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

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Modern Applied Statistics With S

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non-habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal-sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal-sized sample sites rather than for unequal-sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch-scale and landscape-scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/jbi.12130

Rethinking patch size and isolation effects: the habitat amount hypothesis

For this article, I reviewed empirical studies finding significant ecological responses to habitat fragmentation per se—in other words, significant responses to fragmentation independent of the effects of habitat amount (hereafter referred to as habitat fragmentation) I asked these two questions: Are most significant responses to habitat fragmentation negative or positive? And do particular attributes of species or landscapes lead to a predominance of negative or positive significant responses? I found 118 studies reporting 381 significant responses to habitat fragmentation independent of habitat amount Of these responses, 76% were positive Most significant fragmentation effects were positive, irrespective of how the authors controlled for habitat amount, the measure of fragmentation, the taxonomic group, the type of response variable, or the degree of specialization or conservation status of the species or species group No support was found for predictions that most significant responses to fragmenta

Ecological Responses to Habitat Fragmentation Per Se

Globally, forests represent highly productive ecosystems that act as carbon sinks where soil organic matter is formed from residuals after biomass decomposition as well as from rhizodeposited carbon. Forests exhibit a high level of spatial heterogeneity and the importance of trees, the dominant primary producers, for their structure and functioning. Fungi, bacteria and archaea inhabit various forest habitats: foliage, the wood of living trees, the bark surface, ground vegetation, roots and the rhizosphere, litter, soil, deadwood, rock surfaces, invertebrates, wetlands or the atmosphere, each of which has its own specific features, such as nutrient availability or temporal dynamicy and specific drivers that affect microbial abundance, the level of dominance of bacteria or fungi as well as the composition of their communities. However, several microorganisms, and in particular fungi, inhabit or even connect multiple habitats, and most ecosystem processes affect multiple habitats. Forests are dynamic on a broad temporal scale with processes ranging from short-term events over seasonal ecosystem dynamics to long-term stand development after disturbances such as fires or insect outbreaks. The understanding of these processes can be only achieved by the exploration of the complex 'ecosystem microbiome' and its functioning using focused, integrative microbiological and ecological research performed across multiple habitats.

Forest microbiome: diversity, complexity and dynamics

The colonization-extinction dynamics of many species are affected by the dynamics of their patches. For increasing our understanding of the metapopulation dynamics of sessile species confined to dynamic patches, we fitted a Bayesian incidence function model extended for dynamic landscapes to snapshot data on five epiphytic lichens among 2083 mapped oaks (dynamic patches). We estimate the age at which trees become suitable patches for different species, which defines their niche breadth (number of suitable trees). We show that the colonization rates were generally low, but increased with increasing connectivity in accordance with metapopulation theory. The rates were related to species traits, and we show, for the first time, that they are higher for species with wide niches and small dispersal propagules than for species with narrow niches or large propagules. We also show frequent long-distance dispersal in epiphytes by quantifying the relative importance of local dispersal and background deposition of dispersal propagules. Local stochastic extinctions from intact trees were negligible in all study species, and thus, the extinction rate is set by the rate of patch destruction (tree fall). These findings mean that epiphyte metapopulations may have slow colonization-extinction dynamics that are explained by connectivity, species traits, and patch dynamics.

https://esajournals.onlinelibrary.wiley.com/doi/pdf/10.1890/11-0760.1

Epiphyte metapopulation dynamics are explained by species traits, connectivity, and patch dynamics

The relative importance of stand and dead wood types for wood-dependent lichens in managed boreal forests

Oaks (Quercus robur) can reach a considerable age, which makes them an important substrate for many epiphytic lichens, including several red-listed species. We studied the importance of tree size and other environmental factors for the occurrence of six epiphytic lichens at two sites, in southeastern Sweden, differing in quality as judged by tree size distribution and number of old trees. The effects of tree circumference, light availability, trunk inclination and site were analysed. Results showed that different lichen species responded differently to these factors, but, overall, tree size was most important for lichen occurrence. Five species showed a positive relation to tree size, but the 50% probability of occurrence was reached at different tree sizes among these species and there were also site differences. This study shows that the maintenance of old trees is crucial for several lichen species, which highlights the importance of long-term management plans.

/pdf/tree-and-site-quality-preferences-of-six-epiphytic-lichens-1el3bzglj8.pdf

Tree and Site Quality Preferences of Six Epiphytic Lichens Growing on Oaks in Southeastern Sweden

This study aimed at comparing six patch connectivity measures by fitting them to field data. We used occupancy data for eight beetle and two pseudoscorpion species from 281 hollow oaks in southeast Sweden. Species occupancy was modelled in relation to tree characteristics and one measure of patch connectivity at a time. For each connectivity measure we searched for the spatial scale that generated the best fit to field data. Connectivity measures that only include occupied patches provided better model fits than those that include all patches. When occupancy data are absent for surrounding habitat patches, information that reflects occurrence probabilities can be included in the connectivity measure. However, in this study incorporation of such information resulted in only a slight improvement of model fit. A frequently used connectivity measure based on the negative exponential function was relatively poor in explaining species’ occurrence; for eight species out of nine a buffer measure was better. A better fit was obtained when the negative exponential function was modified to take into account that habitat patches may ‘‘compete’’ for the immigrants. The spatial scale with the best fit tended to be larger when we used connectivity measures in which dispersal sources are identified with lower precision. Thus, the outcomes from different multiple-scale studies are not directly comparable if the density of dispersal sources is not measured in the same way. Overall we conclude that buffer measures are useful, as they give good predictions and are easy to understand and use. If a biologically more realistic measure is needed, one that up-weights the closest patches should be used. Finally, the possibility that habitat patches may compete with each other for immigrants should be considered when selecting a connectivity measure.

https://www.glel.carleton.ca/PDF/landPub/10/10RaniusetalEcography.PDF

A comparison of patch connectivity measures using data on invertebrates in hollow oaks

We modelled presence/absence per tree of beetles and pseudoscorpions living in tree hollows in relation to trunk circumference, habitat openness, and connectivity (= density of hollow oaks in the surrounding area), using data from 281 oaks. The presence/absence models were then used to predict species’ occurrences in a county (11,600 km2) in southeastern Sweden. For eight of the nine species, the most parsimonious occupancy model included a positive relationship with connectivity and at least one tree characteristic. Occupancy underestimates from occurrence records—the ratio of the area of occupancy based on our predictive model to the area of occupancy based on occurrence records—varied between 3 and 83 among species when using occurrence records up to 1993, with significantly larger underestimates for smaller beetle species. Today (after extensive surveys), underestimation has decreased to 1.3–25, confirming that calculations solely based on species occurrence records greatly underestimate the area of occupancy. We suggest this should be taken into account to a greater extent and in a clearer way than today when constructing red lists. The radius of the connectivity measure that generated the best fit varied between 135 and 2,857 m among species, with longer distances for more threatened species. Consequently, preservation of the most threatened species (Elater ferrugineus and Tenebrio opacus) requires conservation efforts at larger spatial scales than required to protect Osmoderma eremita, which frequently has been used as an indicator and umbrella species.

/pdf/predicting-spatial-occurrence-of-beetles-and-pseudoscorpions-3l0ady9fr1.pdf

Victor Johansson

Papers

Epiphyte metapopulation dynamics are explained by species traits, connectivity, and patch dynamics

The relative importance of stand and dead wood types for wood-dependent lichens in managed boreal forests

Tree and Site Quality Preferences of Six Epiphytic Lichens Growing on Oaks in Southeastern Sweden

A comparison of patch connectivity measures using data on invertebrates in hollow oaks

Predicting spatial occurrence of beetles and pseudoscorpions in hollow oaks in southeastern Sweden