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

https://bio.uib.no/te/papers/Grimm_2010_The_ODD_protocol_.pdf

The ODD protocol: A review and first update

Landscape modification and habitat fragmentation are key drivers of global species loss. Their effects may be understood by focusing on: (1) individual species and the processes threatening them, and (2) human-perceived landscape patterns and their correlation with species and assemblages. Individual species may decline as a result of interacting exogenous and endogenous threats, including habitat loss, habitat degradation, habitat isolation, changes in the biology, behaviour, and interactions of species, as well as additional, stochastic threats. Human-perceived landscape patterns that are frequently correlated with species assemblages include the amount and structure of native vegetation, the prevalence of anthropogenic edges, the degree of landscape connectivity, and the structure and heterogeneity of modified areas. Extinction cascades are particularly likely to occur in landscapes with low native vegetation cover, low landscape connectivity, degraded native vegetation and intensive land use in modified areas, especially if keystone species or entire functional groups of species are lost. This review (1) demonstrates that species-oriented and pattern-oriented approaches to understanding the ecology of modified landscapes are highly complementary, (2) clarifies the links between a wide range of interconnected themes, and (3) provides clear and consistent terminology. Tangible research and management priorities are outlined that are likely to benefit the conservation of native species in modified landscapes around the world.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1466-8238.2007.00287.x

Landscape modification and habitat fragmentation: a synthesis

Individual-based modeling is a new, exciting discipline that allows ecologists to explore, using computer simulations, how properties of populations and ecosystems might evolve from the characteristics and behaviors of individual organisms. Individual-based Modeling and Ecology, written by Volker Grimm and Steven F. Railsback, gives an excellent introduction and overview of this field. It should be read by everyone interested in individual-based modeling, and especially by anyone contemplating developing, or being involved with a group developing, an individualbased model.

http://www.gbv.de/dms/weimar/toc/396272002_toc.pdf

Individual-based modeling and ecology

Environmental heterogeneity is regarded as one of the most important factors governing species richness gradients. An increase in available niche space, provision of refuges and opportunities for isolation and divergent adaptation are thought to enhance species coexistence, persistence and diversification. However, the extent and generality of positive heterogeneity–richness relationships are still debated. Apart from widespread evidence supporting positive relationships, negative and hump-shaped relationships have also been reported. In a meta-analysis of 1148 data points from 192 studies worldwide, we examine the strength and direction of the relationship between spatial environmental heterogeneity and species richness of terrestrial plants and animals. We find that separate effects of heterogeneity in land cover, vegetation, climate, soil and topography are significantly positive, with vegetation and topographic heterogeneity showing particularly strong associations with species richness. The use of equal-area study units, spatial grain and spatial extent emerge as key factors influencing the strength of heterogeneity–richness relationships, highlighting the pervasive influence of spatial scale in heterogeneity–richness studies. We provide the first quantitative support for the generality of positive heterogeneity–richness relationships across heterogeneity components, habitat types, taxa and spatial scales from landscape to global extents, and identify specific needs for future comparative heterogeneity–richness research.

/pdf/environmental-heterogeneity-as-a-universal-driver-of-species-1jab0wmnve.pdf

Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales

Aim In a selected literature survey we reviewed studies on the habitat heterogeneity–animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support.

Location  World-wide.

Methods  We reviewed 85 publications for the period 1960–2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study.

Main conclusions  The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper, we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to ‘keystone structures’ that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.0305-0270.2003.00994.x

Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Human activities have fundamental impacts on the distribution of species through altered land use, but also directly by dispersal of propagules. Rare long-distance dispersal events have a disproportionate importance for the spread of species including invasions. While it is widely accepted that humans may act as vectors of long-distance dispersal, there are few studies that quantify this process. We studied in detail a mechanism of human-mediated dispersal (HMD). For two plant species we measured, over a wide range of distances, how many seeds are carried by humans on shoes. While over half of the seeds fell off within 5 m, seeds were regularly still attached to shoes after 5 km. Semi-mechanistic models were fitted, and these suggested that long-distance dispersal on shoes is facilitated by decreasing seed detachment probability with distance. Mechanistic modelling showed that the primary vector, wind, was less important as an agent of long-distance dispersal, dispersing seeds less than 250 m. Full dispersal kernels were derived by combining the models for primary dispersal by wind and secondary dispersal by humans. These suggest that walking humans can disperse seeds to very long distances, up to at least 10 km, and provide some of the first quantified dispersal kernels for HMD.

Human-mediated dispersal of seeds over long distances

Aim In a selected literature survey we reviewed studies on the habitat heterogeneity–animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support. Location World-wide. Methods We reviewed 85 publications for the period 1960–2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study. Main conclusions The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper, we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to ‘keystone structures’ that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.

ARTICLE Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Human-mediated dispersal is known as an important driver of long-distance dispersal for plants but underlying mechanisms have rarely been assessed. Road corridors function as routes of secondary dispersal for many plant species but the extent to which vehicles support this process remains unclear. In this paper we quantify dispersal distances and seed deposition of plant species moved over the ground by the slipstream of passing cars. We exposed marked seeds of four species on a section of road and drove a car along the road at a speed of 48 km/h. By tracking seeds we quantified movement parallel as well as lateral to the road, resulting dispersal kernels, and the effect of repeated vehicle passes. Median distances travelled by seeds along the road were about eight meters for species with wind dispersal morphologies and one meter for species without such adaptations. Airflow created by the car lifted seeds and resulted in longitudinal dispersal. Single seeds reached our maximum measuring distance of 45 m and for some species exceeded distances under primary dispersal. Mathematical models were fit to dispersal kernels. The incremental effect of passing vehicles on longitudinal dispersal decreased with increasing number of passes as seeds accumulated at road verges. We conclude that dispersal by vehicle airflow facilitates seed movement along roads and accumulation of seeds in roadside habitats. Dispersal by vehicle airflow can aid the spread of plant species and thus has wide implications for roadside ecology, invasion biology and nature conservation.

/pdf/human-mediated-dispersal-of-seeds-by-the-airflow-of-vehicles-3qalvesmx5.pdf

Human-mediated dispersal of seeds by the airflow of vehicles.

Dispersal is critical step in plant invasions but there is limited information about human-mediated long distance seed dispersal, including in protected areas. Seed dispersal by hikers was quantified for five invasive species (the native Acaena novae-zelandiae, and the non-native weeds Rumex acetosella
 Anthoxanthum odoratum, Dactylis glomerata and Festuca rubra) in part of Australia’s Kosciuszko National Park. The proportion of seeds remaining attached to trousers and socks was quantified for replicated short (150 m) and long (5,000 m) distance walks. Functions were fitted for each dataset, and parameters compared among species and between trousers and socks. Dispersal data were combined with attachment rates and the number of people undertaking walks to estimate the total number of weed seeds that might be dispersed. The power exponential function gave the best fit for the majority of datasets, indicating that detachment probability decreased with distance. Seeds of all five species were more tightly attached to socks than trousers, with some seeds still present on socks at 5,000 m. Anthoxanthum and Acaena seeds were more tightly attached to clothing than the other species. Theoretically 1.9 million seeds could be dispersed on socks or 2.4 million seeds on trousers through a season but the actual numbers are likely to be much lower because of limited weed seed at the start of the walks. Because of differences in attachment and detachment rates, seeds from Acaena were more likely to be dispersed longer distances. Long distance human-mediated seed dispersal is potentially a major cause of spread of invasive weeds into protected areas that favours some invasive species over others.

Matthias C. Wichmann

Papers

Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Human-mediated dispersal of seeds over long distances

ARTICLE Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Human-mediated dispersal of seeds by the airflow of vehicles.

Estimating human-mediated dispersal of seeds within an Australian protected area