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.     

Understanding the nature of wealth and its effects on human fitness

Studying fitness consequences of variable behavioural, physiological and cognitive traits in contemporary populations constitutes the specific contribution of human behavioural ecology to the study of human diversity. Yet, despite 30 years of evolutionary anthropological interest in the determinants of fitness, there exist few principled investigations of the diverse sources of wealth that might reveal selective forces during recent human history. To develop a more holistic understanding of how selection shapes human phenotypic traits, be these transmitted by genetic or cultural means, we expand the conventional focus on associations between socioeconomic status and fitness to three distinct types of wealth-embodied, material and relational. Using a model selection approach to the study of women's success in raising offspring in an African horticultural population (the Tanzanian Pimbwe), we find that the top performing models consistently include relational and material wealth, with embodied wealth as a less reliable predictor. Specifically, child mortality risk is increased with few household assets, parent nonresidency, child legitimacy, and one or more parents having been accused of witchcraft. The use of multiple models to test various hypotheses greatly facilitates systematic comparative analyses of human behavioural diversity in wealth accrual and investment across different kinds of societies.     

The evolution of superstitious and superstition-like behaviour

Superstitious behaviours, which arise through the incorrect assignment of cause and effect, receive considerable attention in psychology and popular culture. Perhaps owing to their seeming irrationality, however, they receive little attention in evolutionary biology. Here we develop a simple model to define the condition under which natural selection will favour assigning causality between two events. This leads to an intuitive inequality--akin to an amalgam of Hamilton's rule and Pascal's wager--that shows that natural selection can favour strategies that lead to frequent errors in assessment as long as the occasional correct response carries a large fitness benefit. It follows that incorrect responses are the most common when the probability that two events are really associated is low to moderate: very strong associations are rarely incorrect, while natural selection will rarely favour making very weak associations. Extending the model to include multiple events identifies conditions under which natural selection can favour associating events that are never causally related. Specifically, limitations on assigning causal probabilities to pairs of events can favour strategies that lump non-causal associations with causal ones. We conclude that behaviours which are, or appear, superstitious are an inevitable feature of adaptive behaviour in all organisms, including ourselves.     

Habitat stability and predation pressure affect temperament behaviours in populations of three-spined sticklebacks

1. There is growing interest in the causes and consequences of animal temperaments. Temperament behaviours often have heritable components, but ecological variables can also affect them. Numerous variables are likely to differ between habitats, and these may interact to influence temperament behaviours. 2. Temperament behaviours may be correlated within populations (behavioural syndromes), although the underlying causes of such correlations are currently unclear. 3. We analysed three different temperament behaviours and learning ability in three-spined sticklebacks, Gasterosteus aculeatus, to determine how different ecological variables influence them both within and between populations. We selected populations from four ponds and four rivers that varied naturally in their exposure to predators. 4. High-predation river populations were significantly less bold than a high-predation pond and low-predation river populations, and low-predation pond populations were significantly less bold than a high-predation pond population. Within populations, temperament behaviours were correlated in one high-predation river population only. 5. These results suggest that multiple ecological factors can interact to affect temperament behaviours between populations, and also correlations in those behaviours within populations.     

Tractable models for testing theories about natural strategies: foraging behaviour and habitat selection of free‐ranging sharks

Marine and terrestrial environments differ fundamentally in space‐time scales of both physical and ecological processes. These differences will have an impact on the animals inhabiting each domain, particularly with respect to their spatial ecology. The behavioural strategies that underpin observed distributions of marine species are therefore important to consider. Comparatively little is known, however, about how wild fishes actually respond to gradients in food supply and temperature, and to potential mates. This paper describes how behavioural theory is being used to elucidate the strategies and tactics of free‐ranging sharks in three specific areas of study, namely, foraging on zooplankton, behavioural energetics and sexual segregation. The studies discussed are novel because shark movements were tracked in the wild using electronic tags in relation to simultaneous measurements of prey densities and thermal resources. The results show that filter‐feeding (basking shark, Cetorhinus maximus ) and predatory (dogfish, Scyliorhinus canicula ) sharks have relatively complex behaviour patterns integrally linked to maximizing surplus power, often through making short and longer term 'trade‐off' decisions between optimal foraging and thermal habitats. Interestingly, female S. canicula exhibit alternative behavioural strategies compared to males, a difference resulting in spatial segregation by habitat. Sexual segregation in this species occurs primarily as a consequence of male avoidance by females. Studies on free‐ranging sharks provide a useful model system for examining how a predator's strategy is shaped by its environment. More theory‐based studies of the behavioural processes of sharks are required however, before critical comparisons with other vertebrate predators are possible. Suggestions for further research to address this knowledge gap are given.     

Cooperation and conflict between women in the family

Here I review recent research on reproductive conflict between females in families and how it influences their reproductive behaviour. Kin selection can favor cooperation between parent and offspring, siblings, or unrelated co‐residents who share interests in other family members such as grand‐offspring. However, these are also the individuals most likely to be sharing resources, and so conflict can also emerge. While substantial interest has arisen in evolutionary anthropology, especially over the last two decades, in the possibility of cooperative breeding in humans, less attention has been paid to reproductive conflict among female kin. Communal breeding in animals is generally understood as emerging from competition over the resources needed to breed. Competition for household resources is a problem that also faces human families. Models suggest that in some circumstances, inclusive fitness can be maximized by sharing reproduction rather than harming relatives by fighting with them, even if the shares that emerge are not equal. Thus, competition and cooperation turn out to be strongly related to each other. Reproductive competition within and between families may have underpinned the biological evolution of fertility patterns (such as menopause) and the cultural evolution of marriage, residence, and inheritance norms (such as late male marriage or primogeniture), which can enhance cooperation and minimize the observed incidence of such conflicts.     

Thermoregulatory behaviour in Tropidurus torquatus (Squamata, Tropiduridae) from Brazilian coastal populations: an estimate of passive and active thermoregulation in lizards

Geographical variation in the degree of thermoregulatory behaviour of the lizard Tropidurus torquatus was studied in 10 restinga populations along approximately 1500 km of Brazilian coast. An index of thermoregulation was estimated using the difference between body and environmental (air and substrate) temperatures and the percentage of negative values for these differences (proportion of body temperatures lower than environmental temperatures). In most populations, the lizards primarily used active thermoregulation, through behavioural means, and mainly in relation to substrate temperatures. Along the restingas, the degree of active thermoregulation increased as a function of the increase in the local environmental temperatures along the restingas. Behavioural thermoregulation of T. torquatus helps the lizards to maintain their body temperatures within an optimal range in which to perform their normal daily activities.     

Habitat use, home range and foraging preferences of the coati Nasua nasua in a pluvial tropical Atlantic forest area

Coatis (genus Nasua ) occupy a large range of forested habitats. The aim of this study was to describe the use of habitat by Nasua nasua in a continuous pluvial tropical Atlantic forest area, the Carlos Botelho State Park (CBSP) in south-eastern Brazil. Use of space by N. nasua in the CBSP was characterized by an average home range area for one group of 445 ha by the minimum convex polygon method and 544 ha by the fixed kernel method, which was used across 2 months; the use of trees for foraging and travelling was superior to that reported for N. nasua and Nasua narica until now, being higher in the drier season. The home range of three coati groups shifted during the 3 years of this study; the home range of one habituated group changed by 91%. The proportion of ground foraging increased with the higher availability of soil invertebrates during the rainy season, but cannot be explained solely on the basis of resource availability.     

Intra-deme molecular diversity in spatially expanding populations

We report here a simulation study examining the effect of a recent spatial expansion on the pattern of molecular diversity within a deme. We first simulate a range expansion in a virtual world consisting in a two-dimensional array of demes exchanging a given proportion of migrants (m) with their neighbors. The recorded demographic and migration histories are then used under a coalescent approach to generate the genetic diversity in a sample of genes. We find that the shape of the gene genealogies and the overall pattern of diversity within demes depend not only on the age of the expansion but also on the level of gene flow between neighboring demes, as measured by the product Nm, where N is the size of a deme. For small Nm values (< approximately 20 migrants sent outwards per generation), a substantial proportion of coalescent events occur early in the genealogy, whereas with larger levels of gene flow, most coalescent events occur around the time of the onset of the spatial expansion. Gene genealogies are star shaped, and mismatch distributions are unimodal after a range expansion for large Nm values. In contrast, gene genealogies present a mixture of both very short and very long branch lengths, and mismatch distributions are multimodal for small Nm values. It follows that statistics used in tests of selective neutrality like Tajima's D statistic or Fu's F(S) statistic will show very significant negative values after a spatial expansion only in demes with high Nm values. In the context of human evolution, this difference could explain very simply the fact that analyses of samples of mitochondrial DNA sequences reveal multimodal mismatch distributions in hunter-gatherers and unimodal distributions in post-Neolithic populations. Indeed, the current simulations show that a recent increase in deme size (resulting in a larger Nm value) is sufficient to prevent recent coalescent events and thus lead to unimodal mismatch distributions, even if deme sizes (and therefore Nm values) were previously much smaller. The fact that molecular diversity within deme is so dependent on recent levels of gene flow suggests that it should be possible to estimate Nm values from samples drawn from a single deme.     

Flexibility in reproductive timing in human females: integrating ultimate and proximate explanations

From an ultimate perspective, the age of onset of female reproduction should be sensitive to variation in mortality rates, and variation in the productivity of non-reproductive activities. In accordance with this prediction, most of the cross-national variation in women's age at first birth can be explained by differences in female life expectancies and incomes. The within-country variation in England shows a similar pattern: women have children younger in neighbourhoods where the expectation of healthy life is shorter and incomes are lower. I consider the proximate mechanisms likely to be involved in producing locally appropriate reproductive decisions. There is evidence suggesting that developmental induction, social learning and contextual evocation may all play a role.     

Risk, resources and state-dependent adaptive behavioural syndromes

Many animals exhibit behavioural syndromes-consistent individual differences in behaviour across two or more contexts or situations. Here, we present adaptive, state-dependent mathematical models for analysing issues about behavioural syndromes. We find that asset protection (where individuals with more 'assets' tend be more cautious) and starvation avoidance, two state-dependent mechanisms, can explain short-term behavioural consistency, but not long-term stable behavioural types (BTs). These negative-feedback mechanisms tend to produce convergence in state and behaviour over time. In contrast, a positive-feedback mechanism, state-dependent safety (where individuals with higher energy reserves, size, condition or vigour are better at coping with predators), can explain stable differences in personality over the long term. The relative importance of negative- and positive-feedback mechanisms in governing behavioural consistency depends on environmental conditions (predation risk and resource availability). Behavioural syndromes emerge more readily in conditions of intermediate ecological favourability (e.g. medium risk and medium resources, or high risk and resources, or low risk and resources). Under these conditions, individuals with higher initial state maintain a tendency to be bolder than individuals that start with low initial state; i.e. later BT is determined by state during an early 'developmental window'. In contrast, when conditions are highly favourable (low risk, high resources) or highly unfavourable (high risk, low resources), individuals converge to be all relatively bold or all relatively cautious, respectively. In those circumstances, initial differences in BT are not maintained over the long term, and there is no early developmental window where initial state governs later BT. The exact range of ecological conditions favouring behavioural syndromes depends also on the strength of state-dependent safety.     

Variation in genetic architecture of olfactory behaviour among wild-derived populations of Drosophila melanogaster

Odour-guided behaviour is a quantitative trait determined by many genes that are sensitive to gene-environment interactions. Different natural populations are likely to experience different selection pressures on the genetic underpinnings of chemosensory behaviour. However, few studies have reported comparisons of the quantitative genetic basis of olfactory behaviour in geographically distinct populations. We generated isofemale lines of Drosophila melanogaster from six populations in Argentina and measured larval and adult responses to benzaldehyde. There was significant variation within populations for both larval and adult olfactory behaviour and a significant genotype x sex interaction (GSI) for adult olfactory behaviour. However, there is substantial variation in the contribution of GSI to the total phenotypic variance among populations. Estimates of evolvability are orders of magnitude higher for larvae than for adults. Our results suggest that the potential for evolutionary adaptation to the chemosensory environment is greater at the larval feeding stage than at the adult reproductive stage.     

Habitat preference of Great Argus Pheasant (Argusianus argus) in Kayan Mentarang National Park, East Kalimantan, Indonesia

Summary Habitat preference and abundance of the Great Argus Pheasant was studied over a ten week period in the lowland forests of the Kayan Mentarang National Park, in the remote interior of Indonesian Borneo. Repeat line transect surveys, counting the number of vocalising males, were conducted in four different habitat types. Argus Pheasants were found to be most common in primary forest, and progressively less common in old and young secondary forests. Numbers of calling males were lowest in riverine forest. Argus Pheasant densities were positively correlated with a number of transect characteristics, viz. increasing tree diameter, tree height, height of the first bough and canopy cover, and with increasing remoteness. Dancing grounds were invariably located on ridges. The conservation status of the species on Borneo is briefly discussed. The Great Argus Pheasant is still widespread but thinly spread. The rapid rate of lowland forest clearance, fragmentation of once continuous habitat and increasing hunting pressure owing to increasing accessibility may become serious threats in the near future.

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Habitatpräferenz des Großen Argusfasans (Argusianus argus) im Kayan Mentarang National Park, Ost Kalimantan, Indonesien

Zusammenfassung Die Habitatpräferenz und das Vorkommen des Großen Argusfasans wurden zehn Wochen lang im Flachlandurwald des Kayan Mentarang National Park, tief im Inneren von Kalimantan, Borneo, Indonesien, untersucht. Dazu wurden in vier verschiedenen Vegetationstypen die Anzahl vokalisierender Männchen entlang von Linientransekten festgestellt. Am häufigsten waren die Vögel im Primär-Urwald, seltener in altem und noch seltener in jüngerem Sekundärwald. Am geringsten war die Anzahl rufender Männchen in den flußnahen Wäldern. Die Häufigkeit des Vorkommens war positiv korreliert mit Stammdurchmesser, Stammhöhe, Höhe des ersten dicken Astes, Dichte der Krone und Entfernung zu Ansiedlungen. Die Balzpläte befanden sich immer auf Hügeln oder Hügelrücken. Der Argusfasan ist noch immer weitverbreitet aber nirgends häufig. Das rasche Abholzen von Flachlandwäldern und die zunehmende Jagd, die durch ein neues Wegenetz immer einfacher wird, sind die wichtigsten Gefährdungsfaktoren.

     

The genetic and environmental basis of adaptive differences in shoaling behaviour among populations of Trinidadian guppies,Poecilia reticulata

The degree of plasticity an individual expresses when moving into a new environment is likely to influence the probability of colonization and potential for subsequent evolution. Yet few empirical examples exist where the ancestral and derived conditions suggest a role for plasticity in adaptive genetic divergence of populations. Here we explore the genetic and plastic components of shoaling behaviour in two pairs of populations of Poecilia reticulata (Trinidadian guppies). We contrast shoaling behaviour of guppies derived from high‐ and low‐predation populations from two separate drainages by measuring the shoaling response of second generation laboratory‐reared individuals in the presence and absence of predator induced alarm pheromones. We find persistent differences in mean shoaling cohesion that suggest a genetic basis; when measured under the same conditions high‐predation guppies form more cohesive shoals than low‐predation guppies. Both high and low‐predation guppies also exhibit plasticity in the response to alarm pheromones, by forming tighter, more cohesive shoals. These patterns suggest a conserved capacity for adaptive behavioural plasticity when moving between variable predation communities that are consistent with models of genetic accommodation.     

Social,biological, and political considerations on fertility in Arab populations

In analyzing fertility in the Arab countries, crude birth rate, total fecundity rate, and age specific fertility rates were measured. The data was obtained from United Nations, UNICEF, and the World Bank. In the early 1980's 13 of the countries had birth rates 40/1000. The majority of countries showed a decline in their crude birth rate (CBR) between 1960-83, except Somalia, which increased. The United Arab Emirates (UAE), Tunisia, Lebanon, and Kuwait, had the largest CBR decreases, followed by Morocco, Egypt, and Saudi Arabia. The global fecundity rate (GFR) shows the number of expected births a woman lives through her reproductive period, having children at the prevailing rate for each age. The GFR in these countries is much higher than those of non Moslem countries in the area. Results show that the fertility of Arab countries are in a gradual decline, but remain high, and many have a CBR over 40/1000. In the last 20 years Saudi Arabia, with the largest population of oil producing countries, has had a decreasing CBR. It is not in agreement with its high GFR, but this can be attributed to the large number of immigration workers in the country. The UAE showed a decrease in CBR from 46/1000 to 27/1000, the largest decrease in these countries. This decline coincided with the economic development due to oil production. Kuwait had a 25.5% decrease in CBR but less than Tunisia and Lebanon. The fertility decline in Kuwait intensified in the middle 1970's; the decline in northern Africa began in the late 1960's. There were declines in birth rates in the North African countries in the early 1970's except for Tunisia. The rapid declines in fertility can be attributed to the countries' socioeconomic and political situations.