Effective population size dynamics reveal impacts of historic climatic events and recent anthropogenic pressure in African elephants

Two hundred years of elephant hunting for ivory, peaking in 1970–1980s, caused local extirpations and massive population declines across Africa. The resulting genetic impacts on surviving populations have not been studied, despite the importance of understanding the evolutionary repercussions of such human-mediated events on this keystone species. Using Bayesian coalescent-based genetic methods to evaluate time-specific changes in effective population size, we analysed genetic variation in 20 highly polymorphic microsatellite loci from 400 elephants inhabiting the greater Samburu-Laikipia region of northern Kenya. This area experienced a decline of between 80% and 90% in the last few decades when ivory harvesting was rampant. The most significant change in effective population size, however, occurred approximately 2500 years ago during a mid–Holocene period of climatic drying in tropical Africa. Contrary to expectations, detailed analyses of four contemporary age-based cohorts showed that the peak poaching epidemic in the 1970s caused detectable temporary genetic impacts, with genetic diversity rebounding as juveniles surviving the poaching era became reproductively mature. This study demonstrates the importance of climatic history in shaping the distribution and genetic history of a keystone species and highlights the utility of coalescent-based demographic approaches in unravelling ancestral demographic events despite a lack of ancient samples. Unique insights into the genetic signature of mid-Holocene climatic change in Africa and effects of recent poaching pressure on elephants are discussed.     

Endless forms: human behavioural diversity and evolved universals

Human populations have extraordinary capabilities for generating behavioural diversity without corresponding genetic diversity or change. These capabilities and their consequences can be grouped into three categories: strategic (or cognitive), ecological and cultural-evolutionary. Strategic aspects include: (i) a propensity to employ complex conditional strategies, some certainly genetically evolved but others owing to directed invention or to cultural evolution; (ii) situations in which fitness payoffs (or utilities) are frequency-dependent, so that there is no one best strategy; and (iii) the prevalence of multiple equilibria, with history or minor variations in starting conditions (path dependence) playing a crucial role. Ecological aspects refer to the fact that social behaviour and cultural institutions evolve in diverse niches, producing various adaptive radiations and local adaptations. Although environmental change can drive behavioural change, in humans, it is common for behavioural change (especially technological innovation) to drive environmental change (i.e. niche construction). Evolutionary aspects refer to the fact that human capacities for innovation and cultural transmission lead to diversification and cumulative cultural evolution; critical here is institutional design, in which relatively small shifts in incentive structure can produce very different aggregate outcomes. In effect, institutional design can reshape strategic games, bringing us full circle.     

The genetical theory of social behaviour

We survey the population genetic basis of social evolution, using a logically consistent set of arguments to cover a wide range of biological scenarios. We start by reconsidering Hamilton''s (Hamilton 1964 J. Theoret. Biol. 7, 1–16 (doi:10.1016/0022-5193(64)90038-4)) results for selection on a social trait under the assumptions of additive gene action, weak selection and constant environment and demography. This yields a prediction for the direction of allele frequency change in terms of phenotypic costs and benefits and genealogical concepts of relatedness, which holds for any frequency of the trait in the population, and provides the foundation for further developments and extensions. We then allow for any type of gene interaction within and between individuals, strong selection and fluctuating environments and demography, which may depend on the evolving trait itself. We reach three conclusions pertaining to selection on social behaviours under broad conditions. (i) Selection can be understood by focusing on a one-generation change in mean allele frequency, a computation which underpins the utility of reproductive value weights; (ii) in large populations under the assumptions of additive gene action and weak selection, this change is of constant sign for any allele frequency and is predicted by a phenotypic selection gradient; (iii) under the assumptions of trait substitution sequences, such phenotypic selection gradients suffice to characterize long-term multi-dimensional stochastic evolution, with almost no knowledge about the genetic details underlying the coevolving traits. Having such simple results about the effect of selection regardless of population structure and type of social interactions can help to delineate the common features of distinct biological processes. Finally, we clarify some persistent divergences within social evolution theory, with respect to exactness, synergies, maximization, dynamic sufficiency and the role of genetic arguments.     

Strategies, Conflict, and the Emergence of Territoriality: The Case of the Maine Lobster Industry

In this article, we use game theory to understand the emergence of various kinds of territorial arrangements in the Maine lobster fishery during the past century. Using the Nash equilibria of models of the fishery as our theoretical framework, we show that informal territorial arrangements in this fishery went through three sequential stages. These stages are the result of decisions by groups of lobster fishermen to defend fishing areas or invade those of other groups. A large number of factors influence these defensive and offensive strategies: concentrations of lobsters, the adoption of better technology, transportation costs, ecological changes, trap monitoring costs, the ability to organize defensive and offensive groups, and better law enforcement---all of which are captured by crucial parameters of our model. We argue that this technique can be applied to elucidate territorial changes more generally.     

Long-term field study of sea kraits in New Caledonia: fundamental issues and conservation

This short review focuses on the findings associated with a long-term field study on two species of sea kraits in New Caledonia. Since 2002, more than 30 sites in the lagoon have been sampled, and in most places mark-recapture was implemented. We collected detailed data on more than 14,000 marked individuals (>6000 recaptures) and used different techniques (stable isotopes, bio-logging, analyses of diet). The objective was fundamental: to examine how amphibious snakes cope with both terrestrial and aquatic environments. As access to abundant food is likely the main evolutionary driver for the return transition toward the sea in marine tetrapods, foraging ecology was an important part of the research and novel information was obtained on this subject. Rapidly however, field observations revealed the potential interest of sea kraits for conservation issues. Our results show that these snakes are useful bio-indicators of marine biodiversity; they also provide a useful signal to monitor levels of contamination by heavy metals in the lagoon, and more generally as a means of studying the functioning of reef ecosystems. Importantly, anecdotal observations (e.g., a krait drinking during rain) provided unsuspected physiological insights of general importance to fundamental problems and conservation. One of the lessons of this long-term study is that key results emerged in an unexpected way, but all were dependent on intensive field work.     

Shoot, shovel and shut up: cryptic poaching slows restoration of a large carnivore in Europe

Poaching is a widespread and well-appreciated problem for the conservation of many threatened species. Because poaching is illegal, there is strong incentive for poachers to conceal their activities, and consequently, little data on the effects of poaching on population dynamics are available. Quantifying poaching mortality should be a required knowledge when developing conservation plans for endangered species but is hampered by methodological challenges. We show that rigorous estimates of the effects of poaching relative to other sources of mortality can be obtained with a hierarchical state-space model combined with multiple sources of data. Using the Scandinavian wolf (Canis lupus) population as an illustrative example, we show that poaching accounted for approximately half of total mortality and more than two-thirds of total poaching remained undetected by conventional methods, a source of mortality we term as 'cryptic poaching'. Our simulations suggest that without poaching during the past decade, the population would have been almost four times as large in 2009. Such a severe impact of poaching on population recovery may be widespread among large carnivores. We believe that conservation strategies for large carnivores considering only observed data may not be adequate and should be revised by including and quantifying cryptic poaching.     

Baleen whales: conservation issues and the status of the most endangered populations

Most species of baleen whales were subject to intensive overexploitation by commercial whaling in this and previous centuries, and many populations were reduced to small fractions of their original sizes. Here, we review the status of baleen whale stocks, with an emphasis on those that are known or thought to be critically endangered. Current data suggest that, of the various threats potentially affecting baleen whales, only entanglement in fishing gear and ship strikes may be significant at the population level, and then only in those populations which are already at critically low abundance. The impact of some problems (vessel harassment, and commercial or aboriginal whaling) is at present probably minor. For others (contaminants, habitat degradation, disease), existing data either indicate no immediate cause for concern, or are insufficient to permit an assessment. While the prospect for many baleen whales appears good, there are notable exceptions; populations that are of greatest concern are those suffering from low abundance and associated problems, including (in some cases) anthropogenic mortality. These include: all Northern Right Whales Eubalaena glacialis, Bowhead Whales Balaena mysticetus of the Okhotsk Sea and various eastern Arctic populations, western Gray Whales Eschrichtius robustus, and probably many Blue Whale Balaenoptera musculus populations. We review the status of these populations and, where known, the issues potentially affecting their recovery. Although Humpback Whales Megaptera novaeangliae and Southern Right Whales Eubalaena australis were also heavily exploited by whaling, existing data indicate strong recovery in most studied populations of these species.     

The problem of precision and trend detection posed by small elephant populations in West Africa

The precision of elephant estimates from aerial sample surveys and dung counts is inversely proportional to abundance. West African elephant populations are already small, and the power of a monitoring programme to detect changes in abundance diminishes as the population shrinks in size. Thus it will be difficult to evaluate the effects on elephant numbers of new management policies in West Africa. The same will be true of monitoring schemes for antelope and primate populations that are hunted for bushmeat. Elephant estimates from dung counts are more precise than those from aerial sample surveys, and changes in elephant numbers are more likely to be detected in the subregion by dung counts than by aerial sample surveys.     

The interaction energy of charged filaments in an electrolyte: Results for all filament spacings

Recent studies have explored interactions between evolutionary game dynamics and population structure. Yet most studies so far mainly paid attention to unweighted and static networks. Here we explore evolutionary games played on dynamically weighted networks. Players update their strategies according to the payoffs they obtain. Players also update weights of their adjacent links depending on payoffs they gain through those links; profitable links are reinforced whereas unprofitable ones are weakened. The system is characterized by two time scales, the one for strategy update, β_S, and the other for weight adjustment, β_W. We find that, under a mean-field approximation, the asymptotic behavior of the system is described by the replicator equation with an effective payoff matrix, which is a combination of the original game matrix A and its transpose, A^T. Both analytical and numerical results show that such an adaptive weight adjustment mechanism dramatically promotes evolution of cooperation.     

The fishing conservation and management act of 1976: Economic issues associated with foreign fishing fees and foreign allocations

Abstract

In April 1976 President Ford signed Public Law 94–265, the Fishery Conservation and Management Act of 1976. As this legislation comes of age, a far‐reaching and new approach to the management of our nation's fishery resources will be set into motion. Some of the most significant provisions of the Act are the restriction on the amount of foreign fishing, the charging of fees for foreign fishing privileges, and an enormous expansion of the area over which the United States exercises fisheries management control. This article briefly explains why the Act came into existence and what its key provisions are. It outlines the proposed foreign fee system for FY1977 and discusses the important economic impacts upon the United States from foreign fishing fees and restricted amounts of foreign fishing. It then examines possible policy questions and the important economic information necessary to assess them.     

The evolution of payoff matrices: providing incentives to cooperate

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.     

The influence of large mammalian herbivores on growth form and utilization of mopane trees, Colophospermum mopane, in Botswana's Northern Tuli Game Reserve

Mopane veld is of great value to the general ungulate spectrum in times of drought, and it is capable of retaining its dominance in a community even in the presence of extremely heavy browsing pressure imposed by large browsing mammals. Scrub mopane (hedges) has been regarded as resulting from excessive browsing pressure by large mammals, especially elephants. Both the nutrient and chemical composition of mopane twig bark were investigated, the seasonal results being related back to the seasonal utilization of branches by large mammals. Mopane twigs were most palatable in winter. Eland feed on mopane throughout the year irrespective of palatability. Elephants were rarely present in the scrub mopane area before the onset of spring rains, when the major mopane leaf flush occurs independently of rainfall. The impact of both species was not excessive and recruitment of mopane seedlings does occur. Herbivore browsing is responsible for a mopane morph which buds early and continues to produce accessible, nutritious leaves even when heavily browsed. Many browsing ungulates are reliant on this resource during the stressful transition from spring to summer in south-eastern Botswana.     

Subsistence hunting and conservation issues in the game reserve of Gile, Mozambique

Ethnozoological research was conducted to gather information on the hunting activities and their relevance for the subsistence of local people in 8 villages around the game reserve of Gile, Mozambique. Two series of data were gathered by questionnaires to: (a) 510 householders from eight villages located in the outskirts of the Reserve; (b) 10 hunters from the village of Gile, the main centre of the study area. Several hunting techniques were recorded: spears, nets, traps (including gin-traps) and wildfires, while the use of guns did not appear relevant. The importance of subsistence hunting for local people was underlined by the high percentage of respondents who declared that they usually conduct this activity and sell bushmeat. The proportion of hunters per village was related to the village size but not to its geographical location of villages and the household composition. A positive relationship existed between the proportion of hunters, crop production and fishing activities, indicating that hunting is part of an integrated system of subsistence activities. Most animals harvested were mammals (89.5%, of which 46.7% were ungulates) and most were captured within the Reserve (96%). A higher percentage of animals was sold (56%), representing a relevant income source for the villagers. Small animals were mainly captured by traps during solitary hunting, medium-sized animals in collective net hunting; larger prey were captured by gin-traps adopted by both solitary and collective hunting. In the diet of the local people wild animals represented a higher protein source than domestic animals.     

Transitions and transversions in evolutionary descent: An approach to understanding

Summary In this paper I lay a quantitative theoretical groundwork for understanding the proportions of the possible types of base substitutions observed between 12 genes sharing a common ancestor and isolated from extant species. The experimentally observed types of base substitution between two sequenced genes do not give a direct measure of the types of base substitutions that occur during evolutionary descent. However, by use of a statistical assemblage of these observations, we can recover, without the assumption of parsimony, the conditional base substitution probabilities that determine this descent. Three methods—direct count, regression, and informational entropy maximization—are described by which these probabilities can be estimated from experimental data. The methods are complementary in that each is most useful for somewhat different types of experimental data. These methods are used to study the ratio of transversions to transitions during gene divergence. Though this ratio is not constant during divergence, it does approach a stable limiting value that in principle can vary from zero, corresponding to 100% transition differences, to infinity, corresponding to 0% transition differences. In practice the limiting ratio tends to hover around a value of two, which is expected on a random basis. However, base substitution pathways that are very nonrandom also may lead to a limiting ratio of exactly two, so that such a value is not diagnostic for random pathways. The limiting ratio can be directly calculated from a knowledge of the twelve conditional probabilities for each type of base substitution, or from a knowledge of the equilibrium base composition of the DNAs compared. An expression is given for this calculation. Fifteen years ago Jean Derancourt, Andrew Lebor and Emile Zuckerkandl (1967), analyzing the amino acid sequence of globin chains coded by nuclear genes, made the original observation that the proportion of transition differences decreases with increasing evolutionary time. Recently Brown et al. (1982) and Brown and Simpson (1982) have reported a decrease in the observed proportion of transition differences in mitochondrial DNA with increasing evolutionary divergence. The conditions that must be satisfied for this type of behavior to occur at stable base composition and with stable base substitution probabilities are defined. Multiple substitutionsper se do not lead to a decrease in transition differences with increasing evolutionary divergence.     

Creating Grasslands: Social Institutions and Environmental Change in Mkambati Area, South Africa

Through a case study of the grassland system of the Mkambati area in the Eastern Cape, South Africa, the paper explores the interaction of social institutions and ecological processes in the creation of a diverse grassland environment. A number of different transitions between grassland states are examined in detail, highlighting a range of social and institutional factors influencing grassland change. The spatial patterning of different grassland types and the frequency of transitions between them are shown to be dependent on the institutional relationships between different social actors. Understanding such complex and multifaceted processes of environmental change requires analytical tools which combine social and ecological perspectives;an extended form of qualitative state-transition modeling, which incorporates institutional dimensions, is therefore explored.     

Randomized controlled experiments in health and social sciences: Some conceptual issues

This brief article outlines some difficulties as well as benefits in conducting randomized controlled trials in social science settings especially in developing countries. Some of the historical developments are summarized and certain applications in health sciences are discussed from methodological and policy standpoints.