Interference and the ideal free distribution: oviposition in a parasitoid wasp

The interference ideal free distribution (IFD) model of Sutherlandmakes a number of predictions that have yet to be tested andthat have implications for the validity of subsequent extensionsto the theory. We tested these predictions in a study usingdifferent densities of the parasitoid wasp, Venturia canescens,foraging on patches containing different densities of its host,Plodia interpunctella. Our results support a number of the interferenceIFD model's general predictions. Gain rate decreased becauseof increased interference at higher density. Although gain rateson the two patches differed slighdy, this would be expectedallowing for some sampling behavior and perceptual constraints.Early in each experiment when patch assessment is likely tooccur, wasp movement was higher and gain rates lower. However,the more specific prediction of Sutherland's model, that proportionalpatch use should be constant and independent of density, wasnot upheld. Contemporary IFD models use only one of severalequally valid potential relationships between gain rate, interference,and competitor density. The results of this study provide supportfor the additive model developed by Tregenza et al. (companionarticle).     

Male mating success, reproductive success and multiple paternity in a natural population of sand lizards: behavioural and molecular genetics data

Sand lizard Lacerta agilis females characteristically mate with several males which, in staged mating experiments, results in multiple paternity of the offspring. In order to investigate multiple paternity in a natural population and interpret male reproductive behaviours in terms of sired young, we sampled the blood of females, potential fathers and hatchlings, and determined paternity using multilocus DNA fingerprinting as well as the variation at a single locus detected by the probe (TC) _n . The paternity analyses were preceded by a laboratory experiment in which we established that the parental alleles identified by the single locus probe were inherited in a Mendelian way. Our molecular data demonstrated that 12 out of 13 males (92%) that sired offspring were correctly identified from the 56 sexually mature males in the population. Also smaller males were accepted as sexual partners by the females, but sired fewer young in competition with larger males and were less able to maintain prolonged post-copulatory mate guarding. This may result in that some sexually successful males are only observed inside a female's home range, but never in pair-association with the female.     

Transition from a random to an ideal free to an ideal despotic distribution: the effect of individual difference in growth

Individual differences in growth can lead to a monopolistic form of food competition. We studied the long-term transition in the mode of competition and the distribution of individuals between food patches of the cloned salmonid fish, Oncorhynchus masou ishikawae, in the laboratory. This transition was accompanied by growth depensation, i.e., the increase over time in the variance of size between individuals resulting from the differences in individual growth rates. The 120-cm experimental tanks were divided into two compartments (patches) between which an opaque partition was placed. Fish were able to move freely between the patches and therefore were able to assess the patch quality using long-term memory, but they were not able to see the food input in the other patch directly. The distribution between the two food patches, the amount of food gained, and the growth and the agonistic behavior of four groups of six individuals were observed over 4 weeks. We found that (1) within-group variation in body weight increased with time; (2) on average, the better patch was used by more individuals than predicted by a random distribution but fewer individuals than predicted by an ideal free distribution, and (3) the distribution and pattern of resource use by the fish changed over the 4-week experimental period from a random distribution to an ideal free distribution and finally to an ideal despotic distribution. We suggest that growth depensation causes the long-term change in the spatial distribution and pattern of resource use by competitors. Received: December 19, 2000 / Accepted: March 19, 2001     

Female-biased sexual size dimorphism in the yellow-pine chipmunk (Tamias amoenus): sex-specific patterns of annual reproductive success and survival

Sexual size dimorphism is ultimately the result of independent, sex-specific selection on body size. In mammals, male-biased sexual size dimorphism is the predominant pattern, and it is usually attributed to the polygynous mating system prevalent in most mammals. This sole explanation is unsatisfying because selection acts on both sexes simultaneously, therefore any explanation of sexual size dimorphism should explain why one sex is relatively large and the other is small. Using mark-recapture techniques and DNA microsatellite loci to assign parentage, we examined sex-specific patterns of annual reproductive success and survival in the yellow-pine chipmunk (Tamias amoenus), a small mammal with female-biased sexual size dimorphism, to test the hypothesis that the dimorphism was related to sex differences in the relationship between body size and fitness. Chipmunks were monitored and body size components measured over three years in the Kananaskis Valley, Alberta, Canada. Male reproductive success was independent of body size perhaps due to trade-offs in body size associated with behavioral components of male mating success: dominance and running speed. Male survival was consistent with stabilizing selection for overall body size and body size components. The relationship between reproductive success and female body size fluctuated. In two of three years the relationship was positive, whereas in one year the relationship was negative. This may have been the result of differences in environmental conditions among years. Large females require more energy to maintain their soma than small females and may be unable to maintain lactation in the face of challenging environmental conditions. Female survival was positively related to body size, with little evidence for stabilizing selection. Sex differences in the relationship between body size and fitness (reproductive success and survival) were the result of different processes, but were ultimately consistent with female-biased sexual size dimorphism evident in this species.     

Density-dependent habitat selection in muskrats: a test of the ideal free distribution model

Summary Two predictions of the ideal free distribution model, a null hypothesis of habitat selection, were examined using free-ranging muskrats. We rejected the prediction that the proportion of the animals found in each of five habitats was independent of population size. Data on over-winter occupancy of muskrat dwellings tend also to refute the prediction of equal fitness reward among habitats. Habitat type and water-level had a profound effect on the suitability of a site for settlement. We concluded that the observed pattern of muskrat distribution followed more closely an ideal despotic distribution where some individuals benefited from a higher fitness because of resource monopolization. Current theories of density-dependent habitat selection, which assume an ideal free distribution, would not apply to muskrats and possibly to many other mammal species.     

Effects of group size and personality on social foraging: the distribution of sheep across patches

Group cohesion in social animals foraging in patchy environmentsdepends on 2 opposite forces, intraspecific competition, andattraction. The decision to leave or stay in a group may varyaccording to the individual personality. We investigated therole of personality when feeding competition increases as aresult of increasing group size. Individuals referred as "bold"and "shy" were identified from an indoor exploration test accordingto their propensity to leave the group to explore a novel environment,using 12 novel objects placed at increasing distances from thegroup. Groups of 2, 4, 6, or 8 shy or bold sheep were testedin 45 x 5–m grass arenas, with one 5 x 5–m patchof preferred vegetation at each end. Sheep grazed on or closeto these patches, but the number grazing the patches seldomreached more than 4–5 individuals, suggesting that crowdingmight affect foraging at the highest densities. The smallestgroups grazed together on the same patch, but there was an increasinglikelihood of splitting into subgroups as group size increased,with equal-sized subgroups most commonly grazing the 2 patchessimultaneously. This effect was greatest in bold sheep, whichtended to split into subgroups at smaller group sizes than shysheep. This study provides new insight into the mechanisms bywhich group-living herbivores distribute themselves across patchyresources in a way that minimizes interference competition anddemonstrates the importance of individual variability for spatialorganization at the level of the group.     

Spatial relationships between mate acquisition probability and aggregation size in a gregarious coreid bug, (Colpula lativentris): A case of the ideal free distribution under perceptual constraints

Spatial relationships of mate acquisition probability for individuals of both sexes of a gregariously-mating coreid bug, Colpula lativentris, were studied in relation to aggregation size. Operational sex ratio was always strongly male biased. Mate acquisition probability of females was rather constant and independent of aggregation size, as predicted by an ideal free distribution. Moreover laboratory experiments showed that both multiple mating and rearing density little affected female fecundity, suggesting ideal free distribution of females in terms of reproductive success. On the other hand, mate acquisition probability of males was higher in larger aggregations, where more receptive females were available. This male discrepancy from an ideal free distribution was similar to the patterns predicted by an ideal free distribution under perceptual constraints (Abrahams, 1986), but not by that under unequal competitive ability.     

The ideal free distribution: a review and synthesis of the game-theoretic perspective

The Ideal Free Distribution (IFD), introduced by Fretwell and Lucas in [Fretwell, D.S., Lucas, H.L., 1970. On territorial behavior and other factors influencing habitat distribution in birds. Acta Biotheoretica 19, 16-32] to predict how a single species will distribute itself among several patches, is often cited as an example of an evolutionarily stable strategy (ESS). By defining the strategies and payoffs for habitat selection, this article puts the IFD concept in a more general game-theoretic setting of the “habitat selection game”. Within this game-theoretic framework, the article focuses on recent progress in the following directions: (1) studying evolutionarily stable dispersal rates and corresponding dispersal dynamics; (2) extending the concept when population numbers are not fixed but undergo population dynamics; (3) generalizing the IFD to multiple species.For a single species, the article briefly reviews existing results. It also develops a new perspective for Parker’s matching principle, showing that this can be viewed as the IFD of the habitat selection game that models consumer behavior in several resource patches and analyzing complications involved when the model includes resource dynamics as well. For two species, the article first demonstrates that the connection between IFD and ESS is now more delicate by pointing out pitfalls that arise when applying several existing game-theoretic approaches to these habitat selection games. However, by providing a new detailed analysis of dispersal dynamics for predator-prey or competitive interactions in two habitats, it also pinpoints one approach that shows much promise in this general setting, the so-called “two-species ESS”. The consequences of this concept are shown to be related to recent studies of population dynamics combined with individual dispersal and are explored for more species or more patches.     

Genetic estimates of annual reproductive success in male brown bears: the effects of body size, age, internal relatedness and population density

1. We studied male yearly reproductive success (YRS) and its determinants (phenotypic characteristics, age, population density) in two Scandinavian brown bear populations, using molecular techniques to determine paternity. 2. We found a significant difference in male YRS between the study areas, with lower YRS in the south than in the north. 3. In general, older and larger males had higher YRS. Older males may be more experienced in competition for reproduction (male dominance). Large body size is of direct benefit in male-male competition and of advantage in endurance competition for the access to females. 4. Age was relatively more important for YRS in the north and body size was more important in the south, due perhaps to differences in male age structure due to illegal killing. A single old male dominated the reproduction in the north during the study, which resulted most probably in the relatively higher importance of age in the north. In the south, with a more even male age structure, no single male was able to dominate, probably resulting in a more intense competition among males, with body size as the deciding factor. 5. Male YRS was correlated positively with population density. This may be related to the structure of the expanding bear population, with female densities declining towards the population edge. 6. Internal relatedness, a measure of genetic heterozygosity, was correlated negatively with YRS, suggesting that outbred individuals have a higher YRS. Individual heterozygosity at key or many loci may reflect male physical qualities and condition-sensitive traits, which may benefit males directly in contest or in sperm competition.     

It takes two to tango: reproductive skew and social correlates of male mating success in a lek-breeding bird

Variance in reproductive success among individuals is a defining characteristic of many social vertebrates. Yet, our understanding of which male attributes contribute to reproductive success is still fragmentary in most cases. Male–male reproductive coalitions, where males jointly display to attract females, are of particular interest to evolutionary biologists because one male appears to forego reproduction to assist the social partner. By examining the relationship between social behaviour and reproductive success, we can elucidate the proximate function of coalitions in the context of mate choice. Here, we use data from a 4-year study of wire-tailed manakins (Pipra filicauda) to provide molecular estimates of reproductive skew and to test the hypothesis that male–male social interactions, in the context of coordinated displays, positively influence a male''s reproductive success. More specifically, we quantify male–male social interactions using network metrics and predict that greater connectivity will result in higher relative reproductive success. Our data show that four out of six leks studied had significant reproductive skew, with success apportioned to very few individuals in each lek. Metrics of male social affiliations derived from our network analysis, especially male connectivity, measured as the number of males with whom the focal male has extended interactions, were strong predictors of the number of offspring sired. Thus, network connectivity is associated with male fitness in wire-tailed manakins. This pattern may be the result of shared cues used by both sexes to assess male quality, or the result of strict female choice for coordinated display behaviour.     

牧民对高寒牧区生态环境的感知——以甘南牧区为例

地处青藏高原的甘南牧区是典型的生态敏感区,牧民作为牧区最主要的经济活动主体与最基本的决策单位,其对生态环境的认识和响应影响着牧区的生态安全.采用参与性农户评估方法(PRA)对甘南牧区的97户牧民家庭进行了调查,并根据分布区域将其分成纯牧区牧户和半农半牧区牧户两组,就牧民的环境意识、对环境变化的感知、对环境变化原因的认识进行比较分析,基于分析结果,提出了相关的政策建议.研究结果表明:研究区牧民对生态环境重要性的认识非常明确,但在生产中却较少真正考虑对生态环境的影响;与20a前相比,85%的牧户认为生态环境有所恶化,纯牧区牧户的感受更为深刻;对于生态环境恶化的原因,超过40%的牧户认为是超载过牧,其次是破坏植被(22.68%)和气候变化(18.56%).两组牧户对于生态环境恶化原因的认识具有一定差别,纯牧区牧户认为主要原因是过度放牧及气候变化,半农半牧区牧户认为是过度放牧与破坏植被、滥垦土地.     

Variance in reproductive success and the opportunity for selection in a serially monogamous species: simulations of the mating system of <Emphasis Type="Italic">Tropheus</Emphasis> (Teleostei: Cichlidae)

Sexual selection is believed to play a major role in speciation processes of cichlid fishes in the East African Great Lakes, in particular Lakes Malawi and Victoria, by driving diversification in male nuptial coloration. In Lake Tanganyika, a high rate of color pattern diversification is found in the genus Tropheus, where more than 100, mostly allopatric, color morphs have been described. Whether color pattern differentiation in Tropheus could follow from sexual selection as well, has not yet been conclusively shown. Unlike typical sexually selected species, Tropheus are sexually monomorphic, establish temporary pair bonds and spawn monogamously. Variance in mating success among individuals would still allow for sexual selection, but due to their high population census sizes, parentage data from natural populations are difficult to obtain. Simulations designed according to existing data on mating behavior in Tropheus suggested that variance in male reproductive success can be substantial and lead to levels of opportunity for selection, which are similar to values estimated, for example, from natural populations of sexually dimorphic songbirds. Variance in male success was mostly affected by the way females perceived and reacted to differences between males, and to a lesser extent by the duration of components of the reproductive cycles of males and females. These results indicate that future work on the importance of sexual selection in Tropheus should concentrate on mate choice cues and female mate choice behavior, but does not depend critically on the acquisition of more detailed life history data.     

Foraging behaviour in tadpoles of the bronze frog<Emphasis Type="Italic">Rana temporalis</Emphasis>: Experimental evidence for the ideal free distribution

The ability of bronze frogRana temporalis tadpoles (pure or mixed parental lines) to assess the profitability of food habitats and distribute themselves accordingly was tested experimentally using a rectangular choice tank with a non-continuous input design. Food (boiled spinach) was placed at two opposite ends of the choice tank in a desired ratio (1:1, 1:2 or 1:4) to create habitat A and B. The tadpoles in Gosner stage 28–33, pre-starved for 24 h, were introduced in an open ended mesh cylinder placed in the center of the choice tank, held for 4 min (for acclimation) and then released to allow free movement and habitat selection. The number of tadpoles foraging at each habitat was recorded at 10, 15, 20, 25 and 30 min time intervals. The actual suitability,S _i (the food available in a habitat after colonization of tadpoles) of each habitat was obtained from the equationS _i =B _i−f _i (d _i) whereB _i is basic suitability (amount of food provided at each habitat before release of tadpoles),f _i is the rate of depletion of food (lowering effect) with introduction of each tadpole, andd _i is the density of tadpoles in habitati. The expected number of tadpoles at each habitat was derived from the actual suitability. With no food in the choice tank, movement of the tadpoles in the test arena was random indicating no bias towards any end of the choice tank or the procedure. In tests with a 1:1 food ratio, the observed ratio of tadpoles (11.71: 12.28) was comparable with the expected 12:12 ratio. The observed number of tadpoles in the habitats with a 1:2 food ratio was 8.71:15.29 and 7.87:16.13 for pure and mixed parental lines respectively. In both cases, the observed ratios were close to the expected values (7:17). Likewise, in experiments with a 1:4 food ratio, the observed number of tadpoles in the two habitats (10.78:37.22) did not differ significantly from the expected ratio of 7:41. In all tests, the number ofR. temporalis tadpoles matched ideally with habitat profitability (undermatching indexK ≜ 1. The study shows that tadpoles of the bronze frog exhibit an ideal free distribution while foraging regardless of whether they are siblings or non-siblings in a group, which correlates well with their group living strategy in nature.