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

Hypothesis-testing methods for multivariate data are needed to make rigorous probability statements about the effects of factors and their interactions in experiments. Analysis of variance is particularly powerful for the analysis of univariate data. The traditional multivariate analogues, however, are too stringent in their assumptions for most ecological multivariate data sets. Non-parametric methods, based on permutation tests, are preferable. This paper describes a new non-parametric method for multivariate analysis of variance, after McArdle and Anderson (in press). It is given here, with several applications in ecology, to provide an alternative and perhaps more intuitive formulation for ANOVA (based on sums of squared distances) to complement the description pro- vided by McArdle and Anderson (in press) for the analysis of any linear model. It is an improvement on previous non-parametric methods because it allows a direct additive partitioning of variation for complex models. It does this while maintaining the flexibility and lack of formal assumptions of other non-parametric methods. The test- statistic is a multivariate analogue to Fisher's F-ratio and is calculated directly from any symmetric distance or dissimilarity matrix. P-values are then obtained using permutations. Some examples of the method are given for tests involving several factors, including factorial and hierarchical (nested) designs and tests of interactions.

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A new method for non-parametric multivariate analysis of variance

For centuries, biologists have studied patterns of plant and animal diversity at continental scales. Until recently, similar studies were impossible for microorganisms, arguably the most diverse and abundant group of organisms on Earth. Here, we present a continental-scale description of soil bacterial communities and the environmental factors influencing their biodiversity. We collected 98 soil samples from across North and South America and used a ribosomal DNA-fingerprinting method to compare bacterial community composition and diversity quantitatively across sites. Bacterial diversity was unrelated to site temperature, latitude, and other variables that typically predict plant and animal diversity, and community composition was largely independent of geographic distance. The diversity and richness of soil bacterial communities differed by ecosystem type, and these differences could largely be explained by soil pH (r(2) = 0.70 and r(2) = 0.58, respectively; P < 0.0001 in both cases). Bacterial diversity was highest in neutral soils and lower in acidic soils, with soils from the Peruvian Amazon the most acidic and least diverse in our study. Our results suggest that microbial biogeography is controlled primarily by edaphic variables and differs fundamentally from the biogeography of "macro" organisms.

/pdf/the-diversity-and-biogeography-of-soil-bacterial-communities-1lkjoz8774.pdf

The diversity and biogeography of soil bacterial communities

Physiological and ecological constraints play key roles in the evolution of plant growth patterns, especially in relation to defenses against herbivores. Phenotypic and life history theories are unified within the growth-differentiation balance (GDB) framework, forming an integrated system of theories explaining and predicting patterns of plant defense and competitive interactions in ecological and evolutionary time. Plant activity at the cellular level can be classified as growth (cell division and enlargement) of differentiation (chemical and morphological changes leading to cell maturation and specialization). The GDB hypothesis of plant defense is premised upon a physiological trade-off between growth and differentiation processes. The trade-off between growth and defense exists because secondary metabolism and structural reinforcement are physiologically constrained in dividing and enlarging cells, and because they divert resources from the production of new leaf area. Hence the dilemma of plants: Th...

https://www.jstor.org/stable/pdfplus/2830650.pdf

The dilemma of plants: To grow or defend.

Identification of the selective forces on plant dispersal engenders theoretical argument, empirical study, and speculation. We separate evidence, testable hypotheses, and conjecture surrounding two major questions in dispersal ecology. The first asks what ecological, and ultimately evolutionary, advantages exist in seed dispersal. Astonishingly little is known about the advantages to a parent plant that are actually conferred by investment in dispersal structures. Does dispersal enable seeds and ultimately seedlings to escape mortality near the parent? Is continual recolonization of unstable habitats the primary advantage? Must seeds find rare microhabitats suitable for reestablishment? Such issues are addressed through joint consideration of dispersal and establishment-those stages both mediated by parental provisioning and subject to the highest mortality in the life of a plant. The second broad question asks what general and explicit environmental forces influence the timing and mode of dispersal. Do climates or seasons favor one dispersal mode over another? Do differences in number, size, morphology, or nutritional quality of fruits influence frugivore choice, and consequently differential dispersal of species or individuals within species? Studies of dispersal process and mode should be intimately connected. A general objective of this paper is to explore the degree to which they are integrated and, in so doing, to catalyze their union. We emphasize topics most in need of critical attention: the evolutionary ecology of dispersal process and mode. Excellent recent reviews consider such related topics as dispersal mechanism (131, 184), seed dormancy (1, 30), phytogeography (11, 115, 146), masting and predator satiation (105, 156), and succession (68, 69, 189).

https://www.jstor.org/stable/info/2097067

Ecology of Seed Dispersal

Mutualistic interactions between ants and plants involve rewards offered by plants and services performed by ants in a mutually advantageous relationship. The rewards are principally food and/or nest sites, and ants in turn perform a number of services for plants: they disperse and plant seeds; they protect foliage, buds, and reproductive structures from enemies such as herbivores and seed predators; they fertilize plants with essential nutrients; and they may sometimes function as pollinators. In this book, initially published in 1985, Professor Beattie reviews the fascinating natural history of ant–plant interactions, discusses the scientific evidence for the mutualistic nature of these relationships, and reaches some conclusions about the ecological and evolutionary processes that mold them. This important work explores the natural history, experimental approach, and integration with contemporary evolutionary and ecological literature of the time will appeal to a wide variety of biologists.

The Evolutionary Ecology of Ant-Plant Mutualisms.

The evolutionary ecology of ant-plant mutualisms

Environmental monitoring and conservation evaluation in terrestrial habitats may be enhanced by the use of invertebrate inventories, but taxonomic and logistic constraints frequently encountered during conventional taxonomic treatment have greatly restricted their use. To overcome this problem we suggest that nonspecialists may be used to classify invertebrates to morphospecies without compromising scientific accuracy. To test this proposition, large pitfall and litter samples of ants, beetles, and spiders from four forest types were sorted to morphospecies by a nonspecialist and to species by specialists. These data were used to generate morphospecies and species inventories and to estimate richness (α diversity) and turnover (β diversity), information frequently used in the above activities. Our results show that the estimates of richness of ants and spiders varied little between morphospecies and species inventories. Differences between estimates of beetle richness were largely influenced by errors of identification in two families, Curculionidae and Staphylinidae. But morphospecies and species inventories yielded identical ranking of forest type using richness. Turnover was assessed by sample ordination, which revealed similar clusters regardless of the type of inventory. Analysis of similarities of assemblages of ants and beetles showed significant differences between all forest types. Spider assemblages showed a lower level of discrimination. The assessment of turnover was consistent among inventories but different between the major taxa. Our findings suggest that morphospecies may be used as surrogates for species in some environmental monitoring and conservation, in particular when decisions are guided by estimates of richness and the assessment of turnover.



El monitoreo ambiental y la evaluacion para la conservacion en los habitats terrestres puede ser mejorado con el uso de inventarios de invertebrados. Sin embargo, las restricciones taxonomicas y logisticas que surgen frecuentemente durante los tratamientos taxonomicos convencionales, han restringido en gran medida su uso. Para poder superar este problema, nosotros sugerimos que podria utilizarse personal no especializado para clasificar invertebrados en morfoespecies sin comprometer la exactitud cientifica. Para probar esta propuesta, muestras de hormigas, escarabajos y aranas provenientes de cuatro tipos de bosques fueron ordenadas en morfoespecies por un noespecialista y en especies por un especialista. Estos datos fueron utilizados para generar inventarios de morfoespecies y especies y para estimar riqueza (diversidad alfa) y tasas de renovacion (diversidad beta), informacion que es utilizada frecuentemente en las actividades mencionadas anteriormente. Nuestros resultados muestran que las estimaciones de la riqueza de hormigas y aranas vario muy poco entre los inventarios de morfoespecies y de especies. Las diferencias entre las estimaciones de la riqueza de escarabajos fue en gran medida influenciada por errores en la identificacion de dos familias, Curculionidae y Staphylinidae. Sin embargo, los inventarios de morfoespecies y especies produjeron un ordenamiento identico de los tipos de bosques utilizando la riqueza como parametro. La tasa de intercambio (“turnover”) fue evaluada por medio de metodos de ordenacion de las muestras que revelaron grupos muy similares independientemente del tipo de inventario. Los analisis de similitud de los agrupamientos de hormigas y escarabajos mostraron diferencias significativas entre todos los tipos de bosques. Los agrupamientos de aranas mostraron un bajo nivel de discriminacion. La evaluacion de la tasa de recambio fue consistente entre los inventarios pero diferente entre los taxones mas importantes. Nuestros resultados sugieren que las morfoespecies podrian ser utilizadas como substitutos de las especies en ciertos casos de monitoreos ambientales y de conservacion, en particular, cuando las decisiones estan guiadas por las estimaciones de riqueza y la evaluacion de la tasa de intercambio.

Invertebrate Morphospecies as Surrogates for Species: A Case Study

Patterns in the spatial distribution of organisms provide important information about mechanisms that regulate the diversity of life and the complexity of ecosystems. Although microorganisms may comprise much of the Earth's biodiversity and have critical roles in biogeochemistry and ecosystem functioning, little is known about their spatial diversification. Here we present quantitative estimates of microbial community turnover at local and regional scales using the largest spatially explicit microbial diversity data set available (> 10(6) sample pairs). Turnover rates were small across large geographical distances, of similar magnitude when measured within distinct habitats, and did not increase going from one vegetation type to another. The taxa-area relationship of these terrestrial microbial eukaryotes was relatively flat (slope z = 0.074) and consistent with those reported in aquatic habitats. This suggests that despite high local diversity, microorganisms may have only moderate regional diversity. We show how turnover patterns can be used to project taxa-area relationships up to whole continents. Taxa dissimilarities across continents and between them would strengthen these projections. Such data do not yet exist, but would be feasible to collect.

Spatial scaling of microbial eukaryote diversity

We investigated three procedures that may lead to rapid and accurate assessment of epigaeic arthropod biodiversity. They are: (1) the identification of taxa whose diversity is correlated with that of others: (2) the identification of times and methods of sampling that produce estimates of diversity representative of more intensive sampling; and (3) the use of morphospecies inventories generated by non-specialists. Ants, beetles, and spiders were sampled from four forest types, in three seasons, using two collecting methods: pitfall trapping and extraction from litter. Specimens were sorted by a non-specialist to morphospecies and by specialist taxonomists to species. Richness (α-diversity) and turnover (β-diversity) were compared for different sampling regimes using morphospecies and species inventories. We found no significant positive correlations between ant, beetle, and spider species richness but there was a strong negative correlation between ant and beetle richness. For beetles alone, richness within the families Carabidae, Scarabaeidae, and Pselaphidae (i.e, avoiding taxonomically problematic families) was significantly correlated with richness within all other families. Assessment of turnover revealed that: (1) the four forest types contained significantly different assemblages of ants and beetles but not spiders and 92) the four forests were less clearly discriminated using species from the three beetle families Carabidae, Scarabaeidae, and Pselaphidae when compared to species from all beetle families pooled. Analyses of single sampling periods and methods revealed that summer and spring pitfall samples were most representative of more intensive sampling. That is: (1) the richness of ants and beetles in these samples was significantly positively correlated with the richness of all other samples and 92) turnover of beetles and ants among the four forests revealed by summer pitfall samples was similar to turnover using all samples. The three beetle surrogate families recorded by pitfall samples in spring, and to a lesser extent summer, showed significant correlations in richness with all other beetle species recorded in the same samples. However, the assessment of turnover was less accurate when only surrogate families were used. The most accurate and cost-effective assessment of turnover was generated by a summer pitfall sample in which data for ants, carabid, and scarab beetles were combined and analyzed as a single data set. Results were largely consistent regardless of whether species or morphospecies were used, which suggests that monitoring and assessment of terrestrial invertebrate biodiversity may be achieved by the careful use of morphospecies. Our results also suggest those invertebrate taxa, sampling methods, and sampling periods that yield the most consistent and reliable assessment of epigaeic invertebrate biodiversity in Australian temperate hardwood forests. However, empirical studies that follow the protocols discussed in this paper are urgently required in different environments. These studies may point the way to more representative monitoring and assessment of terrestrial biodiversity.

https://www.jstor.org/stable/info/2269394

Andrew J. Beattie

Papers

The Evolutionary Ecology of Ant-Plant Mutualisms.

The evolutionary ecology of ant-plant mutualisms

Invertebrate Morphospecies as Surrogates for Species: A Case Study

Spatial scaling of microbial eukaryote diversity

Designing a Cost‐Effective Invertebrate Survey: A Test of Methods for Rapid Assessment of Biodiversity