Game theory, multi-modal signalling and the evolution of communication

We discuss how the theoretical framework related to selection pressures on multi-component and multi-modal signalling introduced by the article of Wilson et al. in this special issue could usefully be built both theoretically and empirically. Theoretically, we suggest that the game theory approach of Wilson et al. could be generalised using the methodology of adaptive dynamics in order to indicate evolutionary trajectories and paths of evolution. This might indicate the relative likelihood of finding different equilibriums empirically in situations where the game theory explorations of Wilson et al. suggest that multiple equilibriums may exist for a given system. We also suggest how the work of Wilson et al. could be extended theoretically and empirically to further capture the effects of receiver psychology on selective pressures on multi-component and multi-modal signals. We also highlight the assumptions and predictions of the existing theory that would most benefit from and would be most amenable to empirical testing.     

Persistence models for mark-recapture

The stable of models available for analyzing mark-recapture data (Otis et al. Wild Momogr 66:135, 1978) includes those having behavioral characteristics, time variation, heterogeneity, along with combinations of those characteristics. This paper proposes use of a series of models based on the persistence model of Ramsey and Usner (Biometrics 59:331–339, 2003). We show that persistence can be modeled in combination with behavior and with time variation. We apply the persistence model to situations in which capture occasions are not equally-spaced in time. Two case studies illustrate the use of these extended persistence models.     

Effects of learning and adaptation on population viability

Cultural adaptation is one means by which conservationists may help populations adapt to threats. A learned behavior may protect an individual from a threat, and the behavior can be transmitted horizontally (within generations) and vertically (between generations), rapidly conferring population-level protection. Although possible in theory, it remains unclear whether such manipulations work in a conservation setting; what conditions are required for them to work; and how they might affect the evolutionary process. We examined models in which a population can adapt through both genetic and cultural mechanisms. Our work was motivated by the invasion of highly toxic cane toads (Rhinella marina) across northern Australia and the resultant declines of endangered northern quolls (Dasyurus hallucatus), which attack and are fatally poisoned by the toxic toads. We examined whether a novel management strategy in which wild quolls are trained to avoid toads can reduce extinction probability. We used a simulation model tailored to quoll life history. Within simulations, individuals were trained and a continuous evolving trait determined innate tendency to attack toads. We applied this model in a population viability setting. The strategy reduced extinction probability only when heritability of innate aversion was low (<20%) and when trained mothers trained >70% of their young to avoid toads. When these conditions were met, genetic adaptation was slower, but rapid cultural adaptation kept the population extant while genetic adaptation was completed. To gain insight into the evolutionary dynamics (in which we saw a transitory peak in cultural adaptation over time), we also developed a simple analytical model of evolutionary dynamics. This model showed that the strength of natural selection declined as the cultural transmission rate increased and that adaptation proceeded only when the rate of cultural transmission was below a critical value determined by the relative levels of protection conferred by genetic versus cultural mechanisms. Together, our models showed that cultural adaptation can play a powerful role in preventing extinction, but that rates of cultural transmission need to be high for this to occur.     

Behavioral transitions with the evolution of cooperative nest founding by harvester ant queens

Research on the evolution of cooperative groups tends to explore the costs and benefits of cooperation, with less focus on the proximate behavioral changes necessary for the transition from solitary to cooperative living. However, understanding what proximate changes must occur, as well as those pre-conditions already in place, is critical to understanding the origins and evolution of sociality. The California harvester ant Pogonomyrmex californicus demonstrates population-level variation in colony founding over a close geographic range. In adjacent populations, queens either found nests as single individuals (haplometrosis) or form cooperative groups of nonrelatives (pleometrosis). We compared aggregation, aggression, and tolerance of queens from one pleometrotic and two haplometrotic populations during nest initiation, to determine which behaviors show an evolutionary shift and which are present at the transition to pleometrosis. Surprisingly, within-nest aggregative behavior was equally present among all populations. In nesting boxes with multiple available brood-rearing sites, both queen types readily formed and clustered around a single common brood pile, suggesting that innate attraction to brood (offspring) facilitates the transition to social aggregation. In contrast, queens from the three populations differed in their probabilities of attraction on the ground to nest sites occupied by other queens and in levels of aggression. Our results suggest that some key behavioral mechanisms facilitating cooperation in P. californicus are in place prior to the evolution of pleometrosis and that the switch from aggression to tolerance is critical for the evolution of stable cooperative associations.     

Colony level and within colony level selection in honeybees

Summary Although honeybee workers are usually infertile, in queenless colonies some workers can develop ovaries and produce offspring. Therefore the classical Darwinian fitness of workers is not zero. Experimental studies in the Cape honey bee (Apis mellifera capensis) reveal a huge genetic variation for individual fitness of workers. The present study with a one locus, two allele model for reproductive dominance of workers shows that a balanced system between colony level and individual within colony selection plausibly explains the phenomenon of a high genetic variance of worker fitness. In particular, a frequent occurrence of queenless colonies in the population leads to stable polymorphic equilibria. Also the multiple mating system of the honey bee queen supports the propagation of alleles causing reproductive dominance of workers.     

Indications of fitness differences among vocal clans of sperm whales

Cultural variation can affect the genetic evolution of a species if there are consistent cultural differences that contribute to fitness variation between groups of individuals. In this study, measures of the reproductive success of groups of sperm whales from different cultural clans are used as proxies for fitness. We measure reproductive success using population length distributions from acoustic and photographic measurements and visual observations of the presence of calves. The results obtained are generally consistent between methods; there are large and significant differences between the clans in the measures of reproductive success. The results from this study strengthen the case for cultural hitchhiking in sperm whales by indicating that differences in culture between clans correlate with differences in fitness.     

Local‐Scale Dynamics and Local Drivers of Bushmeat Trade

Bushmeat management policies are often developed outside the communities in which they are to be implemented. These policies are also routinely designed to be applied uniformly across communities with little regard for variation in social or ecological conditions. We used fuzzy‐logic cognitive mapping, a form of participatory modeling, to compare the assumptions driving externally generated bushmeat management policies with perceptions of bushmeat trade dynamics collected from local community members who admitted to being recently engaged in bushmeat trading (e.g., hunters, sellers, consumers). Data were collected during 9 workshops in 4 Tanzanian villages bordering Serengeti National Park. Specifically, we evaluated 9 community‐generated models for the presence of the central factors that comprise and drive the bushmeat trade and whether or not models included the same core concepts, relationships, and logical chains of reasoning on which bushmeat conservation policies are commonly based. Across local communities, there was agreement about the most central factors important to understanding the bushmeat trade (e.g., animal recruitment, low income, and scarcity of food crops). These matched policy assumptions. However, the factors perceived to drive social‐ecological bushmeat trade dynamics were more diverse and varied considerably across communities (e.g., presence or absence of collaborative law enforcement, increasing human population, market demand, cultural preference). Sensitive conservation issues, such as the bushmeat trade, that require cooperation between communities and outside conservation organizations can benefit from participatory modeling approaches that make local‐scale dynamics and conservation policy assumptions explicit. Further, communities’ and conservation organizations’ perceptions need to be aligned. This can improve success by allowing context appropriate policies to be developed, monitored, and appropriately adapted as new evidence is generated. Dinámicas a Escala Local y Conductores Locales del Mercado de Carne de Caza     

Sexual selection and the evolutionary effects of copying mate choice

We examine the evolutionary consequences of copying mate choice using models in which the preferences of younger females are affected by the mate choices that they observe older females making. We introduce two models of copying, termed single mate copying and mass copying, corresponding to situations in which immature females imprint on the choices of only one or of a very large number of older females, respectively. Female mating preferences are assumed to evolve only through cultural evolution, while the male trait on which they act is inherited either via a haploid autosomal or a Y-linked locus. Results show that the preference and male trait can rapidly coevolve, with a positive frequencey-dependent advantage to the more common male trait allele. This process can cause a display trait that lowers male viability to increase in a population. Mass copying results in stronger frequency dependence than does single mate copying. Mass copying and, under some conditions, single mate copying lead to two alterative stable equilibria for the male trait. Neither copying model supports variation at the male trait locus, and copying makes it more difficult for a novel male trait phenotype to spread. Correspondence to: M. Kirkpatrick     

Estimating dispersal potential for marine larvae: dynamic models applied to scleractinian corals

Dispersal influences ecological dynamics, evolution, biogeography, and biodiversity conservation, but models of larval dispersal in marine organisms make simplifying assumptions that are likely to approximate poorly the temporal dynamics of larval survival and capacity for settlement. In particular, larval mortality rates are typically assumed to be constant throughout larval life; and all larvae are frequently assumed to acquire and lose competence at the same time. To improve upon these assumptions, we here develop simple models of dispersal potential that incorporate rates of mortality, and acquisition and loss of settlement competence. We fit these models to empirical competence and survival data for five scleractinian coral species, to test the models' ability to characterize empirical survival and competence patterns, and to estimate the dispersal potential implied by those patterns. The models fit the data well, incorporating qualitative features of competence and survival that traditional approaches to modeling dispersal do not, with important implications for dispersal potential. Most notably, there was high within-cohort variation in the duration of the competent period in all species, and this variation increases both self-recruitment and long-distance dispersal compared with models assuming a fixed competent period. These findings help to explain the seeming paradox of high genetic population structure, coupled with large geographic range size, observed in many coral species. More broadly, our approach offers a way to parsimoniously account for variation in competence dynamics in dispersal models, a phenomenon that our results suggest has important effects on patterns of connectivity in marine metapopulations.     

Grass feedbacks on fire stabilize savannas

Savannas commonly consist of a discontinuous cover of overstory trees and a groundcover of grasses. Savanna models have previously demonstrated that vegetation feedbacks on fire frequency can limit the density of overstory trees, thereby maintaining savannas. Positive feedbacks of either savanna trees alone or trees and grasses together on fire frequency have been shown to result in a stable savanna equilibrium. Grass feedbacks on fire frequency, in contrast, have resulted in stable equilibria in either a grassland or forest state, but not in a savanna. These results, however, were derived from a system of differential equations that assumes that fire occurrence is strictly deterministic and that vegetation losses due to fire are continuous in time. We develop an alternative formulation of the grass-fire feedback model that assumes that fires are discrete and occur stochastically in time to examine the influence of these assumptions on the predicted state of the system. We show that incorporating fire as a discrete event can produce a recurring temporal refuge in which both grass and trees co-occur in a stable, bounded savanna. In our model, tree abundance is limited without invoking demographic bottlenecks in the transition from fire-sensitive to fire-resistant life history stages. An increasing strength of grass feedback on fire results in regular, predictable fires, which suggests that the system can also be modeled using a set of difference equations. We implement this discrete system using modified Leslie/Gower difference equations and demonstrate the existence of a bounded savanna state in this model framework. Our results confirm the potential for grass feedbacks to result in stable savannas, and indicate the importance of modeling fire as a discrete event rather than as a loss rate that is continuous in time.     

Beiträge zur Temperaturadaptation des Aales Anguilla anguilla IV. Einfluß der Adaptationstemperatur auf den denervierten Seitenmuskel

In order to discover whether the spinal cord plays a role in muscular temperature adaptation, the responses of spinalectomized and intact eels Anguilla anguilla L. were compared. Eight and 14 days after denervation, the activities of the succinate oxidase-system and of cytochromoxidase were examined in 4 groups of eels (adapted to 14° or 24°C before and after operation; adapted to 14°C before and to 24°C after operation; or vice versa). Only the 24°C eels show significant differences which refer to the second phase of atrophy according to Muscatello et al. (1965). All differences measured at the various adaptation temperatures can be explained as adaptation effects allowing for the consequences of the operation (atrophy) on the denervated individuals. Therefore, a significant influence of the spinal cord on temperature adaptation of the lateral muscle of the eel is improbable.     

The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.     

Genetic relatedness and evolution of insect sociality

Summary Genetic relatedness is a vital concept in the study of the evolution of social behaviour because it can be used to predict gene transmission into future generations. Any models using genetic relatedness to predict the course of social evolution have to take into account that this relatedness, when defined as genotypic regression, is affected by selection and that the evolution also depends on specific assumptions concerning the cost/benefit ratios. In this paper these problems are shown to exist in recent studies examining the evolution of worker behaviour in insects; these studies are reanalysed using allele frequency models, and the results compared with those from other models.It is shown that cyclic inbreeding (alternating outbred and inbred generations) can favour worker evolution in diploid populations but is a necessary condition only within a very limited range of the cost/benefit ratio.Male-haploid models show that both female and male workers evolve more easily if they can manipulate the brood produced by the queen by biassing the brood sex ratio in favour of females (the optimal proportion of females is ca. 0.75). The threshold for female workers to evolve is also lowered if they are allowed to produce some of the colony's male offspring (which arise from unfertilized eggs), in which case the optimal brood sex ratio becomes closer to 1:1. It is apparent that the two factors — sex-ratio bias and worker-produced males — are also important in the comparison of worker evolution in male-haploid and diploid insects.     

Aggregation of Dynamic Systems and the Existence of a Regeneration Function

In environmental economics, complex ecosystems are often represented by low-dimensional models. The question of whether the results of these models can be applied to a more complex reality leads to the investigation of an aggregation problem in a nonlinear dynamic setting. We show that restrictive assumptions are needed for aggregation and that the low-dimensional model has to be linear. On the basis of these results, we argue that the aggregation of complex ecosystems is often oversimplifying and that substantial gains can be expected from the use of more complex models.     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

Agricultural heritage in disintegration: Trends of agropastoral transhumance on the southeast Tibetan Plateau

This paper examines agropastoral transhumance in 12 sampled villages on the southeast Tibetan Plateau. By analysing trends in this indigenous livelihood system and examining causes and potential consequences, we conclude that Tibetan agropastoral transhumance is a fragile system extremely vulnerable to external impacts. Within the context of China's rapid development and globalisation, this agricultural heritage is changing rapidly, as is most saliently reflected in herd size and structure, movement patterns, relative economic importance of pastoral activities and the relations between pastoralism and crop cultivation. Immediate causes of changes are local changes in livestock management objectives, land uses and labour availability, which are driven by more fundamental changes of environmental, political, socioeconomic, technological and cultural profiles, from local to global scales. Changes in local livelihood systems can have multifaceted and profound political, socioeconomic, cultural and ecological consequences, in both constructive and destructive terms. Policy-makers and/or project designers must have a holistic perspective so as to integrate multiple objectives of promoting sustainable socioeconomic development, preserving biological and cultural diversities and maintaining the environmentally stable balance of human society, animal population and ecosystem that has existed in the area for centuries.     

Ornaments and condition: plumage patch sizes,nutritional reserve state,reserve accumulation,and reserve depletion

Behavioral Ecology and Sociobiology - Cooperation plays a crucial role in primate social life. However, the evolution of large-scale human cooperation from the cognitive fundamentals found in other...     

Temporal, spatial, and body size effects on growth rates of loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

In response to a call from the US National Research Council for research programs to combine their data to improve sea turtle population assessments, we analyzed somatic growth data for Northwest Atlantic (NWA) loggerhead sea turtles (Caretta caretta) from 10 research programs. We assessed growth dynamics over wide ranges of geography (9–33°N latitude), time (1978–2012), and body size (35.4–103.3 cm carapace length). Generalized additive models revealed significant spatial and temporal variation in growth rates and a significant decline in growth rates with increasing body size. Growth was more rapid in waters south of the USA (<24°N) than in USA waters. Growth dynamics in southern waters in the NWA need more study because sample size was small. Within USA waters, the significant spatial effect in growth rates of immature loggerheads did not exhibit a consistent latitudinal trend. Growth rates declined significantly from 1997 through 2007 and then leveled off or increased. During this same interval, annual nest counts in Florida declined by 43 % (Witherington et al. in Ecol Appl 19:30–54, 2009) before rebounding. Whether these simultaneous declines reflect responses in productivity to a common environmental change should be explored to determine whether somatic growth rates can help interpret population trends based on annual counts of nests or nesting females. Because of the significant spatial and temporal variation in growth rates, population models of NWA loggerheads should avoid employing growth data from restricted spatial or temporal coverage to calculate demographic metrics such as age at sexual maturity.     

Nonequilibrium coexistence in a competition model with nutrient storage

Resource competition theory predicts that, in equilibrium, the number of coexisting species cannot exceed the number of limiting resources. In some competition models, however, competitive interactions may result in nonequilibrium dynamics, allowing the coexistence of many species on few resources. The relevance of these findings is still unclear, since some assumptions of the underlying models are unrealistic. Most importantly, these models assume that individual growth directly reflects the availability of external resources, whereas real organisms can store resources, thereby decoupling their growth from external fluctuations. Here we study the effects of resource storage by extending the well-known Droop model to the context of multiple species and multiple resources. We demonstrate that the extended Droop model shows virtually the same complex dynamics as models without storage. Depending on the model parameters, one may obtain competitive exclusion, stable equilibrium coexistence, periodic and non-periodic oscillations, and chaos. Again, nonequilibrium dynamics allows for the coexistence of many species on few resources. We discuss our findings in the light of earlier work on resource competition, highlighting the role of luxury consumption, trade-offs in competitive abilities, and ecological stoichiometry.     

Using the negative binomial distribution to model overdispersion in ecological count data

A Poisson process is a commonly used starting point for modeling stochastic variation of ecological count data around a theoretical expectation. However, data typically show more variation than implied by the Poisson distribution. Such overdispersion is often accounted for by using models with different assumptions about how the variance changes with the expectation. The choice of these assumptions can naturally have apparent consequences for statistical inference. We propose a parameterization of the negative binomial distribution, where two overdispersion parameters are introduced to allow for various quadratic mean-variance relationships, including the ones assumed in the most commonly used approaches. Using bird migration as an example, we present hypothetical scenarios on how overdispersion can arise due to sampling, flocking behavior or aggregation, environmental variability, or combinations of these factors. For all considered scenarios, mean-variance relationships can be appropriately described by the negative binomial distribution with two overdispersion parameters. To illustrate, we apply the model to empirical migration data with a high level of overdispersion, gaining clearly different model fits with different assumptions about mean-variance relationships. The proposed framework can be a useful approximation for modeling marginal distributions of independent count data in likelihood-based analyses.