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.     

Changes in age-specific behaviour and spatial structure of masked greenling, Hexagrammos octogrammus Hexagrammidae)

We studied spatial distribution, social behaviour and temporal patterns of daily activity of two different size classes of masked greenling, Hexagrammos octogrammus, during the non-breeding period (June to early September 1986-1988) in the Sea of Japan because of appreciable differences in young and adult fish social behaviour and habitat preferences. Small, young fish (1 year old, SL 50–130 mm) inhabited the rocky littoral zone where aquatic predators were absent and food sources were limited. Small greenling were territorial and occupied and guarded areas ranging from 1.5 to 2.5 m2. Adult fish (> 2 years old, SL 140–240 mm) occupied the upper sublittoral zone, with high food abundance. They were non-territorial and were moving within home ranges up to several hundreds of square meters. A few young greenling were observed in sublittoral adult habitats. However, they displayed territorial behaviour. Similarly, several adults, present consistently in rocky sublittoral habitat were non-territorial. Aggressive bouts very rarely observed between adult fish. We suggest that young and adult greenling have different social behaviour resulting from the interval of ontogenetic development, and not related to food abundance.     

Adaptive evolution of butterfly wing shape: from morphology to behaviour

Butterflies display extreme variation in wing shape associated with tremendous ecological diversity. Disentangling the role of neutral versus adaptive processes in wing shape diversification remains a challenge for evolutionary biologists. Ascertaining how natural selection influences wing shape evolution requires both functional studies linking morphology to flight performance, and ecological investigations linking performance in the wild with fitness. However, direct links between morphological variation and fitness have rarely been established. The functional morphology of butterfly flight has been investigated but selective forces acting on flight behaviour and associated wing shape have received less attention. Here, we attempt to estimate the ecological relevance of morpho‐functional links established through biomechanical studies in order to understand the evolution of butterfly wing morphology. We survey the evidence for natural and sexual selection driving wing shape evolution in butterflies, and discuss how our functional knowledge may allow identification of the selective forces involved, at both the macro‐ and micro‐evolutionary scales. Our review shows that although correlations between wing shape variation and ecological factors have been established at the macro‐evolutionary level, the underlying selective pressures often remain unclear. We identify the need to investigate flight behaviour in relevant ecological contexts to detect variation in fitness‐related traits. Identifying the selective regime then should guide experimental studies towards the relevant estimates of flight performance. Habitat, predators and sex‐specific behaviours are likely to be major selective forces acting on wing shape evolution in butterflies. Some striking cases of morphological divergence driven by contrasting ecology involve both wing and body morphology, indicating that their interactions should be included in future studies investigating co‐evolution between morphology and flight behaviour.     

Lek behaviour of Drosophila cnecopleura in Hawaii

ABSTRACT.

• 1 Male spacing, aggression, and female visitation are described for a lek of Drosophila cnecopleura Hardy within a Hawaiian rainforest. The lek was located at a single tree of Pisonia umbellifera, and the males present defended individual leaves as mating territories. Observations were made in October 1986 and April 1987.
• 2 Approximately twice as many males were present at the lek in October than in April. At both times, males generally perched on the lowest leaves in the lek. Individuals did not remain on particular leaves for long periods of time but frequently moved among different territories.
• 3 Males engaged in two types of aggressive encounters, face-offs and chases, and in all encounters the resident displaced the intruder.
• 4 Females were sighted at the lek 0.69 times/h in October and 0.23 times/h in April. Most of the leaves visited by females were among the lowest in the lek.

     

Nonconvergence in the evolution of primate life history and socio-ecology

The goal of this study was to investigate the extent of convergence in four basic life history and socio-ecological traits among the primates of Africa, Asia, South America and Madagascar. The convergence hypothesis predicts that similar abiotic conditions should result in similar adaptations in independent taxa. Because primates offer a unique opportunity among mammals to examine adaptations of independent groups to tropical environments, we collected information on body mass, activity pattern, diet and group size from all genera for quantitative tests of this hypothesis. We revealed a number of qualitative and quantitative differences among the four primate groups, indicating a lack of convergence in these basic aspects of life history and socio-ecology. Our analyses demonstrated that New World primates are on average significantly smaller than primates in other regions and characterized by a lack of species larger than about 10 kg. Madagascar harbours significantly more nocturnal species than the other regions and is home to all but one of the primates with irregular bursts of activity. Asia is the only region with strictly faunivorous primates, but lacks primarily gummivorous ones. The Neotropics are characterized by the absence of primarily folivorous primates. Solitary species are not represented in the New World, whereas solitary and pair-living species make up the majority of Malagasy primates. Lemurs live in significantly smaller groups than other primates, even after controlling for differences in body size. The lack of convergence among the major primate groups is neither primarily due to phylogenetic constraints as a result of founder effects, nor can it be sufficiently explained as a passive consequence of body size differences. However, because the role of adaptive forces, such as interspecific competition, predation or phenology in shaping the observed differences is largely unexplored, we conclude that it is premature to discard the convergence hypothesis without further tests.     

Body temperature and territory selection by males of the speckled wood butterfly (Pararge aegeria): what makes a forest sunlit patch a rendezvous site?

1. Insects locate mobile resources like prey items or mates using either sit‐and‐wait (‘perching’) or active (‘patrolling’) searching strategies. The sit‐and‐wait strategy can be accompanied by defending and monopolising a site through territorial behaviour. 2. The present study focuses on the territorial perching behaviour in males of the speckled wood butterfly (Pararge aegeria L.). Recent studies suggested that the selection of territories (i.e. sunlit patches on the forest floor) is driven by structural characteristics of the site that affect male visual detection. However, given that adult butterflies are heliothermic organisms and that forests provide a diverse array of light environments, it seems likely that thermal aspects may also be used for territory selection. 3. We tested whether used and unused sunlit patches differed in thermal profile under field conditions in a Belgian woodland. We also used dummy butterflies to quantify variation in operative thoracic temperature and to calculate heating rates within (i.e. different vegetation structures) and between patches. 4. Sunlit patches occupied by a territorial male were larger, and were more frequently characterised by low vegetation structures compared with empty sunlit patches. It took longer to reach optimal thorax temperature (starting from a fixed suboptimal body temperature) in small patches compared with large patches. 5. We suggest that aspects of visual detection need to be combined with thermal aspects to fully understand territory selection in the speckled wood butterfly, as synergetic and/or trade‐off effects of ambient temperature, solar radiation, and canopy/vegetation structure may be involved.     

Butterfly wing morphology variation in the British Isles: the influence of climate, behavioural posture and the hostplant-habitat

Gradients (isophenes) in modifications of butterfly wing morphology (colour, pattern, size) to the north and west of Britain are shown to correlate closely with contemporary environmental gradients, whereas their alleged formation as infra-specific units in Devensian refugia off western Britain is unsubstantiated. A model is described which explains the transformation in phenotypes in relationship to climate, especially ambient temperatures and radiation levels. In cooler, less predictable summer conditions to the north and west, selection has favoured modifications in adult phenotypes that maintain efficiency in thermoregulation, mate advertisement and predator escape. The form that wing modifications take depends mainly on basking posture (lateral, dorsal-absorption and reflectance), which determines the allocation and interaction of functions on different wing surfaces. It is also dependent on hostplant-habitat structure, which influences thermal stability and the milieu of predators and conspecifics, and other behavioural norms (mate-locating behaviour) and biological attributes (size, robustness, speed and mode of flight, chemical defences) which affect their relationships with predators and conspecifics. The significance of Quaternary palaeoenvironments to phenetic transformations is discussed as is the relevance of the model to the development of phenotypes in arctic endemic butterflies. Differences in phenotypes of butterflies which occupy arctic and temperate montane environments are also predicted by the model.     

Multiregional comparison of the ecological and phylogenetic structure of butterfly species richness gradients

Aim To examine butterfly species richness gradients in seven regions/countries and to quantify geographic mean root distance (MRD) patterns. My primary goal is to determine the extent to which an explanation for butterfly richness patterns based on tropical niche conservatism and the evolution of cold tolerance, proposed for the fauna of Canada and the USA, applies to other parts of the world. Location USA/Canada, Mexico, Europe/NW Africa, Transbaikal Siberia, Chile, South Africa and Australia. Methods Digitized range maps for butterfly species in each region were used to map richness patterns in summer (for all areas) and winter (for USA/Canada, Europe/NW Africa and Australia). A phylogeny resolved to subfamily was used to map the geographic MRD patterns. Regression trees and general linear models examined climatic and vegetation correlates of species richness and MRD within and among regions. Results Various combinations of climate and vegetation were strong predictors of species richness gradients within regions, but unresolved ‘regional’ factors contributed to the multiregional pattern. Regionally based differences in phylogenetic structure also exist, but MRD is negatively correlated with temperature both within and across areas. MRD patterns consistent with tropical niche conservatism occur in most areas. With a possible partial exception of Mexico, faunas in cold climates and in mountains are more derived than faunas in lowlands and tropical/subtropical climates. In USA/Canada, Europe and Australia, winter faunas are more derived than summer faunas. Main conclusions The phylogenetic pattern previously found in the USA and Canada is widespread in both the Northern and Southern Hemispheres, and niche conservatism and the evolution of cold tolerance is the likely explanation for the development of the global butterfly species richness gradient over evolutionary time. Contemporary climate also influences species richness patterns but is unlikely to be a complete explanation globally. The importance of climate is also manifested in the seasonal loss of more basal butterfly elements outside the tropics in winter.     

Variation in number of copulations in Theclini butterfly females (Lycaenidae: Lepidoptera)

The number of spermatophores inseminated in the bursa copulatrix of females of 21 species of Theclini butterflies varied from 0 to 38, a maximum much larger than reported so far. The variation of spermatophore number observed was analyzed on a phylogenetic tree recently determined, together with variation of wing colors and male mating tactics. Monandrous species tended to be sexually dimorphic in wing color and associated with male territoriality.     

Contest outcome in a territorial butterfly: the role of motivation

In many butterfly species, males compete over areas advantageous for encountering females. Rules for contest settlement are, however, largely unknown and neither morphological nor physiological traits can reliably predict the contest outcome. Here, we test the hypothesis that contests are settled in accordance with a motivation asymmetry. We staged contests between males of Pararge aegeria and after removing the resident, the non-resident was allowed (i) either to interact with a non-receptive female for 30 min (n = 30) or (ii) to spend 30 min alone in the cage (n = 30), after which the initial resident was reintroduced. The results show that males that had interacted with a female had a higher probability of becoming dominant and reversing contest outcome. Moreover, males that were faster to take over a vacant territory when the resident was removed were more likely to become dominant. Here, we show for the first time, to our knowledge, that frequent encounters with a mated female can increase male motivation to persist in a territorial contest in a butterfly. Further, we suggest that variation in intrinsic motivation reflects male eagerness to take over vacant territory. This study indicates that variation in resource value and motivational asymmetries are important for settling contests in butterflies.     

Ageing, reproductive value, and the evolution of lifetime fighting behaviour

The study of sexual selection has recently been enriched by an explicit life history focus. This lifetime perspective has much to offer the field; however, most existing interdisciplinary exchanges have restricted themselves to the context of mate choice and associated phenomena. The present study explores the consequences of an explicit life history view upon the evolution of male–male contest competition. Key to this view is the fact that fighting typically has lifetime consequences, and thus the costs involved with contests are best considered in lifetime currencies such as residual reproductive value. The rate of contemporary contest cost accrual may vary among contestants due to differences between them in what they 'stand to lose', in terms of future reproductive opportunities. It is also suggested that it may be fruitful to partition key life history parameters into components that are either dependent or independent of future choice of mating strategy. Using a simple simulation, it is demonstrated that the optimum pattern of lifetime contest participation may vary depending upon lifetime variation in 'strategy-independent' components of reproductive value (such as the rate of mortality experienced in contexts unrelated to mating behaviour). In line with previous models of age-specific sexual advertisement, increasing lifetime aggression appears as the most favoured strategy; however, young age aggression is predicted if resource holding potential is expected to decline with age irrespective of prior contest participation. The model is discussed, along with key concepts surrounding the life history viewpoint, in light of prior life history/sexual selection models and the existing empirical data regarding lifetime fighting strategies in the wild.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 565–578.     

Experimental test of the influence of light availability on the evolution of eye size and behaviour in Daphnia

There exists extensive variation in eye size. Much work has provided a connection between light availability and differences in eye size across taxa. Experimental tests of the role of the light environment on the evolution of eye size are lacking. Here, we performed a selection experiment that examined the influence of light availability on shifts in eye size and the connection between eye size and phototactic (anti-predator) behaviour in Daphnia. We set-up replicate experimental populations of Daphnia, repeatedly evaluated phenotypic shifts in eye size during the ~50-day experiment, and performed a common garden experiment at the end of the experiment to test for evolutionary shifts in eye size and behaviour. Our phenotypic analyses showed that eye size rapidly diverged between the light treatments; relative eye size was consistently larger in the low versus high light treatments. Selection on eye size was also modified by variation in density as increases in Daphnia density favoured a larger eye. However, we did not observe differences in eye size between the light treatments following two generations of common garden rearing at the end of the experiment. We instead observed strong shifts in anti-predator behaviour. Daphnia from the low light treatment exhibited decreased phototactic responses to light. Our results show that decreased light relaxes selection on anti-predator behaviour. Such trends provide new insights into selection on eye size and behaviour.     

Speciation and the evolution of behaviour in prosimians

A mechanism is suggested, based on a biphasic approach-withdrawal theory proposed byT. C. Schneirla, to account for how behaviours may be selected by environmental forces for transmission over generations. The basic postulates of Schneirla's theory are presented followed by an examination of some aspects of lorisid behaviour in the light of this theory.     

Male–female evolutionary game on mate-locating behaviour and evolution of mating systems in insects

We present a model of a male–female evolutionary game on mate-locating behaviour. Two major mating systems are considered: "lek polygyny" (in which males aggregate to wait for females searching for males) and "searching polygyny" (in which males search for females emerging or waiting for males). The model predicts that lek polygyny is favoured (i) when male survivorship during lekking is sufficiently higher than that during mate searching, (ii) when female survivorship while visiting a lek is sufficiently higher than at the emergence site, or (iii) when searching efficiency is higher at a lek than at an emergence site. Furthermore, the model shows that a reduction in the reproductive value of females later in the day, which prevents males from performing riskier mate-locating behaviour, can result in a change of mating system. In addition, mixed mating systems can be realized as transient states during this shift.     

Sex‐specific differences in spatial behaviour in the flesh fly Sarcophaga crassipalpis

Abstract Territoriality in the flesh fly Sarcophaga crassipalpis (Diptera: Sarcophagidae) is studied in the laboratory. In rectangular enclosures, male flies exhibit a lower tolerance (occupation of the same physical space) of same‐sex conspecifics than do females. In circular arenas, male flies show significantly higher levels of spatial separation among themselves (as determined from nearest neighbour analyses) than do females: males show a slight tendency towards a uniform distribution, whereas females exhibit a slight tendency towards clustering. The male spatial behaviour occurs during the photophase but not the scotophase, suggesting that visual cues are required for maintenance of inter‐individual spacing. No significant differences in male spacing behaviour occur between subjective day and subjective night in either constant dark or constant light conditions, suggesting that spatial patterning is not driven by a circadian rhythm.