Social selection offers an alternative to sexual selection by reversing its logic. Social selection starts with offspring production and works back to mating, and starts with behavioural dynamics and works up to gene pool dynamics. In social selection, courtship can potentially be deduced as a negotiation, leading to an optimal allocation of tasks during offspring rearing. Ornaments facilitate this negotiation and also comprise 'admission tickets' to cliques. Mating pairs may form 'teams' based on the reciprocal sharing of pleasure. The parent-offspring relation can be managed by the parent considered as the owner of a 'family firm' whose product is offspring. The cooperation in reproductive social behaviour evolves as a mutual direct benefit through individual selection rather than as some form of altruism requiring kin or multi-level selection.
View details for DOI 10.1098/rstb.2011.0282
View details for Web of Science ID 000306193500005
View details for PubMedID 22777017
We model direct fitness benefits of genetic mosaicism for a long-lived tree in coevolution with a short-lived herbivore to test four hypotheses: that mosaicism reduces selection on the herbivore for resistance to plant defenses; that module-level selection allows the individual tree to adapt to its herbivore; and that this benefits the tree population, increasing average tree fitness and reducing local adaptation of the herbivore. We show that: mosaicism does not sufficiently reduce selection for resistance in the herbivore to benefit the tree; that individual trees do benefit from module-level selection when somatic mutation introduces new defenses; and that mosaicism does reduce local adaptation in the herbivore, which increases average tree fitness. These results are robust to varying genetic assumptions of dominance and the somatic mutation rate, but only hold for sufficiently long-lived trees with relatively strong selection. We also show that a mixed reproductive strategy of primarily asexual reproduction interspersed with occasional sexual reproduction is effective in coevolving with the herbivore, as it maintains beneficial allele combinations. Finally, we argue that intraorganismal genetic heterogeneity need not threaten the integrity of the individual and may be adaptive when selection acts concordantly between levels.
View details for DOI 10.1111/j.1558-5646.2011.01500.x
View details for Web of Science ID 000302546700011
View details for PubMedID 22486691
Most of the work in evolutionary game theory starts with a model of a social situation that gives rise to a particular payoff matrix and analyses how behaviour evolves through natural selection. Here, we invert this approach and ask, given a model of how individuals behave, how the payoff matrix will evolve through natural selection. In particular, we ask whether a prisoner's dilemma game is stable against invasions by mutant genotypes that alter the payoffs. To answer this question, we develop a two-tiered framework with goal-oriented dynamics at the behavioural time scale and a diploid population genetic model at the evolutionary time scale. Our results are two-fold: first, we show that the prisoner's dilemma is subject to invasions by mutants that provide incentives for cooperation to their partners, and that the resulting game is a coordination game similar to the hawk-dove game. Second, we find that for a large class of mutants and symmetric games, a stable genetic polymorphism will exist in the locus determining the payoff matrix, resulting in a complex pattern of behavioural diversity in the population. Our results highlight the importance of considering the evolution of payoff matrices to understand the evolution of animal social systems.
View details for DOI 10.1098/rspb.2010.2105
View details for Web of Science ID 000291464200017
View details for PubMedID 21147797
Most biologists implicitly define an individual organism as "one genome in one body." This definition is based on physiological and genetic criteria, but it is problematic for colonial organisms. We propose a definition based instead on the evolutionary criteria of alignment of fitness, export of fitness by germ-soma specialization, and adaptive functional organization. We consider how these concepts apply to various putative individual organisms. We conclude that complex multicellular organisms and colonies of eusocial insects satisfy these three criteria, but that, in most cases (with at least one notable exception), colonies of modular organisms and genetic chimeras do not. While species do not meet these criteria, they may meet the criteria for a broader concept--that of an evolutionary individual--and sexual reproduction may be a species-level exaptation for enhancing evolvability. We also review the costs and benefits of internal genetic heterogeneity within putative individuals, demonstrating that high relatedness is neither a necessary nor a sufficient condition for individuality, and that, in some cases, genetic variability may have adaptive benefits at the level of the whole.
View details for Web of Science ID 000285039800003
View details for PubMedID 21243964
Although much previous work describes evolutionary mechanisms that promote or stabilize different social behaviors, we still have little understanding of the factors that drive animal behavior proximately. Here we present a modeling approach to answer this question. Our model rests on motivations to achieve objectives as the proximate determinants of behavior. We develop a two-tiered framework by first modeling the dynamics of a social interaction at the behavioral time scale and then find the evolutionarily stable objectives that result from the outcomes these dynamics produce. We use this framework to ask whether "other-regarding" motivations, which result from a kind of nonselfish objective, can evolve when individuals are engaged in a social interaction that entails a conflict between their material payoffs. We find that, at the evolutionarily stable state, individuals can be other-regarding in that they are motivated to increase their partners' payoff as well as their own. In contrast to previous theories, we find that such motivations can evolve because of their direct effect on fitness and do not require kin selection or a special group structure. We also derive general conditions for the evolutionary stability of other-regarding motivations. Our conditions indicate that other-regarding motivations are more likely to evolve when social interactions and behavioral objectives are both synergistic.
View details for DOI 10.1073/pnas.0904357106
View details for Web of Science ID 000271637500035
View details for PubMedID 19858492
Previous theoretical work on parental decisions in biparental care has emphasized the role of the conflict between evolutionary interests of parents in these decisions. A prominent prediction from this work is that parents should compensate for decreases in each other's effort, but only partially so. However, experimental tests that manipulate parents and measure their responses fail to confirm this prediction. At the same time, the process of parental decision making has remained unexplored theoretically. We develop a model to address the discrepancy between experiments and the theoretical prediction, and explore how assuming different decision making processes changes the prediction from the theory. MODEL DESCRIPTION: We assume that parents make decisions in behavioral time. They have a fixed time budget, and allocate it between two parental tasks: provisioning the offspring and defending the nest. The proximate determinant of the allocation decisions are parents' behavioral objectives. We assume both parents aim to maximize the offspring production from the nest. Experimental manipulations change the shape of the nest production function. We consider two different scenarios for how parents make decisions: one where parents communicate with each other and act together (the perfect family), and one where they do not communicate, and act independently (the almost perfect family).The perfect family model is able to generate all the types of responses seen in experimental studies. The kind of response predicted depends on the nest production function, i.e. how parents' allocations affect offspring production, and the type of experimental manipulation. In particular, we find that complementarity of parents' allocations promotes matching responses. In contrast, the relative responses do not depend on the type of manipulation in the almost perfect family model. These results highlight the importance of the interaction between nest production function and how parents make decisions, factors that have largely been overlooked in previous models.
View details for DOI 10.1371/journal.pone.0007345
View details for Web of Science ID 000270808300005
View details for PubMedID 19823687
View details for Web of Science ID 000263624900004
Spatial models commonly assume that dispersal does not depend on environmental conditions or phenotype. For example, these assumptions underpin explanations for clines on the basis of a trade-off between dispersal and local adaptation. We reexamine clines when an individual's decisions over whether and where to disperse depend on its fitness. We compare fitness-dependent dispersal with cases where dispersal responds to juvenile survivorship only. Clines are steeper the more responsive dispersal is to environmental conditions for all dispersal behaviors that we consider. Clines eventually become stepped as the responsiveness of dispersal to environmental conditions is increased for half of the dispersal behaviors we consider, but smooth clines are maintained for the remaining cases. Smooth clines are maintained by the biased movement of individuals out of the hybrid zone when individuals move directionally in response to gradients in juvenile survivorship, which is a different mechanism to that maintaining smooth clines in classic cline theory.
View details for DOI 10.1086/591685
View details for Web of Science ID 000260186000005
View details for PubMedID 18828744
View details for Web of Science ID 000260229000007
Anisogamy refers to gametes that differ in size, and characterizes the difference between males and females. The evolution of aniosgamy is widely interpreted as involving conflict between gamete producers with small sperm parasitizing on the investment made by the eggs. Using a population genetic model for evolution at a locus that codes jointly for sperm and egg sizes of a hermaphrodite, we show that the origin of anisogamy in an externally spawning population need not involve conflict between gamete producers. Gamete size dimorphism may be an adaptation that increases gamete encounter rates when large zygotes are selected, and we show this in a mechanistically general individual selection model. We use the Vance survival function without specific allometric assumptions to model the zygote fitness dependence on its size, and hence obtain ecological and life-history correlates of isogamy and anisogamy, which we successfully compare with data from Volvocales.
View details for DOI 10.1016/j.tpb.2008.02.002
View details for Web of Science ID 000256996000001
View details for PubMedID 18485981
View details for Web of Science ID 000252652600001
View details for Web of Science ID 000248192000009
View details for Web of Science ID 000245912900003
The evolution and persistence of biological cooperation have been an important puzzle in evolutionary theory. Here, we suggest a new approach based on bargaining theory to tackle the question. We present a mechanistic model for negotiation of benefits between a nitrogen-fixing nodule and a legume plant. To that end, we first derive growth rates for the nodule and plant from metabolic models of each as a function of material fluxes between them. We use these growth rates as pay-off functions in the negotiation process, which is analogous to collective bargaining between a firm and a workers' union. Our model predicts that negotiations lead to the Nash bargaining solution, maximizing the product of players' pay-offs. This work introduces elements of cooperative game theory into the field of mutualistic interactions. In the discussion of the paper, we argue for the benefits of such an approach in studying the question of biological cooperation.
View details for DOI 10.1098/rspb.2006.3689
View details for Web of Science ID 000242765300004
View details for PubMedID 17015340
View details for Web of Science ID 000237655400005
Theories about sexual selection can be traced back to Darwin in 1871. He proposed that males fertilize as many females as possible with inexpensive sperm, whereas females, with a limited supply of large eggs, select the genetically highest quality males to endow their offspring with superior capabilities. Since its proposal, problems with this narrative have continued to accumulate, and it is our view that sexual selection theory needs to be replaced. We suggest an approach that relies on the exchange of direct ecological benefits among cooperating animals without reference to genetic benefits. This approach can be expressed mathematically in a branch of game theory that pertains to bargaining and side payments.
View details for DOI 10.1126/science.1110105
View details for Web of Science ID 000235456900035
View details for PubMedID 16484485
Spatial models commonly assume that dispersal rates are constant across individuals and environments and that movement directions are unbiased. These random-movement assumptions are inadequate to capture the range of dispersal behaviors revealed in diverse case studies. We examine an alternative assumption of directed movement, in which dispersal is a conditional and directional response by individuals to varying environmental conditions. Specifically, we assume individuals bias their movements to climb spatial fitness gradients. We compare the consequences of random and directed movement for local adaptation, the evolution of dispersal, and the reinforcement process. The implications of each movement strategy depend on the nature of environmental disturbance, and we examine the outcomes for undisturbed environments and with uncorrelated and autocorrelated disturbances. Both movement strategies offer advantages over sedentary life histories by allowing colonization of suitable habitats. However, random movement eventually becomes costly in stable environments because it inhibits local adaptation. In contrast, directed movement accelerates local adaptation. In disturbed environments, random movement offers bet-spreading advantages by distributing offspring across habitats. Despite being a more targeted strategy, an intermediate amount of directed movement provides similar bet-spreading benefits. These fitness consequences have implications for the evolution of dispersal. Dispersiveness is lost by random movers in undisturbed environments, is maintained in polymorphism with infrequent disturbances, and evolves when disturbances are uncorrelated. Directed movement becomes selectively neutral in the absence of disturbance, evolves when disturbances are autocorrelated, and is maintained in polymorphism with uncorrelated disturbances. Disturbance also determines the outcome of the reinforcement process for each strategy. For example, directed movers show no progress toward reinforcement in undisturbed environments, evolve random mating with uncorrelated disturbances, and can evolve assortative mating in infrequently disturbed environments.
View details for Web of Science ID 000232941400003
View details for PubMedID 16405154
An improved understanding of dispersal behavior is needed to predict how populations and communities respond to habitat fragmentation. Most spatial dynamic theory concentrates on random dispersal, in which movement rates depend neither on the state of an individual nor its environment and movement directions are unbiased. We examine the neglected dispersal component of directed movement in which dispersal is a conditional and directional response of individuals to varying environmental conditions. Specifically, we assume that individuals bias their movements along local gradients in fitness. Random movers, unable to track heterogeneous environmental conditions, face source-sink dynamics, which can result in deterministic extinction or increase their vulnerability to stochastic extinction. Directed movers track environmental conditions closely. In fluctuating environments, random movers "spread their bets" across patches, while directed movers invest offspring in habitats currently enjoying propitious conditions. The autocorrelation in the environment determines each strategy's success. Random movers permeate entire landscapes, but directed movers are more geographically constrained. Local information constraints limit the ranges of directed movers and introduce a role for historical contingency in determining their ultimate distribution. These geographic differences have implications for biodiversity. Random movement maintains biodiversity through local coexistence, but directed movement favors a spatial partitioning of species.
View details for Web of Science ID 000227517000007
View details for PubMedID 15791537
Thomas et al. argue, contrary to Sala et al. that climate change poses an equal or greater threat to global biodiversity than land-use change. We contest this claim, however, on the grounds that Thomas et al. incorrectly apply species-area relationships.
View details for DOI 10.1038/nature02717
View details for Web of Science ID 000222356800033
View details for PubMedID 15233130
View details for Web of Science ID 000222174000009
View details for Web of Science ID 000186802200017
The diversity, frequency, and scale of human impacts on coral reefs are increasing to the extent that reefs are threatened globally. Projected increases in carbon dioxide and temperature over the next 50 years exceed the conditions under which coral reefs have flourished over the past half-million years. However, reefs will change rather than disappear entirely, with some species already showing far greater tolerance to climate change and coral bleaching than others. International integration of management strategies that support reef resilience need to be vigorously implemented, and complemented by strong policy decisions to reduce the rate of global warming.
View details for Web of Science ID 000184755900027
View details for PubMedID 12920289
Seemingly intangible ecosystem characteristics that preoccupy ecologists, like ecosystem stability and the responsiveness of populations to environmental variation, have quantifiable economic values. We show how to derive these values, and how their consideration should change environmental decision making. To illustrate these concepts, we use a simple reserve design model. When resource managers choose a particular landscape configuration, their decision affects both the mean abundance of species and the temporal variation in abundances. Population stability and related phenomena have economic value, because management actions affect the variance of ecosystem components. In our example, a larger reserve size is recommended when accounting for the stability of the managed ecosystem.
View details for DOI 10.1073/pnas.0832226100
View details for Web of Science ID 000183493500047
View details for PubMedID 12777632
This article presents a theory of territoriality that integrates optimal foraging and conflict resolution through negotiation. Using a spatially explicit model of a sit-and-wait forager, we show that when resources are scarce, there is a conflict between foragers: there is not enough space for all individuals to have optimal home ranges. We propose that a division of space that solves this conflict over resources is the outcome of a negotiation between foragers. We name this outcome the socially stable territories (SST). Using game theory we show that in a homogenous patch occupied by two interacting foragers, both individuals receive identical energy yields at the socially stable territories; that is, there is economic equity. Economic inequity can arise in a heterogeneous patch or from asymmetries in fighting abilities between the foragers. Opportunity costs play a role in reducing economic inequity. When the asymmetry in fighting abilities is very large, a negotiated division of space is not possible and the forager with lowest fighting ability may be evicted from the habitat patch. A comparison between territories and overlapping home ranges shows that energy yields from territories are generally higher. We discuss why there are instances in which individuals nevertheless overlap home ranges.
View details for Web of Science ID 000180186600011
View details for PubMedID 12650468
On the eve of the World Summit on Sustainable Development, it is timely to assess progress over the 10 years since its predecessor in Rio de Janeiro. Loss and degradation of remaining natural habitats has continued largely unabated. However, evidence has been accumulating that such systems generate marked economic benefits, which the available data suggest exceed those obtained from continued habitat conversion. We estimate that the overall benefit:cost ratio of an effective global program for the conservation of remaining wild nature is at least 100:1.
View details for Web of Science ID 000177325400027
View details for PubMedID 12169718
We introduce a model for the dynamics of a patchy population in a stochastic environment and derive a criterion for its persistence. This criterion is based on the geometric mean (GM) through time of the spatial-arithmetic mean of growth rates. For the population to persist, the GM has to be >/=1. The GM increases with the number of patches (because the sampling error is reduced) and decreases with both the variance and the spatial covariance of growth rates. We derive analytical expressions for the minimum number of patches (and the maximum harvesting rate) required for the persistence of the population. As the magnitude of environmental fluctuations increases, the number of patches required for persistence increases, and the fraction of individuals that can be harvested decreases. The novelty of our approach is that we focus on Malthusian local population dynamics with high dispersal and strong environmental variability from year to year. Unlike previous models of patchy populations that assume an infinite number of patches, we focus specifically on the effect that the number of patches has on population persistence. Our work is therefore directly relevant to patchily distributed organisms that are restricted to a small number of habitat patches.
View details for Web of Science ID 000173879300002
View details for PubMedID 18707409
View details for Web of Science ID 000176072000015
View details for Web of Science ID 000179463800019
View details for Web of Science ID 000170125600001
View details for Web of Science ID 000173048000016
The emerging interdisciplinary field of ecological economics should be a recognized research priority. Only through a combination of sound ecology and good economics can we hope to manage our exploitation of the biosphere in a manner that is both sustainable and efficient. This article is an invitation to ecologists to use economic tools and to participate in ecological economic debate. To this end, we review basic ecological economic concepts and discuss how the field has arisen, what benefits it offers, and what challenges it must overcome.
View details for PubMedID 11301151
View details for Web of Science ID 000086995000003
View details for Web of Science ID 000081746100002
View details for Web of Science ID 000079309300024
View details for Web of Science ID 000079677900006
Although humanity depends on the continued, aggregate functioning of natural ecosystems, few studies have explored the impact of community structure on the stability of aggregate community properties. Here we derive the stability of the aggregate property of community biomass as a function of species' competition coefficients for a two-species model. The model predicts that the stability of community biomass is relatively independent of the magnitude of the interaction strengths. Instead, the degree of asymmetry of the interactions appears to be key to community stability.
View details for Web of Science ID 000074131900049
View details for PubMedID 9618499
View details for Web of Science ID 000073841800022
Intertidal systems have been models for the study of the roles of competition, predation, and disturbance in determining community structure. These systems exhibit considerable regional variability in percentage cover and in the strength of interspecific interactions, which may be due largely to effects of varying larval supply. In Oregon and Washington, experimental studies of space allocation among sessile invertebrates have emphasized the role of benthic processes such as competition and predation. In contrast, studies in central California have emphasized the importance of larval supply. In this article, we identify a gradient in percentage cover in the middle and upper intertidal zone that is consistent with an oceanographically based explanation for these differences: percentage cover of mussels and barnacles is much higher in Oregon, where nearshore circulation promotes high recruitment, than in California, where strong offshore currents inhibit recruitment. A mathematical model incorporating larval transport and interspecific competition for space offers an explanation for the one violation of the hypothesis-higher percentage cover of Chthamalus spp. in California. The findings illustrate that attempts to synthesize regional differences in community structure and dynamics can benefit from considering both the benthic adult and pelagic larval environments.
View details for Web of Science ID 000072752400002
View details for PubMedID 18811323
View details for Web of Science ID 000072166600018
View details for Web of Science ID 000071611800011
View details for Web of Science ID A1997YL29100009
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With the collapse of fisheries in many parts of the world causing widespread economic harm, attention is focused on a possible cause and remedy of fishery collapse. Economic theory for managing a renewable resource, such as a fishery, leads to an ecologically unstable equilibrium as difficult to maintain as balancing a marble on top of a dome. A fishery should be managed for ecological stability instead--in the analogy, as easy to maintain as keeping a marble near the base of a bowl. The goal of ecological stability is achieved if the target stock is above that producing maximum sustainable yield and harvested at less than the maximum sustainable yield. The cost of managing for ecological stability, termed "natural insurance," is low if the fishery is sufficiently productive. This cost is shown to pay for itself over the long term in a variable and uncertain environment. An ecologically stable target stock may be attained either with annually variable quotas following current practice or, preferably, through a market mechanism whereby fish are taxed at dockside if caught when the stock was below target and are untaxed otherwise. In this regulatory environment, the goal of maximizing short-term revenue coincides with the goal of ecological stability, thereby also maximizing long-term revenue. This new approach to fishery management is illustrated with the recently collapsed Newfoundland fishing industry. The Newfoundland cod fishery is expected to rebuild to an ecologically stable level in about 9 years and thereafter support an annual harvest of about 75% of the 1981-1990 average.
View details for Web of Science ID A1996UL25500102
View details for PubMedID 11607680
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Organisms living in the marine rocky intertidal zone compete for space. This, together with predation, physical disruption, and differing species tolerances to physiological stress, explains the structure of the ecological communities at some sites. At other sites the supply of larvae is limiting, and events in the offshore waters, such as wind-driven upwelling, explain the composition of intertidal communities. Whether the community ecology at a site is governed by adult-adult interactions within the site, or by limitations to the supply of larvae reaching the site, is determined by the regional pattern of circulation in the coastal waters. Models combining larval circulation with adult interactions can potentially forecast population fluctuations. These findings illustrate how processes in different ecological habitats are coupled.
View details for Web of Science ID A1988Q047900029
View details for PubMedID 11538249
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View details for Web of Science ID A1987H283600007
Kelp forests along the coast of central California harbor juvenile rockfish that prey on the larvae of invertebrates from the rocky intertidal zone. This predation reduces recruitment to barnacle populations to 1/50 of the level in the absence of fish. The dynamics of the intertidal community are thus strongly coupled to the dynamics of the offshore kelp community.
View details for Web of Science ID A1987F689100031
View details for PubMedID 17810342
View details for Web of Science ID A1986E902500004
View details for Web of Science ID A1986C026600003
View details for Web of Science ID A1985AQH3500019
Field studies demonstrate that the population structure of the barnacle Balanus glandula differs between locations of high and low larval settlement rate. These observations, together with results from a model for the demography of an open, space-limited population, suggest that the settlement rate may be a more important determinant of rocky intertidal community structure than is presently realized. Locations with a low larval settlement rate exhibit a generally low abundance of barnacles that varies slightly within years and greatly between years, reflecting yearly differences in settlement. Locations with a high-settlement rate exhibit a generally high abudance of barnacles. However, the abundance varies greatly within years with a significant oscillatory component (period, 30 weeks) and only slightly between years regardless of yearly differences in settlement. At the low-settlement location mortality of barnacles is independent of the area occupied by barnacles. At the high-settlement location mortality is cover-dependent due to increased predation by starfish on areas of high barnacle cover. In both locations the cover-independent component of mortality does not vary with age during the first 60 weeks. As assumed in the demographic model, the kinetics of larval settlement can be described as a process in which the rate of settlement to a quadrat is proportional to the fraction of vacant space within the quadrat. Generalizations that the highest species diversity in a rocky intertidal community is found at locations of intermediate disturbance, and that competition causes zonation between species of the barnacle genera Balanus and Chthamalus, seem to apply only to locations with high-settlement rates.
View details for Web of Science ID A1985AKB7900039
View details for PubMedID 16593571
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Population experiments with Anolis lizard species demonstrate a relation between the amount of between-species competition and the degree of interspecific resource partitioning (the more the partitioning the less the competition). Specifically, the amount of resource partitioning between the two species (Anolis gingivinus and Anolis wattsi pogus) on the island of St. Maarten is less than that between the two species (Anolis bimaculatus and Anolis wattsi schwartzi) on the island of St. Eustatius. The presence of Anolis wattsi both lowers the growth rates and raises the perch heights of Anolis gingivinus individuals. In contrast, Anolis wattsi has no effect on Anolis bimaculatus. Thus, when there is less resource partitioning, Anolis wattsi has a greater competitive effect. This verifies, for these species, a central assumption of competition theory: the strength of between-species competition is inversely related to the amount of interspecific resource partitioning.
View details for Web of Science ID A1982NY36400025
View details for PubMedID 17782979
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In one specific model of a density-regulated population undergoing natural selection in a fluctuating environment there is a systematic evolutionary pressure favoring a lower intrinsic rate of increase, which can sometimes even overcome an evolutionary pressure favoring a higher carrying capacity.
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View details for PubMedID 16592938
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