Alternative models of conspecific attraction in flies and crabs

Animals are often attracted to one another when selecting habitats,but little is known about the rules governing conspecific attraction.We use Akaike Information Criterion to evaluate alternativemodels of the effects of conspecifics on individual choice inthe context of habitat selection. One set of models was testedusing data collected on virgin female flies, Drosophila melanogaster,selecting habitats in the laboratory; a second set of modelswas tested using data collected on crabs, Ocypode rotundata,selecting foraging patches in the field. Patterns of space usein the flies were most consistent with models indicating thatindividuals were attracted to other females that selected trapsduring the same hour, as well as to cues left by females thathad entered traps during the previous hour. Results for thecrabs were most consistent with a model which assumes that individualspreferred to join the patch with the most crabs but that theirability to assess the number of crabs in alternative patcheswas constrained by basic psychophysical principles (Weber'slaw). These results provide support for hypotheses about thefunctional significance of conspecific attraction in the fliesand the crabs and illustrate the richness of information aboutconspecific attraction that can be obtained when the same dataset is confronted with alternative models of the ways that animalsrespond to one another when selecting habitats.     

The effect of conspecific attraction on metapopulation dynamics

Random dispersal direction is assumed in all current metapopulation models. This assumption is called into question by recent experiments demonstrating that some species disperse preferentially to sites occupied by conspecifies. We incorporate conspecific attraction into two metapopulation models which differ in type of dispersal, the Levins model and a two-dimensional stepping-stone model. In both models, conspecific attraction lowers the proportion of occupied habitat patches within a metapopulation at equilibrium.     

The evolution of conspecific gamete precedence and its effect on reinforcement

Conspecific gamete precedence, the usage of conspecific sperm by a female that mates with both a conspecific and a heterospecific male, has been found in many taxa. We construct a population genetic model to examine the evolution of conspecific gamete precedence and its coevolution with premating isolation in the process of reinforcement. Our findings suggest that conspecific gamete precedence can evolve via a process very similar to reinforcement. We explore the nature of the selection against hybridization necessary to drive this evolution. Moreover, our results confirm the prediction of Marshall et al. (Trends Ecol. Evol. 2002;17:558-563) that conspecific gamete precedence will inhibit the evolution of reinforcement between two species. We further find that reinforcement will inhibit the evolution of conspecific gamete precedence. Both reinforcement and conspecific gamete precedence increase reproductive isolation and contribute to the process of speciation. We discuss factors that may affect which of these phenomena are likely to become predominant between incipient species.     

Managing breeding forest songbirds with conspecific song playbacks

Recent studies confirm that several territorial songbird species use conspecific cues, including song, when selecting habitat. We conducted a controlled experiment with a population of black-throated blue warblers Dendroica caerulescens , a species of concern in boreal hardwood transition forests, to determine whether song playbacks influence settlement patterns. We tested whether conspecific song broadcast during spring arrival attracts black-throated blue warblers during territory establishment, and whether song playbacks shift territories from the previous year's breeding distributions. The results of our experiment demonstrate that song playbacks significantly increased territorial occupancy and shifted territories closer to the speakers. Loss of breeding habitat is implicated in population declines of many songbirds, and our results suggest that song playbacks should be further investigated as a tool for mitigating the effects of disruptive management actions. Additional work should consider the spatial arrangement of playback speakers to avoid attracting birds from safe areas. The demographic consequences of conspecific attraction must also be explored before this technique is widely implemented.     

Allee effects and conspecific cueing jointly lead to conspecific attraction

Conspecific attraction is the preferential settlement into habitat patches with conspecifics. To be a good proximate strategy, fitness gains from settling with conspecifics must outweigh the costs of higher conspecific densities, such as intraspecific competition. Two types of benefits have been proposed to explain conspecific attraction: Allee effects (i.e., positive density dependence) and conspecific cueing (using conspecifics as an indicator of habitat quality). I present empirical evidence for conspecific attraction in the settlement of the porcelain crab, Petrolisthes cinctipes Randall (Anomura: Porcellanidae). Previous work demonstrated that P. cinctipes experiences strong intraspecific competition and that both Allee effects and conspecific cueing are present in P. cinctipes life-history. I developed an empirically-based fitness model of the costs and benefits of settling with conspecifics. Based on this model, I simulated optimal settlement to habitat patches that varied in conspecific density and habitat quality, where the correlation between density and habitat quality determined the level of conspecific cueing. I tested whether Allee effects alone, conspecific cueing alone, or Allee effects and conspecific cueing together could provide an ultimate explanation for the proximate settlement behavior of P. cinctipes. The settlement simulation was consistent with empirical settlement only when Allee effects and conspecific cueing were both included. Three life-history features are critical to this conclusion: (1) fitness is maximized at intermediate density, (2) fitness depends on the decisions of previous settlers, and (3) conspecific density provides good information about habitat quality. The quality of information garnered from conspecifics determines whether conspecific attraction is a good proximate strategy for settlement. I present a graphical illustration demonstrating how Allee effects and conspecific cueing work together to influence fitness, providing a conceptual framework for other systems.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

The use of conspecific reproductive success for breeding patch selection in terrestrial migratory species

Classical models of breeding habitat selection rarely deal with the question of information gathering for patch quality assessment. In this paper, we present two models comparing the fitness outcomes of behavioural strategies based on conspecific reproductive success as a cue to assess local environmental quality before selecting a new breeding habitat. The models deal with two phases of the life-cycle of a territorial migratory species: recruitment to a breeding population (model 1) and breeding site fidelity of subsequent breeding attempts (model 2). The first model shows that prospecting breeding patches before recruiting is the best strategy if the environment is predictable and contains a low proportion of good patches, even if it implies losing a breeding opportunity. The second model shows that dispersing after a breeding attempt according to the patch's breeding success rather than the individual's own success is the bests own success is the best strategy if the environment is patchy. These results underline the importance of studying the spatio-temporal variations of factors affecting reproductive success when considering the importance of habitat selection strategies based on conspecifics. Moreover, they allow the understanding of individual behaviour patterns observed in natural populations and their potential consequences at the metapopulation level.     

A game theoretical approach to conspecific brood parasitism

We constructed a game theoretical model to predict optimal patternsof egg laying in systems where individuals lay in the nestsof others as well as in their own nests. We show that decreasingthe effect of position within an egg-laying sequence on theworth of an egg should lead to reduced parasitism. Indeed,parasitism can only flourish if the worth of an egg to its biological parent declines with the total number of eggs laid in that nest.Further, we found that increasing the intrinsic costs of eggproduction should lead to an increased propensity for conspecificbrood parasitism. The model also predicts that variation inhosts' ability to reject parasitic eggs has little effect on parasitism until this ability is well developed.     

Distribution dynamics of a great bustard metapopulation throughout a decade: influence of conspecific attraction and recruitment

Dispersing individuals can use conspecifics as indicators of habitat quality and aggregate at traditionally occupied sites, leaving other favourable patches unoccupied. Here we test the predictions of the conspecific-based habitat selection hypothesis on a Spanish great bustard (Otis tarda) metapopulation, currently fragmented due to recent human-induced habitat changes. The number of birds had increased by 23% between 1988 and 1998, but not consistently among leks. Leks that were large in 1988 increased, while those that were small decreased, which suggests that dispersing individuals used the numbers of conspecifics as cues for breeding-site selection. Moreover, leks with high productivity increased, while those with low productivity decreased. Finally, lek distribution was markedly stable throughout the decade, with no establishment of new leks, and suitable habitat patches remained unoccupied, as predicted by the conspecific attraction hypothesis. These results were corroborated by a simulation model which incorporated natal dispersal rates between leks as obtained through radio-tracking of 15 birds that survived throughout their 4-year dispersal period. In conclusion, in spite of the apparent increase in total numbers throughout the decade, both conspecific attraction and local differences in reproductive success contributed to a more aggregated distribution, increasing the species' vulnerability to local catastrophes, and the risks of reduced genetic diversity and extinction of small leks.     

Harvesting resources in groups or alone: the case of renewing patches

Group foraging has been proposed to be the most efficient mannerwith which to exploit habitats with renewing patches as individualsin groups are less likely to revisit patches that have alreadybeen exploited recently by others. However, to avoid a group-selectionargument, it is necessary to compare the success of solitaryand group foraging tactics when each competes with the other.We used a genetic algorithm approach to examine the costs andbenefits of exploiting renewing resources in a spatially andtemporally explicit habitat, thus controlling the time courseof resource renewal and including the time cost of travelingbetween patches, which may be a significant factor for groupforagers that deplete patches more quickly. Results indicatethat group foragers fare more poorly than an equivalent numberof solitary foragers in the same habitat unless the rate ofresource renewal is very low. The low revisitation rate by groupforagers allows resources to replenish more fully, thus maintainingthe resource level across the habitat at a higher level. Incontrast, solitary foragers, who revisit previously exploitedpatches more often, maintain the same resources at a lower level.Nevertheless, a pure population of group foragers can be readilyinvaded by solitary foragers even when the rate of renewal isat low levels. We conclude that while group foraging may bean efficient tactic to exploit renewing resources, it is nota stable strategy under the circumstances examined in this model.     

Heterospecific attraction and food resources in migrants' breeding patch selection in northern boreal forest

We studied experimentally how heterospecific attraction may affect habitat selection of migrant passerine birds in Finnish Lapland. We manipulated the densities of resident tit species (Parus spp.). In four study plots residents were removed before the arrival of the migrants in the first study year, and in four other plots their densities were increased by releasing caught individuals. In the second year the treatments of the areas were reversed, allowing paired comparisons within each plot. We also investigated the relative abundance of arthropods in the study plots by the sweep-net method. This allowed us to estimate the effect of food resources on the abundance of birds. The heterospecific attraction hypothesis predicts that densities of migrant species (especially habitat generalists) would be higher during increased resident density. Results supported this prediction. Densities and number of the most abundant migrant species were significantly higher when resident density was increased than when they were removed. On the species level the redwing (Turdus iliacus) showed the strongest positive response to the increased abundance of tits. Migrant bird abundances seemed not to vary in parallel with relative arthropod abundance, with the exception of the pied flycatcher (Ficedula hypoleuca) which showed a strongly positive correlation with many arthropod groups. The results of the experiment indicate that migrants can use resident tit species as a cue to a profitable breeding patch. The relationship between the abundance of the birds and arthropods suggests that annual changes in food resources during the breeding season probably do not have a very important effect on bird populations in these areas. The results stress the importance of positive interspecific interactions in structuring northern breeding bird communities. Received: 1 September 1997 / Accepted: 22 January 1998     

Analysis of spatial genetic structure in an expanding Pinus halepensis population reveals development of fine-scale genetic clustering over time

We analysed the change of spatial genetic structure (SGS) of reproductive individuals over time in an expanding Pinus halepensis population. To our knowledge, this is the first empirical study to analyse the temporal component of SGS by following the dynamics of successive cohorts of the same population over time, rather than analysing different age cohorts at a single time. SGS is influenced by various factors including restricted gene dispersal, microenvironmental selection, mating patterns and the spatial pattern of reproductive individuals. Several factors that affect SGS are expected to vary over time and as adult density increases. Using air photo analysis, tree-ring dating and molecular marker analysis we reconstructed the spread of reproductive individuals over 30 years beginning from five initial individuals. In the early stages, genotypes were distributed randomly in space. Over time and with increasing density, fine-scale (< 20 m) SGS developed and the magnitude of genetic clustering increased. The SGS was strongly affected by the initial spatial distribution and genetic variation of the founding individuals. The development of SGS may be explained by fine-scale environmental heterogeneity and possibly microenvironmental selection. Inbreeding and variation in reproductive success may have enhanced SGS magnitude over time.