carry large batteries and transmitters. New technologies have opened up new frontiers in animal tracking remote data collection. Hussey et al. review the unique directions such efforts have taken for marine systems, while Kays et al. review recent advances for terrestrial species. We have entered a new era of animal ecology, where animals act as both subjects and samplers of their environments. Science, this issue 10.1126/science.1255642, 10.1126/science.aaa2478 BACKGROUND The movement of animals makes them fascinating but difficult study subjects. Animal movements underpin many biological phenomena, and understanding them is critical for applications in conservation, health, and food. Traditional approaches to animal tracking used field biologists wielding antennas to record a few dozen locations per animal, revealing only the most general patterns of animal space use. The advent of satellite tracking automated this process, but initially was limited to larger animals and increased the resolution of trajectories to only a few hundred locations per animal. The last few years have shown exponential improvement in tracking technology, leading to smaller tracking devices that can return millions of movement steps for ever-smaller animals. Finally, we have a tool that returns high-resolution data that reveal the detailed facets of animal movement and its many implications for biodiversity, animal ecology, behavior, and ecosystem function. ADVANCES Improved technology has brought animal tracking into the realm of big data, not only through high-resolution movement trajectories, but also through the addition of other on-animal sensors and the integration of remote sensing data about the environment through which these animals are moving. These new data are opening up a breadth of new scientific questions about ecology, evolution, and physiology and enable the use of animals as sensors of the environment. High–temporal resolution movement data also can document brief but important contacts between animals, creating new opportunities to study social networks, as well as interspecific interactions such as competition and predation. With solar panels keeping batteries charged, “lifetime” tracks can now be collected for some species, while broader approaches are aiming for species-wide sampling across multiple populations. Miniaturized tags also help reduce the impact of the devices on the study subjects, improving animal welfare and scientific results. As in other disciplines, the explosion of data volume and variety has created new challenges and opportunities for information management, integration, and analysis. In an exciting interdisciplinary push, biologists, statisticians, and computer scientists have begun to develop new tools that are already leading to new insights and scientific breakthroughs. OUTLOOK We suggest that a golden age of animal tracking science has begun and that the upcoming years will be a time of unprecedented exciting discoveries. Technology continues to improve our ability to track animals, with the promise of smaller tags collecting more data, less invasively, on a greater variety of animals. The big-data tracking studies that are just now being pioneered will become commonplace. If analytical developments can keep pace, the field will be able to develop real-time predictive models that integrate habitat preferences, movement abilities, sensory capacities, and animal memories into movement forecasts. The unique perspective offered by big-data animal tracking enables a new view of animals as naturally evolved sensors of environment, which we think has the potential to help us monitor the planet in completely new ways. A massive multi-individual monitoring program would allow a quorum sensing of our planet, using a variety of species to tap into the diversity of senses that have evolved across animal groups, providing new insight on our world through the sixth sense of the global animal collective. We expect that the field will soon reach a transformational point where these studies do more than inform us about particular species of animals, but allow the animals to teach us about the world. Big-data animal tracking. The red trajectory shows how studies can now track animals with unprecedented detail, allowing researchers to predict the causes and consequences of movements, and animals to become environmental sensors. Multisensor tracking tags monitor movement, behavior, physiology, and environmental context. Geo- and biosciences merge now using a multitude of remote-sensing data. Understanding how social and interspecific interactions affect movement is the next big frontier. Moving animals connect our world, spreading pollen, seeds, nutrients, and parasites as they go about the their daily lives. Recent integration of high-resolution Global Positioning System and other sensors into miniaturized tracking tags has dramatically improved our ability to describe animal movement. This has created opportunities and challenges that parallel big data transformations in other fields and has rapidly advanced animal ecology and physiology. New analytical approaches, combined with remotely sensed or modeled environmental information, have opened up a host of new questions on the causes of movement and its consequences for individuals, populations, and ecosystems. Simultaneous tracking of multiple animals is leading to new insights on species interactions and, scaled up, may enable distributed monitoring of both animals and our changing environment.

Geometric constraints explain much of the species richness pattern in African birds

The world contains boundaries (e.g., continental edge for terrestrial taxa) that impose geometric constraints on the distribution of species ranges. Thus, contrary to traditional thinking, the expected species richness pattern in absence of ecological or physiographical factors is unlikely to be uniform. Species richness has been shown to peak in the middle of a bounded one-dimensional domain, even in the absence of ecological or physiographical factors. Because species ranges are not linear, an extension of the approach to two dimensions is necessary. Here we present a two-dimensional null model accounting for effects of geometric constraints. We use the model to examine the effects of continental edge on the distribution of terrestrial animals in Africa and compare the predictions with the observed pattern of species richness in birds endemic to the continent. Latitudinal, longitudinal, and two-dimensional patterns of species richness are predicted well from the modeled null effects alone. As expected, null effects are of high significance for wide ranging species only. Our results highlight the conceptual significance of an until recently neglected constraint from continental shape alone and support a more cautious analysis of species richness patterns at this scale.