The sensory ecology of adaptive landscapes

In complex environments, behavioural plasticity depends on the ability of an animal to integrate numerous sensory stimuli. The multidimensionality of factors interacting to shape plastic behaviour means it is difficult for both organisms and researchers to predict what constitutes an adaptive response to a given set of conditions. Although researchers may be able to map the fitness pay-offs of different behavioural strategies in changing environments, there is no guarantee that the study species will be able to perceive these pay-offs. We thus risk a disconnect between our own predictions about adaptive behaviour and what is behaviourally achievable given the umwelt of the animal being studied. This may lead to erroneous conclusions about maladaptive behaviour in circumstances when the behaviour exhibited is the most adaptive possible given sensory limitations. With advances in the computational resources available to behavioural ecologists, we can now measure vast numbers of interactions among behaviours and environments to create adaptive behavioural surfaces. These surfaces have massive heuristic, predictive and analytical potential in understanding adaptive animal behaviour, but researchers using them are destined to fail if they ignore the sensory ecology of the species they study. Here, we advocate the continued use of these approaches while directly linking them to perceptual space to ensure that the topology of the generated adaptive landscape matches the perceptual reality of the animal it intends to study. Doing so will allow predictive models of animal behaviour to reflect the reality faced by the agents on adaptive surfaces, vastly improving our ability to determine what constitutes an adaptive response for the animal in question.     

Towards a definition of ecological disturbance

The terms disturbance, perturbation, and stress have been used in various ecological contexts, often synonymously, inconsistently and ambiguously. Consequently, the meaning of these terms lack any ecological rigor upon which to construct a coherent theory of ecosystem response to disturbance. Herein are some of the semantic and conceptual problems involved in defining disturbance, perturbation and stress, and proposals of working definitions as a basis for further discussion.     

The ecology of visual landscapes: Exploring the conceptual common ground of visual and ecological landscape indicators

This paper explores the conceptual common ground between visual and ecological landscape indicators. Indicators can be used to summarise complex information about landscape functions and the ability to extract a common set of indicators for analysing different landscape functions may provide valuable support for multiple-use planning.The development of landscape ecological indicators has been a very active research field that has resulted in a wide range of landscape metrics and composite indices with a strong conceptual base in landscape ecological principles. For the visual aspects of landscapes, however, this conceptual base is often missing and thus hindering progress in the development of indicators. This paper presents the results of an analysis of the correspondence between ecological indicators and visual indicators in order to explore whether there is common ground in both concept and operation. The conceptual level is presented for both ecological and visual indicators, as it is the concepts that define their aim and scope. We feel that transparency over this issue is crucial to the development and use of environmental indicators. The approach identified a candidate set of indicators that capture important aspects of both ecological and visual quality. The strength of the approach is that it forces us to focus on the identification of what we wish to indicate by means of landscape theory and assessment that are relevant to a specific landscape context. We believe that the approach presented here forms the basis for development of new methods for understanding the impact of landscape change in a management and planning context.     

Towards a general theory of adaptive walks on rugged landscapes

Adaptive evolution, to a large extent, is a complex combinatorial optimization process. In this article we take beginning steps towards developing a general theory of adaptive "walks" via fitter variants in such optimization processes. We introduce the basic idea of a space of entities, each a 1-mutant neighbor of many other entities in the space, and the idea of a fitness ascribed to each entity. Adaptive walks proceed from an initial entity, via fitter neighbors, to locally or globally optimal entities that are fitter than their neighbors. We develop a general theory for the number of local optima, lengths of adaptive walks, and the number of alternative local optima accessible from any given initial entity, for the baseline case of an uncorrelated fitness landscape. Most fitness landscapes are correlated, however. Therefore we develop parts of a universal theory of adaptation on correlated landscapes by adaptive processes that have sufficient numbers of mutations per individual to "jump beyond" the correlation lengths in the underlying landscape. In addition, we explore the statistical character of adaptive walks in two independent complex combinatorial optimization problems, that of evolving a specific cell type in model genetic networks, and that of finding good solutions to the traveling salesman problem. Surprisingly, both show similar statistical features, encouraging the hope that a general theory for adaptive walks on correlated and uncorrelated landscapes can be found. In the final section we explore two limits to the efficacy of selection. The first is new, and surprising: for a wide class of systems, as the complexity of the entities under selection increases, the local optima that are attainable fall progressively closer to the mean properties of the underlying space of entities. This may imply that complex biological systems, such as genetic regulatory systems, are "close" to the mean properties of the ensemble of genomic regulatory systems explored by evolution. The second limit shows that with increasing complexity and a fixed mutation rate, selection often becomes unable to pull an adapting population to those local optima to which connected adaptive walks via fitter variants exist. These beginning steps in theory development are applied to maturation of the immune response, and to the problem of radiation and stasis. Despite the limitations of the adaptive landscape metaphor, we believe that further development along the lines begun here will prove useful.     

Human behavioral ecology: I

New fields of inquiry rarely spring fully grown from the forehead of a single genius, and in addition, it is often difficult to decide when a related set of inquiries has coalesced sufficiently to define a field. As measured by the solicitation and publication of review articles, human behavioral ecology has recently become a self-conscious field, for this is the third essay to review it^1,2 and a book-length survey will appear later this year.³ In this two-part article, I will try to give readers a sense of the field by outlining its theoretical and methodological principles and key issues and by summarizing representative studies and unresolved questions in three main topical areas: subsistence strategies (Part I) and reproductive strategies and social interactions (Part II).     

Human behavioral ecology: II

Human behavioral ecology is an interdisciplinary field of study applying theory from evolutionary ecology to a variety of anthropological questions. In Part I of this essay,¹ which appeared in an earlier issue, I surveyed key theoretical and methodological elements of the field, and summarized representative studies and issues in the topical area of subsistence strategies. In Part II, I turn to studies of reproductive strategies, and those analyzing patterns of cooperation and competition in an ecological and adaptive framework. I conclude with a brief look at possible future developments in the field.     

Towards a general formalization of encounter rates in ecology

Although encounters between organisms are fundamental to many ecological processes, a general theory of encounters that accounts for random movements and probabilistic events has yet to be proposed. We present a framework for examining probabilistic encounters between arbitrarily moving searchers and immobile targets in continuous space and time. We define and contrast first encounter rates and mean encounter rates, which are generally not equal and depend on several properties of the process, including movement behaviors, the spatial scales of the encounter kernel, spatial distribution and birth–death dynamics of targets, and whether the encounters are destructive. Based on these considerations, we propose a taxonomy of encounter processes and discuss their functional relationships. Analytical approximations in several special cases are derived, leading to inference about general patterns. We identify, for example, cases (nondestructive, mean encounters) in which encounter rates are completely independent of movement velocity or tortuosity, and we quantify the dependence for cases (e.g., hard, first encounters and destructive encounters in a dynamic landscape) where there is a relationship. The analytical results lead to general qualitative conclusions, while the mathematical formalization and taxonomic organization provides a framework for studying and contrasting a broad range of encounter processes in ecology.     

Population genetics meets behavioral ecology

Populations are often composed of more than just randomly mating subpopulations - many organisms from social groups with distinct patterns of mating and dispersal. Such patterns have recieved much attention in behavioral ecology, yet theories of population genetics rarely take social structures into account. Consequently, population geneticists often report high levels of apparent in breeding and concomitantly low efective sizes, even for species that avoid mating between close kin. Recently, a view of gene dynamics has been introduced that takes dispersal and social structure into account. Accounting for social structure in population genetics leads to a different perspective on how genetic variation is partitoned and the rate at which genic diversity is lost in natural populations - a view that is more consistent with observed behaviors for the minimization of inbreeding.     

Towards an ecology of protective coloration

The strategies underlying different forms of protective coloration are well understood but little attention has been paid to the ecological, life-history and behavioural circumstances under which they evolve. While some comparative studies have investigated the ecological correlates of aposematism, and background matching, the latter particularly in mammals, few have examined the ecological correlates of other types of protective coloration. Here, we first outline which types of defensive coloration strategies may be exhibited by the same individual; concluding that many protective coloration mechanisms can be employed simultaneously, particularly in conjunction with background matching. Second, we review the ecological predictions that have been made for each sort of protective coloration mechanism before systematically surveying phylogenetically controlled comparative studies linking ecological and social variables to antipredator defences that involve coloration. We find that some a priori predictions based on small-scale empirical studies and logical arguments are indeed supported by comparative data, especially in relation to how illumination affects both background matching and self-shadow concealment through countershading; how body size is associated with countershading, motion dazzle, flash coloration and aposematism, although only in selected taxa; how immobility may promote background matching in ambush predators; and how mobility may facilitate motion dazzle. Examination of nearly 120 comparative tests reveals that many focus on ecological variables that have little to do with predictions derived from antipredator defence theory, and that broad-scale ecological studies of defence strategies that incorporate phylogenetics are still very much in their infancy. We close by making recommendations for future evolutionary ecological research.     

The behavioral ecology of mountain baboons

Chacma baboons (Papio ursinus)were studied in a mountain habitat where the effects of high altitude and latitude combine to produce conditions as harsh as those experienced by the desert or hamadryas baboon (P. hamadryas).The population density was as low as that of hamadryas baboons. A survey of populations at altitudes between 1400 and 3000 m showed a strong negative correlation between altitude and group size, with the highest-living groups averaging just 13 individuals and, like hamadryas baboons, seasonally retreating from marginal habitat on the fringes of the range. Foraging activities in these groups relied heavily on the underground storage organs of plants and other items that were time-consuming to find, harvest, and process, placing severe constraints on the time budget. High-altitude and low-altitude groups were nevertheless able to maintain similar activity budgets. This is explicable through an interaction between the patterns of foraging and range usage and observed altitude differences in group size, population density, and home-range size. The behavior of mountain baboons provides insights into ecological effects on behavior both through local altitudinal variation and through similarities to other populations inhabiting marginal environments, notably P. hamadryas.Mountain baboons may represent a significant southern highland population which does not fit into the neat socioecological dichotomy of desert versus savannah baboons.     

Towards an integration of ecological stoichiometry and the metabolic theory of ecology to better understand nutrient cycling

Ecologists have long recognized that species are sustained by the flux, storage and turnover of two biological currencies: energy, which fuels biological metabolism and materials (i.e. chemical elements), which are used to construct biomass. Ecological theories often describe the dynamics of populations, communities and ecosystems in terms of either energy (e.g. population-dynamics theory) or materials (e.g. resource-competition theory). These two classes of theory have been formulated using different assumptions, and yield distinct, but often complementary predictions for the same or similar phenomena. For example, the energy-based equation of von Bertalanffy and the nutrient-based equation of Droop both describe growth. Yet, there is relatively little theoretical understanding of how these two distinct classes of theory, and the currencies they use, are interrelated. Here, we begin to address this issue by integrating models and concepts from two rapidly developing theories, the metabolic theory of ecology and ecological stoichiometry theory. We show how combining these theories, using recently published theory and data along with new theoretical formulations, leads to novel predictions on the flux, storage and turnover of energy and materials that apply to animals, plants and unicells. The theory and results presented here highlight the potential for developing a more general ecological theory that explicitly relates the energetics and stoichiometry of individuals, communities and ecosystems to subcellular structures and processes. We conclude by discussing the basic and applied implications of such a theory, and the prospects and challenges for further development.     

Towards a more plant physiological perspective on soil ecology

Soil respiration almost balances carbon fixation by terrestrial photosynthesis and exceeds all anthropogenic carbon emissions by an order of magnitude, yet we lack precise knowledge of the sources of, and controls upon, the release of carbon dioxide from soils. Here, we discuss the increasing evidence that half of this carbon release is from living plant roots, their mycorrhizal fungi and other root-associated microbes, and that this release is driven directly by recent photosynthesis. The new studies challenge the widespread view that soil activity is dominated by decomposer organisms using older detrital material and that root litter inputs equal those of aboveground litter. The new observations emphasize the physiological continuity and dynamic interdependence of the plant-microbe-soil system and highlight the need for closer cooperation between plant and soil scientists.     

The behavioral ecology of threshold evolution in a polyphenic beetle

Facultative expression of alternative male morphologies is thoughtto allow individual males to select the phenotype with the highestfitness gain given their competitive status relative to othermales with which they compete for females. Choice of, or switchingbetween, morphs commonly relies on developmental threshold responses.Evolutionary changes in developmental threshold responses arethought to provide an important avenue for phenotypic diversificationand the evolution of morphological and behavioral novelties.However, the extent to which alternative male phenotypes andtheir underlying threshold responses actually evolve in naturalpopulations is unclear. Likewise, the ecological factors thatshape the evolution of threshold responses in natural populationsare unexplored for most organisms, as are the consequences ofsuch modifications for patterns of morphological diversity.I examined the ecological basis of rapid threshold evolutionin exotic populations of the horn-polyphenic dung beetle Onthophagustaurus. Male O. taurus vary continuously in body size as a functionof larval feeding conditions. Only males that exceed a criticalthreshold body size develop a pair of long horns on their heads,whereas males below this threshold remain hornless. Populationsin two exotic ranges of this species, the eastern United Statesand western Australia, have diverged in the mean threshold bodysize, which has resulted in the evolution of highly divergentand novel horn length–body size allometries in these populations.Populations in a third and previously unstudied exotic rangeof O. taurus in eastern Australia exhibit threshold body sizesroughly intermediate between the eastern U.S. and western Australianpopulations. I tested three hypothesis to explain how differencesin ecological and demographic factors can drive allometric divergencesbetween populations, using data derived from comparative, standardizedsampling of a large number of populations in each exotic range.Results suggest that differences in the intensity of both intra-and interspecific competition have contributed to the evolutionof divergent thresholds in these populations. My results donot support the hypothesis that shifts in threshold body sizesto larger body sizes are a consequence of increases in the meanbody size of competing males. I discuss my results in the contextof Onthophagus mating systems and the evolutionary implicationsof threshold evolution.     

The behavioral ecology of a cognitive constraint: limited attention

Limited attention may constrain animal behavior in situationsin which the rate of relevant information exceeds the thresholdprocessing capacity of the brain. In the present study, we examinewhy attention is limited by quantifying how attention affectsthe ubiquitous problem of balancing foraging and antipredatoractivity. We analyze how a given attentional capacity affectsfeeding requirements, the optimal attentional focus during predatorscanning, and the probability of detecting predators. Our modelindicates that because of the complex interplay between thecosts and benefits associated with a given attentional capacity,limited attention can be an optimal strategy, which allows effectiveand economical search for cryptic objects.     

Towards a multi‐trophic extension of metacommunity ecology

Metacommunity theory provides an understanding of how spatial processes determine the structure and function of communities at local and regional scales. Although metacommunity theory has considered trophic dynamics in the past, it has been performed idiosyncratically with a wide selection of possible dynamics. Trophic metacommunity theory needs a synthesis of a few influential axis to simplify future predictions and tests. We propose an extension of metacommunity ecology that addresses these shortcomings by incorporating variability among trophic levels in ‘spatial use properties’. We define ‘spatial use properties’ as a set of traits (dispersal, migration, foraging and spatial information processing) that set the spatial and temporal scales of organismal movement, and thus scales of interspecific interactions. Progress towards a synthetic predictive framework can be made by (1) documenting patterns of spatial use properties in natural food webs and (2) using theory and experiments to test how trophic structure in spatial use properties affects metacommunity dynamics.