Conservation biogeography of the Antarctic

Aim

To present a synthesis of past biogeographic analyses and a new approach based on spatially explicit biodiversity information for the Antarctic region to identify biologically distinct areas in need of representation in a protected area network.

Location

Antarctica and the sub‐Antarctic.

Methods

We reviewed and summarized published biogeographic studies of the Antarctic. We then developed a biogeographic classification for terrestrial conservation planning in Antarctica by combining the most comprehensive source of Antarctic biodiversity data available with three spatial frameworks: (1) a 200‐km grid, (2) a set of areas based on physical parameters known as the environmental domains of Antarctica and (3) expert‐defined bioregions. We used these frameworks, or combinations thereof, together with multivariate techniques to identify biologically distinct areas.

Results

Early studies of continental Antarctica typically described broad bioregions, with the Antarctic Peninsula usually identified as biologically distinct from continental Antarctica; later studies suggested a more complex biogeography. Increasing complexity also characterizes the sub‐Antarctic and marine realms, with differences among studies often attributable to the focal taxa. Using the most comprehensive terrestrial data available and by combining the groups formed by the environmental domains and expert‐defined bioregions, we were able to identify 15 biologically distinct, ice‐free, Antarctic Conservation Biogeographic Regions (ACBRs), encompassing the continent and close lying islands.

Main conclusions

Ice‐free terrestrial Antarctica comprises several distinct bioregions that are not fully represented in the current Antarctic Specially Protected Area network. Biosecurity measures between these ACBRs should also be developed to prevent biotic homogenization in the region.     

物种保护的理论基础——从岛屿生物地理学理论到集合种群理论

全球面临着生境破碎化的危机,物种保护已成为人类面临的重大课题,并不是所有的人对岛屿生物地理学理论的产生及其关注的海洋岛屿都很熟悉,但是越来越多生物赖以生存的自然栖息地的丧失和破碎化都是有目共睹的,岛屿生物地理学和集合种群理论是目前物种保护的两个基本理论,物种迁入率和绝灭率的动态变化决策岛屿上的物种丰富度是岛屿生物地理学理论的核心内容,而集合种群理论关注的是局部种群之间个体迁移的动态以及物种的续存条件,在概述两个理论形成、发展及其核心内容的基础上,着重比较它们的异同点以及在生态学理论和实践中的应用,并论述物种保护理论范式从岛屿生物地理学向集合种群理论转变的基本背景和原因。     

Population structure, propagule pressure, and conservation biogeography in the sub-Antarctic: lessons from indigenous and invasive springtails

The patterns in and the processes underlying the distribution of invertebrates among Southern Ocean islands and across vegetation types on these islands are reasonably well understood. However, few studies have examined the extent to which populations are genetically structured. Given that many sub‐Antarctic islands experienced major glaciation and volcanic activity, it might be predicted that substantial population substructure and little genetic isolation‐by‐distance should characterize indigenous species. By contrast, substantially less population structure might be expected for introduced species. Here, we examine these predictions and their consequences for the conservation of diversity in the region. We do so by examining haplotype diversity based on mitochondrial cytochrome c oxidase subunit I sequence data, from two indigenous (Cryptopygus antarcticus travei, Tullbergia bisetosa) and two introduced (Isotomurus cf. palustris, Ceratophysella denticulata) springtail species from Marion Island. We find considerable genetic substructure in the indigenous species that is compatible with the geological and glacialogical history of the island. Moreover, by employing ecological techniques, we show that haplotype diversity is likely much higher than our sequenced samples suggest. No structure is found in the introduced species, with each being represented by a single haplotype only. This indicates that propagule pressure is not significant for these small animals unlike the situation for other, larger invasive species: a few individuals introduced once are likely to have initiated the invasion. These outcomes demonstrate that sampling must be more comprehensive if the population history of indigenous arthropods on these islands is to be comprehended, and that the risks of within‐ and among‐island introductions are substantial. The latter means that, if biogeographical signal is to be retained in the region, great care must be taken to avoid inadvertent movement of indigenous species among and within islands. Thus, quarantine procedures should also focus on among‐island movements.     

Identifying complementary areas for conservation in Thailand: an example using owls, hawkmoths and tiger beetles

Setting priorities in biodiversity conservation requires that explicit, efficient and appropriate methods are developed and made available to conservation managers. The principle of complementarity is fundamental to the most efficient of these methods. Given a goal of only a single representation of each taxon, complementarity analysis of data sets of owls, hawkmoths and tiger beetles in Thailand yields near-minimum sets of 6, 14 and 34 areas, respectively. The consensus of these sets comprises 48 areas. However, when the data are combined into a single data set, complementarity analysis gives a more efficient solution of 46 areas. Over 90% of the owls, hawkmoths and tiger beetles are already represented as a minimum of one population within the current protected areas network of Thailand. However, an additional 18 areas are still required for complete representation. Several of these additional areas are discussed and their potential contribution to biodiversity conservation in Thailand examined. Khao Pok Yo and Doi Pa Hom Pok are noted as being of particular importance. The case of Sanpatong emphasizes the need to remain aware of the biology and ecology of the species under study.     

Is a new paradigm emerging for oceanic island biogeography?

Following several decades during which two dissimilar and incompatible models (equilibrium and vicariance) dominated island biogeography, recent publications have documented patterns that point the way towards a new paradigm that includes elements of both models, as well as some novel aspects. Many of these seminal contributions have been made possible by the recent development of robust, temporally calibrated phylogenies used in concert with increasingly precise and reliable geological reconstructions of oceanic regions. Although a new general model of oceanic island biogeography has not yet been proposed, in this brief overview I present six hypotheses that summarize aspects of the emerging paradigm. These hypotheses deal with: the frequency of dispersal over oceanic water barriers by terrestrial organisms; the existence of substantial variation in the amount of dispersal (and gene flow) within a given set of related species within a given archipelago; the frequency, extent and impact on species richness of diversification within archipelagos; the frequent correlation of island age and the age of the species that live on the island; the long-term persistence of species on oceanic islands; and the occasional recolonization of continents by species from clades that diversified on islands. Identifying, testing, and seeking means of synthesizing these and other emerging hypotheses may allow a new conceptual paradigm to emerge.     

Taxonomic homogenization in ungulates: patterns and mechanisms at local and global scales

Aim The aim of this paper is to examine taxonomic homogenization in ungulates globally and at the local scale in South Africa. Specifically, we aim to examine the roles of distance, scale, time, extinctions vs. introductions, and extralimital vs. extraregional introductions in the homogenization of ungulate biotas, and to determine pathways of introduction of ungulate species globally and the proximate explanatory variables of ungulate introductions in South Africa. Location Forty‐one countries globally and three spatial resolutions in South Africa. Methods Indigenous, extirpated and established introduced ungulate species data were obtained for countries globally, and at a quarter‐degree grid‐cell resolution in South Africa. Homogenization was calculated using Jaccard’s index of similarity (JI) for countries globally and for three spatial resolutions in South Africa. Zoo holdings and transfer data from the International Species Information System database were used to investigate the relationship between non‐indigenous ungulate species introductions and the number of non‐indigenous ungulate species in zoos. Relationships between JI and species richness, and between numbers of introductions and several environmental and social factors were examined using generalized linear models. Results Homogenization in ungulates was 2% for countries globally and 8% at the coarsest resolution in South Africa. Homogenization increased with increasing resolution and with time, but it decreased with increasing percentage change in species richness. Globally, introductions contributed more to homogenization than did extinctions. Within South Africa, extralimital introductions contributed more to the homogenization of ungulate assemblages than did extraregional ones, and ungulates were typically introduced to high‐income areas with high human population and livestock densities. The same was not true in the past, when ungulates were introduced to ungulate species‐poor areas. The number of non‐indigenous ungulate species established in a country is significantly related to the number of non‐indigenous ungulate species in zoos in the country, possibly owing to sales of surplus animals from zoos. Main conclusions Ungulate faunas are homogenized at both the global scale and in South Africa, with extralimital introductions being of considerable significance regionally. In consequence, increasing attention will have to be given to the conservation consequences of ungulate translocations, both within particular geopolitical regions and across the globe.     

Biotic and environmental dynamics through the Late Jurassic–Early Cretaceous transition: evidence for protracted faunal and ecological turnover

The Late Jurassic to Early Cretaceous interval represents a time of environmental upheaval and cataclysmic events, combined with disruptions to terrestrial and marine ecosystems. Historically, the Jurassic/Cretaceous (J/K) boundary was classified as one of eight mass extinctions. However, more recent research has largely overturned this view, revealing a much more complex pattern of biotic and abiotic dynamics than has previously been appreciated. Here, we present a synthesis of our current knowledge of Late Jurassic–Early Cretaceous events, focusing particularly on events closest to the J/K boundary. We find evidence for a combination of short‐term catastrophic events, large‐scale tectonic processes and environmental perturbations, and major clade interactions that led to a seemingly dramatic faunal and ecological turnover in both the marine and terrestrial realms. This is coupled with a great reduction in global biodiversity which might in part be explained by poor sampling. Very few groups appear to have been entirely resilient to this J/K boundary ‘event’, which hints at a ‘cascade model’ of ecosystem changes driving faunal dynamics. Within terrestrial ecosystems, larger, more‐specialised organisms, such as saurischian dinosaurs, appear to have suffered the most. Medium‐sized tetanuran theropods declined, and were replaced by larger‐bodied groups, and basal eusauropods were replaced by neosauropod faunas. The ascent of paravian theropods is emphasised by escalated competition with contemporary pterosaur groups, culminating in the explosive radiation of birds, although the timing of this is obfuscated by biases in sampling. Smaller, more ecologically diverse terrestrial non‐archosaurs, such as lissamphibians and mammaliaforms, were comparatively resilient to extinctions, instead documenting the origination of many extant groups around the J/K boundary. In the marine realm, extinctions were focused on low‐latitude, shallow marine shelf‐dwelling faunas, corresponding to a significant eustatic sea‐level fall in the latest Jurassic. More mobile and ecologically plastic marine groups, such as ichthyosaurs, survived the boundary relatively unscathed. High rates of extinction and turnover in other macropredaceous marine groups, including plesiosaurs, are accompanied by the origin of most major lineages of extant sharks. Groups which occupied both marine and terrestrial ecosystems, including crocodylomorphs, document a selective extinction in shallow marine forms, whereas turtles appear to have diversified. These patterns suggest that different extinction selectivity and ecological processes were operating between marine and terrestrial ecosystems, which were ultimately important in determining the fates of many key groups, as well as the origins of many major extant lineages. We identify a series of potential abiotic candidates for driving these patterns, including multiple bolide impacts, several episodes of flood basalt eruptions, dramatic climate change, and major disruptions to oceanic systems. The J/K transition therefore, although not a mass extinction, represents an important transitional period in the co‐evolutionary history of life on Earth.     

Extreme homogenization: The past,present and future of mammal assemblages on islands

Aim

We documented how the similarity of mammal assemblages on continental and oceanic islands has changed since initial human colonization, since European arrival and overall. We investigated how levels of similarity might change in the future.

Location

Continental and oceanic islands worldwide.

Time period

Human settlement of islands to the present, as well as projections for the future.

Major taxa studied

Mammals.

Methods

We used mammal occurrence data on islands to calculate the change in similarity using a pairwise approach based on Jaccard's index and a multisite approach based on Jaccard's and Sørensen's measures. We divided the mammal assemblages into two time periods, before and after island colonization or trade began with Europeans. We unpacked the mechanisms driving changes in similarity, exploring how initial similarity interacts with seven types of species turnover events to determine overall change. Finally, we calculated how future similarity levels will change if past trends in introductions and extinctions continue.

Results

Mammals, on both continental and oceanic islands, show one of the most pronounced cases of homogenization ever observed, and on oceanic islands mammals show the largest increase in homogenization ever observed for a terrestrial group. Most of the homogenization observed to date has been driven by recent historical changes, not by changes that occurred before European arrival. If current patterns of species introductions and extinctions continue, then oceanic islands will experience little additional homogenization, whereas continental islands will homogenize greatly beyond current levels.

Main conclusions

Mammal assemblages on oceanic islands show nearly an order of magnitude greater change in similarity than plant and bird assemblages. Projections of future similarity indicate that continental and oceanic islands are on different trajectories of change. These trajectories could be altered by management actions, but in some cases those actions that would be impactful run counter to current conservation norms.     

Human impacts on environment–diversity relationships: evidence for biotic homogenization from butterfly species richness patterns

Aim Broad‐scale spatial variation in species richness relates to climate and physical heterogeneity but human activities may be changing these patterns. We test whether climate and heterogeneity predict butterfly species richness regionally and across Canada and whether these relationships change in areas of human activity. Location Canada. Methods We modelled the ranges of 102 butterfly species using genetic algorithms for rule‐set production (GARP). We then measured butterfly species richness and potentially important aspects of human activity and the natural environment. These were included in a series of statistical models to determine which factors are likely to affect butterfly species richness in Canada. We considered patterns across Canada, within predominantly natural areas, human‐dominated areas and particular ecozones. We examined independent observations of butterfly species currently listed under Canada's endangered species legislation to test whether these were consistent with findings from statistical models. Results Growing season temperature is the main determinant of butterfly species richness across Canada, with substantial contributions from habitat heterogeneity (measured using elevation). Only in the driest areas does precipitation emerge as a leading predictor of richness. The slope of relationships between all of these variables and butterfly species richness becomes shallower in human‐dominated areas, but butterfly richness is still highest there. Insecticide applications, habitat loss and road networks reduce butterfly richness in human‐dominated areas, but these effects are relatively small. All of Canada's at‐risk butterfly species are located in these human‐dominated areas. Main conclusions Temperature affects butterfly species richness to a greater extent than habitat heterogeneity at fine spatial scales and is generally far more important than precipitation, supporting both the species richness–energy and habitat heterogeneity hypotheses. Human activities, especially in southern Canada, appear to cause surprisingly consistent trends in biotic homogenization across this region, perhaps through range expansion of common species and loss of range‐restricted species.     

Phylogeography of the bushmaster (Lachesis tnuta: Viperidae): implications for neotropical biogeography, systematics, and conservation

We used mitochondrial gene sequences to reconstruct phylogenetic relationships among subspecies of the bushmaster, Lachesis muta. These large vipers are widely distributed in lowland tropical forests in Central and South America, where three of four allopatric subspecies are separated by montane barriers. Our phylogeny indicates that the four subspecies belong to two clades, the Central American and South American lineages. We use published molecular studies of other taxa to estimate a 'reptilian mtDNA rate' and thus temporal boundaries for major lineage divergences in Lachesis. We estimate that the Central and South American forms diverged 18-6 Mya, perhaps due to the uplifting of the Andes, whereas the two Central American subspecies may have diverged 11-4 Mya with the uprising of the Cordillera de Talamanca that separates them today. South American bushmasters from the Amazon Basin and the Atlantic Forest are not strongly differentiated, perhaps due to episodic gene flow during the Pleistocene, when suitable habitat for this species was at times more continuous. Our results agree with previous evidence that genetic divergence among some neotropical vertebrates pre-dated Pleistocene forest fragmentation cycles and the appearance of the Panamanian Isthmus. Based on morphological, behavioral, and molecular evidence, we recognize three species of Lachesis. In addition to L. muta, the widespread South American form, the Central American forms are treated as distinct species (L. meknocephak and L. stenophrys), each deserving of special conservation status due to restricted distribution and habitat destruction.     

Old and new challenges in using species diversity for assessing biodiversity

Although the maintenance of diversity of living systems is critical for ecosystem functioning, the accelerating pace of global change is threatening its preservation. Standardized methods for biodiversity assessment and monitoring are needed. Species diversity is one of the most widely adopted metrics for assessing patterns and processes of biodiversity, at both ecological and biogeographic scales. However, those perspectives differ because of the types of data that can be feasibly collected, resulting in differences in the questions that can be addressed. Despite a theoretical consensus on diversity metrics, standardized methods for its measurement are lacking, especially at the scales needed to monitor biodiversity for conservation and management purposes. We review the conceptual framework for species diversity, examine common metrics, and explore their use for biodiversity conservation and management. Key differences in diversity measures at ecological and biogeographic scales are the completeness of species lists and the ability to include information on species abundances. We analyse the major pitfalls and problems with quantitative measurement of species diversity, look at the use of weighting measures by phylogenetic distance, discuss potential solutions and propose a research agenda to solve the major existing problems.     

Evolutionary biology and practical conservation: bridging a widening gap

At the ecosystem, species and population level, available measures suggest that average rates of loss of populations and habitats are now around 1% per year and seem likely to increase in the future. Habitat conversion continues in most parts of the world, especially in areas of high species richness, and novel threats, especially climate change, will pose new challenges. With this pressure, maintaining evolutionary processes in natural populations will be critical to longer term persistence, and will often require specific planning relevant to the context. However, in many areas of policy and practice, urgent actions tend to focus on pattern-based analyses and considerations of evolutionary and ecological processes are neglected. At a variety of levels, from setting goals to implementing conservation management at the site or species level, there are simple adjustments that can be made. Improved methods for integrating the work of scientists and policymakers is recommended, from the beginning to end of the planning process.     

Balance between climate change mitigation benefits and land use impacts of bioenergy: conservation implications for European birds

Both climate change and habitat modification exert serious pressure on biodiversity. Although climate change mitigation has been identified as an important strategy for biodiversity conservation, bioenergy remains a controversial mitigation action due to its potential negative ecological and socio-economic impacts which arise through habitat modification by land use change. While the debate continues, the separate or simultaneous impacts of both climate change and bioenergy on biodiversity have not yet been compared. We assess projected range shifts of 156 European bird species by 2050 under two alternative climate change trajectories: a baseline scenario, where the global mean temperature increases by 4 °C by the end of the century, and a 2 degrees scenario, where global concerted effort limits the temperature increase to below 2 °C. For the latter scenario, we also quantify the pressure exerted by increased cultivation of energy biomass as modelled by IMAGE2.4, an integrated land use model. The global bioenergy use in this scenario is in the lower end of the range of previously estimated sustainable potential. Under the assumptions of these scenarios, we find that the magnitude of range shifts due to climate change is far greater than the impact of land conversion to woody bioenergy plantations within the European Union, and that mitigation of climate change reduces the exposure experienced by species. However, we identified potential for local conservation conflict between priority areas for conservation and bioenergy production. These conflicts must be addressed by strict bioenergy sustainability criteria that acknowledge biodiversity conservation needs beyond existing protected areas and apply also to biomass imported from outside the European Union.     

Rivers in peril inside and outside protected areas: a systematic approach to conservation assessment of river ecosystems

This paper establishes a framework within which a rapid and pragmatic assessment of river ecosystems can be undertaken at a broad, subcontinental scale, highlighting some implications for achieving conservation of river biodiversity in water‐limited countries. The status of river ecosystems associated with main rivers in South Africa was assessed based on the extent to which each ecosystem had been altered from its natural condition. This requires consistent data on river integrity for the entire country, which was only available for main rivers; tributaries were thus excluded from the analyses. The state of main river ecosystems in South Africa is dire: 84% of the ecosystems are threatened, with a disturbing 54% critically endangered, 18% endangered, and 12% vulnerable. Protection levels were measured as the proportion of conservation target achieved within protected areas, where the conservation target was set as 20% of the total length of each river ecosystem. Sixteen of the 112 main river ecosystems are moderately to well represented within protected areas; the majority of the ecosystems have very low levels of representation, or are not represented at all within protected areas. Only 50% of rivers within protected areas are intact, but this is a higher proportion compared to rivers outside (28%), providing some of the first quantitative data on the positive role protected areas can play in conserving river ecosystems. This is also the first assessment of river ecosystems in South Africa to apply a similar approach to parallel assessments of terrestrial, marine, and estuarine ecosystems, and it revealed that main river ecosystems are in a critical state, far worse than terrestrial ecosystems. Ecosystem status is likely to differ with the inclusion of tributaries, since options may well exist for conserving critically endangered ecosystems in intact tributaries, which are generally less regulated than main rivers. This study highlights the importance of healthy tributaries for achieving river conservation targets, and the need for managing main rivers as conduits across the landscape to support ecological processes that depend on connectivity. We also highlight the need for a paradigm shift in the way protected areas are designated, as well as the need for integrated river basin management plans to include explicit conservation visions, targets, and strategies to ensure the conservation of freshwater ecosystems and the services they provide.     

Modelling the dynamic biogeography of the wildcat: implications for taxonomy and conservation

There is still no clear consensus on how to relate geographical variation in the morphology and genetics of the globally widespread wildcat Felis silvestris to its taxonomy and systematics. Reconstructing the dynamic biogeography of the wildcat provides insight into how current geographical patterns of morphological and molecular variation may have developed. A geographical information system was used to infer climate-change influences using a deduced distribution model (DDM) to reconstruct the wildcat's geographical distribution at four points in time from the Last Glacial Maximum [LGM; 18 000 years before present ( bp )] until today. The DDM for 9000  bp , when mean global temperatures were 2 °C more than today, provides insight into how current global warming will affect the wildcat's distribution 50–100 years into future. Modelled distributions were assessed against known geographical barriers or unsuitable habitats, which may have separated populations and led to known morphological and genetic divergence. The DDM_today corresponds well with known contemporary wildcat distribution records, except where wildcats would be expected to be excluded (e.g. high human population densities, potential competitors, inaccessible islands). The DDM_today also corresponds closely with the results of recent studies on skull morphometrics and phylogeography, which support hypothesized colonizations of Africa and Asia from Europe during the late Pleistocene. Although DDM palaeo-distributions are more uncertain, they correspond to expected dramatic declines in northern Eurasia during the LGM, and significant distributional decline in central Asia, the Sahara and southern Africa, owing to increased aridity during climate cooling. From the DDM₉₀₀₀ model moderate global warming is hypothesized to impact minimally on wildcats, except in the Middle East and south-west Asia.