Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

Drafting Authors: Neil Adger, Pramod Aggarwal, Shardul Agrawala, Joseph Alcamo, Abdelkader Allali, Oleg Anisimov, Nigel Arnell, Michel Boko, Osvaldo Canziani, Timothy Carter, Gino Casassa, Ulisses Confalonieri, Rex Victor Cruz, Edmundo de Alba Alcaraz, William Easterling, Christopher Field, Andreas Fischlin, Blair Fitzharris, Carlos Gay García, Clair Hanson, Hideo Harasawa, Kevin Hennessy, Saleemul Huq, Roger Jones, Lucka Kajfež Bogataj, David Karoly, Richard Klein, Zbigniew Kundzewicz, Murari Lal, Rodel Lasco, Geoff Love, Xianfu Lu, Graciela Magrín, Luis José Mata, Roger McLean, Bettina Menne, Guy Midgley, Nobuo Mimura, Monirul Qader Mirza, José Moreno, Linda Mortsch, Isabelle Niang-Diop, Robert Nicholls, Béla Nováky, Leonard Nurse, Anthony Nyong, Michael Oppenheimer, Jean Palutikof, Martin Parry, Anand Patwardhan, Patricia Romero Lankao, Cynthia Rosenzweig, Stephen Schneider, Serguei Semenov, Joel Smith, John Stone, Jean-Pascal van Ypersele, David Vaughan, Coleen Vogel, Thomas Wilbanks, Poh Poh Wong, Shaohong Wu, Gary Yohe

Contribution of Working Group II to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change

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Modern Applied Statistics With S

The distributions of many terrestrial organisms are currently shifting in latitude or elevation in response to changing climate Using a meta-analysis, we estimated that the distributions of species have recently shifted to higher elevations at a median rate of 110 meters per decade, and to higher latitudes at a median rate of 169 kilometers per decade These rates are approximately two and three times faster than previously reported The distances moved by species are greatest in studies showing the highest levels of warming, with average latitudinal shifts being generally sufficient to track temperature changes However, individual species vary greatly in their rates of change, suggesting that the range shift of each species depends on multiple internal species traits and external drivers of change Rapid average shifts derive from a wide diversity of responses by individual species

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Rapid Range Shifts of Species Associated with High Levels of Climate Warming

Habitat loss has pervasive and disruptive impacts on biodiversity in habitat remnants. The magnitude of the ecological impacts of habitat loss can be exacerbated by the spatial arrangement -- or fragmentation -- of remaining habitat. Fragmentation per se is a landscape-level phenomenon in which species that survive in habitat remnants are confronted with a modified environment of reduced area, increased isolation and novel ecological boundaries. The implications of this for individual organisms are many and varied, because species with differing life history strategies are differentially affected by habitat fragmentation. Here, we review the extensive literature on species responses to habitat fragmentation, and detail the numerous ways in which confounding factors have either masked the detection, or prevented the manifestation, of predicted fragmentation effects. Large numbers of empirical studies continue to document changes in species richness with decreasing habitat area, with positive, negative and no relationships regularly reported. The debate surrounding such widely contrasting results is beginning to be resolved by findings that the expected positive species-area relationship can be masked by matrix-derived spatial subsidies of resources to fragment-dwelling species and by the invasion of matrix-dwelling species into habitat edges. Significant advances have been made recently in our understanding of how species interactions are altered at habitat edges as a result of these changes. Interestingly, changes in biotic and abiotic parameters at edges also make ecological processes more variable than in habitat interiors. Individuals are more likely to encounter habitat edges in fragments with convoluted shapes, leading to increased turnover and variability in population size than in fragments that are compact in shape. Habitat isolation in both space and time disrupts species distribution patterns, with consequent effects on metapopulation dynamics and the genetic structure of fragment-dwelling populations. Again, the matrix habitat is a strong determinant of fragmentation effects within remnants because of its role in regulating dispersal and dispersal-related mortality, the provision of spatial subsidies and the potential mediation of edge-related microclimatic gradients. We show that confounding factors can mask many fragmentation effects. For instance, there are multiple ways in which species traits like trophic level, dispersal ability and degree of habitat specialisation influence species-level responses. The temporal scale of investigation may have a strong influence on the results of a study, with short-term crowding effects eventually giving way to long-term extinction debts. Moreover, many fragmentation effects like changes in genetic, morphological or behavioural traits of species require time to appear. By contrast, synergistic interactions of fragmentation with climate change, human-altered disturbance regimes, species interactions and other drivers of population decline may magnify the impacts of fragmentation. To conclude, we emphasise that anthropogenic fragmentation is a recent phenomenon in evolutionary time and suggest that the final, long-term impacts of habitat fragmentation may not yet have shown themselves.

Confounding factors in the detection of species responses to habitat fragmentation

Species-energy theory indicates that recent climate warming should have driven increases in species richness in cool and species-poor parts of the Northern Hemisphere. We confirm that the average species richness of British butterflies has increased since 1970–82, but much more slowly than predicted from changes of climate: on average, only one-third of the predicted increase has taken place. The resultant species assemblages are increasingly dominated by generalist species that were able to respond quickly. The time lag is confirmed by the successful introduction of many species to climatically suitable areas beyond their ranges. Our results imply that it may be decades or centuries before the species richness and composition of biological communities adjusts to the current climate.

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Species richness changes lag behind climate change

Many factors, including climate, resource availability, and habitat diversity, have been proposed as determinants of global diversity, but the links among them have rarely been studied. Using structural equation modeling (SEM), we investigated direct and indirect effects of climate variables, host-plant richness, and habitat diversity on butterfly species richness across Britain, at 20-km grid resolution. These factors were all important determinants of butterfly diversity, but their relative contributions differed between habitat generalists and specialists, and whether the effects were direct or indirect. Climate variables had strong effects on habitat generalists, whereas host-plant richness and habitat diversity contributed relatively more for habitat specialists. Considering total effects (direct and indirect together), climate variables had the strongest link to butterfly species richness for all groups of species. The results suggest that different mechanistic hypotheses to explain species richness may be more appropriate for habitat generalists and specialists, with generalists hypothesized to show direct physiological limitations and specialists additionally being constrained by trophic interactions (climate affecting host-plant richness).

Direct and indirect effects of climate and habitat factors on butterfly diversity

1. A major, and largely unexplored, uncertainty in projecting the impact of climate change on biodiversity is the consequence of altered interspecific interactions, for example between parasitoids and their hosts. The present study investigated parasitism in the Brown Argus butterfly, Aricia agestis; a species that has expanded northward in Britain during the last 30 years in association with climate warming. 2. Aricia agestis larvae suffered lower mortality from parasitoids in newly colonised areas compared with long-established populations. This result was consistent over four consecutive generations (2 years) when comparing one population of each type, and also when several populations within the historical and recently colonised range of the species were compared within a single year. Thus, A. agestis appears to be partially escaping from parasitism as it expands northwards. 3. Reduced parasitism occurred despite the fact that several of the parasitoid species associated with A. agestis were already present in the newly colonised areas, supported predominantly by an alternative host species, the Common Blue butterfly, Polyommatus icarus. 4. As the species expand their distributions into areas of increased climatic suitability, invasion fronts may escape from natural enemies, enhancing rates of range expansion. The results suggest that the decoupling of interspecific interactions may allow some species to exploit a wider range of environments and to do so more rapidly than previously thought possible.

Escape from natural enemies during climate-driven range expansion: a case study

Global average surface temperature has increased by around 0.6 oC during the past century and will continue to rise in the future. Understanding how these changes in climate have affected biological systems has attracted a vast research effort during the last two decades. Here I review the existing empirical evidence of how insects have responded to these changes in climate, especially to the increases in temperature. Evidence provided here indicates that insects are good indicators of current human-driven climate change. They have responded to warming in all the predicted ways, from changes in phenology and distribution, to undergoing evolutionary changes albeit at the population level. Insects have also provided examples of how biodiversity and community structure is affected by current climate change. However, there are still many unknowns in our understanding of the detrimental and beneficial effects of climate change to biological systems. Future research needs to consider other climatic factors, geographic and taxonomic bias and the effect of individual responses on species interaction.

How are insects responding to global warming

The effects of dispersal ability, measured as two wing size categories (brachypterous vs. macropterous), on the distribution, abundance and body size, and on the relationships between these variables were examined in eighty-four species of carabid beetles over twenty-two sites in the northern Iberian peninsula. Geographic ranges of species (restricted to the northern Iberian peninsula vs. widespread European or wider range) were also taken into account in the analyses because macropterous species significantly tended to exhibit wider geographic ranges than did brachypterous species. Regional distributions were wider in brachypterous-restricted and brachypterous-widespread species than in macropterous-widespread species. The three groups did not differ in abundance. Differences in regional distributions between groups may be explained by referring to a trade-off between dispersal ability and establishment ability indicated in the literature. Macropterous species would occupy relatively few sites due to a high frequency of unsuccessful colonizations. The relationships between regional distribution and abundance were positive for all the three groups, brachypterous-restricted, brachypterous- widespread and macropterous-widespread species. The regression line for the last group showed a lower elevation than those for brachypterous-restricted and brachypterous- widespread species. This fact was probably due to differences in regional distributions between groups. No relationship between abundance and body size was significant. Regressions of regional distribution on body size were positive in brachypterous-restricted and brachypterous- widespread carabids, but the relationship was not significant in macropterous-widespread carabids. These results were interpreted in terms of differences in body size-dependency of travelling velocities between flying and running carabids.

Rosa Menéndez

Papers

Species richness changes lag behind climate change

Direct and indirect effects of climate and habitat factors on butterfly diversity

Escape from natural enemies during climate-driven range expansion: a case study

How are insects responding to global warming

Patterns in the distribution, abundance and body size of carabid beetles (Coleoptera: Caraboidea) in relation to dispersal ability