Oviposition site selection and oviposition stimulation by conspecifics in the golden egg bug (<Emphasis Type="Italic">Phyllomorpha laciniata</Emphasis>): implications for female fitness

Phyllomorpha laciniata Vill. (Heteroptera, Coreidae) females lay eggs on the host plant and on the backs of conspecifics. Since egg survival is greater when eggs develop on the backs of conspecifics than when laid on plants, we predict that females should prefer to lay eggs on conspecifics. In addition, because conspecifics are a high-quality site that represents a limiting resource, females should experience oviposition stimulation upon an encounter with a conspecific. Our results reveal that, when both the host plant and conspecifics are available simultaneously, females lay eggs preferentially on conspecifics. The results also support the second prediction, since females housed with conspecifics lay more than twice the number of eggs than isolated females. Isolated females do not seem to retain eggs, suggesting that oviposition stimulation is the result of an acceleration of egg-maturation rates. Other studies have found oviposition stimulation by mating and have suggested that it is the result of male strategies to increase short-term male reproductive success at some cost to females. The evolutionary scenario of our model organism seems to be quite different since females benefit greatly from increasing egg laying when there are conspecifics, because the advantages in terms of offspring survival are likely to translate into substantial increases in female reproductive success.     

Malaria infection and host behavior: a comparative study of Neotropical primates

Parasites are ubiquitous in populations of free-ranging animals and impact host fitness, but virtually nothing is known about the factors that influence patterns of disease risk across species and the effectiveness of behavioral defenses to reduce this risk. We investigated the correlates of malaria infection (prevalence) in Neotropical primates using data from the literature, focusing on host traits involving group size, body mass, and sleeping behavior. Malaria is spread to these monkeys through anopheline mosquitoes that search for hosts at night using olfactory cues. In comparative tests that used two different phylogenetic trees, we confirmed that malaria prevalence increases with group size in Neotropical primates, as suggested by a previous non-phylogenetic analysis. These results are consistent with the hypothesis that larger groups experience increased risk of attack by mosquitoes, and counter to the hypothesis that primates benefit from the encounter-dilution effect of avoiding actively-seeking insects by living in larger groups. In contrast to non-phylogenetic tests, body mass was significant in fewer phylogeny-based analyses, and primarily when group size was included as a covariate. We also found statistical support for the hypothesis that sleeping in closed microhabitats, such as tree holes or tangles of vegetation, reduces the risk of malaria infection by containing the host cues used by mosquitoes to locate hosts. Due to the small number of evolutionary transitions in sleeping behavior in this group of primates, however, this result is considered preliminary until repeated with a larger sample size. In summary, risk of infection with malaria and other vector-borne diseases are likely to act as a cost of living in groups, rather than a benefit, and sleeping site selection may provide benefits by reducing rates of attack by malaria vectors.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Mycorrhizal benefit differs among the sexes in a gynodioecious species

Both plant sex and arbuscular mycorrhizal (AM) symbiosis influence resource acquisition and allocation in plants, but the interaction between these two components is not well established. As the different plant sexes differ in their resource needs and allocation patterns, it is logical to presume that they might differ in their relationship with AM as well. We investigate whether the association with AM symbiosis is different according to the host plant sex in the gynodioecious Geranium sylvaticum, of which, besides female and hermaphrodite plants, intermediate plants are also recognized. Specifically, we examine the effects of two different AM fungi in plant mass allocation and phosphorus acquisition using a factorial greenhouse/common garden experiment. Cloned G. sylvaticum material was grown in symbiosis with AM fungi or in non-mycorrhizal condition. We evaluated both the symbiotic plant benefit in terms of plant mass and plant P content and the fungal benefit in terms of AM colonization intensity in the plant roots and spore production. Our results suggest that G. sylvaticum plants benefit from the symbiosis with both AM fungal species tested but that the benefits gained from the symbiosis depend on the sex of the plant and on the trait investigated. Hermaphrodites suffered most from the lack of AM symbiosis as the proportion of flowering plants was dramatically reduced by the absence of AM fungi. However, females and intermediates benefited from the symbiosis relatively more than hermaphrodites in terms of higher P acquisition. The two AM fungal species differed in the amount of resources accumulated, and the fungal benefit was also dependent on the sex of the host plant. This study provides the first evidence of sex-specific benefits from mycorrhizal symbiosis in a gynodioecious plant species.     

Current Constraints and Future Directions in Estimating Coextinction

Abstract: Coextinction is a poorly quantified phenomenon, but results of recent modeling suggest high losses to global biodiversity through the loss of dependent species when hosts go extinct. There are critical gaps in coextinction theory, and we outline these in a framework to direct future research toward more accurate estimates of coextinction rates. Specifically, the most critical priorities include acquisition of more accurate host data, including the threat status of host species; acquisition of data on the use of hosts by dependent species across a wide array of localities, habitats, and breadth of both hosts and dependents; development of models that incorporate correlates of nonrandom host and dependent extinctions, such as phylogeny and traits that increase extinction‐proneness; and determination of whether dependents are being lost before their hosts and adjusting models accordingly. Without synergistic development of better empirical data and more realistic models to estimate the number of cothreatened species and coextinction rates, the contribution of coextinction to global declines in biodiversity will remain unknown and unmanaged.     

Rank-related partner choice in the fission–fusion society of the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>)

When social partners vary in their relative value, individuals should theoretically initiate partnerships with conspecifics of the highest value. Here, we tested this prediction in a wild population of spotted hyenas (Crocuta crocuta). Crocuta live in complex, fission–fusion societies structured by dominance hierarchies in which individuals vary greatly in their value as social companions. Because patterns of association among Crocuta reflect social preferences, we calculated association indices (AIs) to examine how social rank influences intrasexual partner choice among unrelated adults of both sexes. The highest-ranking individuals were generally most gregarious in both sexes. Females associated most often with dominant and adjacent-ranking females. Females joined subgroups based on the presence of particular conspecifics such that subordinates joined focal females at higher rates than did dominants. Dominants benefit from associations with subordinates by enjoying priority of access to resources obtained and defended by multiple group members, but the benefits of these associations to subordinates are unknown. To investigate this, we tested three hypotheses suggesting how subordinates might benefit from rank-related partner choice among unrelated females. We found that subordinates who initiated group formation benefited by gaining social and feeding tolerance from dominants. However, rates at which dominants provided coalitionary support to subordinates did not vary with AIs. Overall, our data resemble those documenting patterns of association among cercopithecine primates. We consider our results in light of optimal reproductive skew theory, Seyfarth’s rank attractiveness model, and biological market theory. Our data are more consistent with the predictions of Seyfarth’s model and of biological market theory than with those of skew theory.     

Do social spiders cooperate in predator defense and foraging without a web?

Nearly all social spiders spin prey-capture webs, and many of the benefits proposed for sociality in spiders, such as cooperative prey capture and reduced silk costs, appear to depend on a mutually shared web. The social huntsman spider, Delena cancerides (Sparassidae), forms colonies under bark with no capture web, yet these spiders remain in tightly associated, long-lasting groups. To investigate how the absence of the web may or may not constrain social evolution in spiders, we observed D. cancerides colonies in the field and laboratory for possible cooperative defense and foraging benefits. We observed spiders’ responses to three types of potential predators and to prey that were introduced into retreats. We recorded all natural prey capture over 447 h both inside and outside the retreats of field colonies. The colony’s sole adult female was the primary defender of the colony and captured most prey introduced into the retreat. She shared prey with younger juveniles about half the time but never with older subadults. Spiders of all ages individually captured and consumed the vast majority of prey outside the retreat. Young spiders benefited directly from maternal defense and prey sharing in the retreat. However, active cooperation was rare, and older spiders gained no foraging benefit by remaining in their natal colony. D. cancerides does not share many of the benefits of group living described in other web-building social spiders. We discuss other reasons why this species has evolved group living.     

Behavioral transitions with the evolution of cooperative nest founding by harvester ant queens

Research on the evolution of cooperative groups tends to explore the costs and benefits of cooperation, with less focus on the proximate behavioral changes necessary for the transition from solitary to cooperative living. However, understanding what proximate changes must occur, as well as those pre-conditions already in place, is critical to understanding the origins and evolution of sociality. The California harvester ant Pogonomyrmex californicus demonstrates population-level variation in colony founding over a close geographic range. In adjacent populations, queens either found nests as single individuals (haplometrosis) or form cooperative groups of nonrelatives (pleometrosis). We compared aggregation, aggression, and tolerance of queens from one pleometrotic and two haplometrotic populations during nest initiation, to determine which behaviors show an evolutionary shift and which are present at the transition to pleometrosis. Surprisingly, within-nest aggregative behavior was equally present among all populations. In nesting boxes with multiple available brood-rearing sites, both queen types readily formed and clustered around a single common brood pile, suggesting that innate attraction to brood (offspring) facilitates the transition to social aggregation. In contrast, queens from the three populations differed in their probabilities of attraction on the ground to nest sites occupied by other queens and in levels of aggression. Our results suggest that some key behavioral mechanisms facilitating cooperation in P. californicus are in place prior to the evolution of pleometrosis and that the switch from aggression to tolerance is critical for the evolution of stable cooperative associations.     

Offspring sex ratio response to host size in the parasitoid wasp Spalangia endius

The host size model, an adaptive model for maternal manipulation of offspring sex ratio, was examined for the parasitoid wasp Spalangia endius. In a Florida strain, as the model predicts, daughters emerged from larger hosts than sons, but only when mothers received both small and large hosts simultaneously. The pattern appeared to result from the mother's ovipositional choice and not from differential mortality of the sexes during development. If sex ratio manipulation is adaptive in the Florida strain, it appears to be through a benefit to daughters of developing on large hosts rather than through a benefit to sons of developing on small hosts. Both female and male parasitoids were larger when they developed on larger hosts. For females, developing on a larger host (1) increased offspring production, except for the largest hosts, (2) increased longevity, (3) lengthened development, and (4) had no effect on wing loading. For males, development on a larger host had no effect on any measure of male fitness – mating success, longevity, development duration, or wing loading. In contrast, a strain from India showed no difference in the size of hosts from which daughters versus sons emerged, although both female and male parasitoids were larger when they developed on larger hosts. These results together with previous studies of Spalangia reveal no consistent connection between host-size-dependent sex ratio and host-size-dependent parasitoid size among strains of S. endius or among species of Spalangia. Received: 28 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Differential use of conspecific-derived information by sexual and asexual parasitic wasps exploiting partially depleted host patches

When foraging partially depleted patches (i.e., a fraction of hosts are already parasitized), female parasitoids must decide: 1—whether to superparasitize, and 2—whether to stay in their current patch (thus missing the opportunity of finding a better patch elsewhere). To make these decisions, parasitoids may rely on different cues, produced both by the environment and by conspecifics. Animals thriving in different environments may differ in cues they use. In the solitary parasitoid Venturia canescens, thelytokous (asexual) and arrhenotokous (sexual) individuals are found in two contrasting environments. Thelytokous females, from anthropogenic conditions, are known to cope with superparasitism in an adaptive way. On the other hand, little is known about superparasitism by arrhenotokous females. We compared the host exploitation strategies of thelytokous and arrhenotokous females in partially depleted patches. Hosts parasitized by thelytokous females were more frequently avoided than those parasitized by arrhenotokous females, suggesting a stronger chemical marking of the former. Only thelytokous females used information from conspecifics for patch-leaving decisions. The conformity of the differences in the behavior of thelytokous and arrhenotokous females with the environmental conditions they experience in their habitat is discussed.     

Ecological drivers of group living in two populations of the communally rearing rodent, Octodon degus

Intraspecific variation in sociality is thought to reflect a trade-off between current fitness benefits and costs that emerge from individuals' decision to join or leave groups. Since those benefits and costs may be influenced by ecological conditions, ecological variation remains a major, ultimate cause of intraspecific variation in sociality. Intraspecific comparisons of mammalian sociality across populations facing different environmental conditions have not provided a consistent relationship between ecological variation and group-living. Thus, we studied two populations of the communally rearing rodent Octodon degus to determine how co-variation between sociality and ecology supports alternative ecological causes of group living. In particular, we examined how variables linked to predation risk, thermal conditions, burrowing costs, and food availability predicted temporal and population variation in sociality. Our study revealed population and temporal variation in total group size and group composition that covaried with population and yearly differences in ecology. In particular, predation risk and burrowing costs are supported as drivers of this social variation in degus. Thermal differences, food quantity and quality were not significant predictors of social group size. In contrast to between populations, social variation within populations was largely uncoupled from ecological differences.     

Parent-absent begging in the Brown-headed Cowbird (<Emphasis Type="Italic">Molothrus ater</Emphasis>): the role of short-term need and nestmate size

Although it is well-established that nestlings of many altricial species beg when parents are away from the nest, we have a poor understanding of parent-absent begging in brood parasites, including the proximate factors that may influence begging frequency and intensity. In this study, I examined how parent-absent begging was influenced by competitive asymmetries between host and Brown-headed Cowbird (Molothrus ater) nestlings under disparate levels of short-term need. Food-deprived cowbird nestlings begged more frequently and for a greater proportion of parent-absent period than when food-supplemented, with similar patterns observed in hosts of different sizes. In contrast, three metrics of cowbird begging intensity varied relative to host size but not due to differences in short-term need. Cowbirds consistently begged more frequently and intensively than host nestlings for a given level of short-term need, providing evidence that cowbird begging displays are more frequent and intense than non-parasitic nestlings during both feeding visits and parent-absent periods. In sum, the frequency of begging by cowbirds was only influenced by short-term need, whereas begging intensity during parent-absent events was only influenced by the host against which cowbirds competed. This study demonstrates that host size and short-term need had differing influences on the frequency and intensity of parent-absent begging in cowbirds, although both factors are likely important in limiting the evolution of parent-absent begging in cowbirds. Because it appears to provide no immediate benefits yet may decrease fitness, parent-absent begging should be included in future theoretical models investigating the evolution of begging displays in nestling birds.     

Intraspecific brood parasitism as a conditional reproductive tactic in the treehopper<Emphasis Type="Italic"> Publilia concava</Emphasis>

In species exhibiting egg guarding as well as communal egg laying, females may adopt the strategy of laying eggs in the nests of conspecifics and leaving without providing care (termed intraspecific brood parasitism). This study is the first to describe such a behavior in the insect Publilia concava (Hemiptera: Membracidae) through field studies that followed 849 marked females across 1,828 brood associations. While brood parasitism increased the total number of eggs in a host brood, it did not reduce the overall hatching success of host broods. Solitary females exhibited a range of guarding durations while parasitic females rarely remained to guard eggs. Females exhibiting the parasitic tactic increased their lifetime number of clutches without decreasing the number of solitary clutches that they were able to initiate. Estimates of egg number for these individual broods revealed that females adopting the parasitic tactic (in addition to solitary breeding) had higher lifetime fecundity relative to females that did not parasitize. In females that exhibited both tactics (solitary and parasitic), the parasitic tactic yielded a higher rate of oviposition. The major component of oviposition rate was the time to find hosts and this time decreased with increasing host availability across 222 replicate groups. Females exhibited a shift toward the parasitic tactic when host broods were more abundant (i.e. in larger groups and later in the season). However, the time to find hosts increased as the frequency of the parasitic tactic increased, suggesting that this tactic may be maintained through negative frequency dependence. The results of this study suggest that brood parasitism may be the preferred tactic, as part of a conditional strategy, when hosts are readily available with solitary breeding being the preferred tactic when hosts are in short supply.     

Constraints on foraging success due to resource ecology limit colony productivity in social insects

The benefit of group living is a fundamental question in social evolution. For sociality to evolve, each individual must gain in terms of some fitness component by living in larger groups. However, in social insects, a decrease in per capita success in brood production has been observed in larger groups. While it has been proposed that this decrease could be outweighed by an increase in the predictability of success, a functional basis to this hypothesis has so far never been demonstrated. In this paper, using foraging economics as a functional proxy to colony productivity, we construct a model to explore how number of foragers in the colony interacts with the ecology of resources to influence per capita foraging success and its predictability. The results of the model show that there is no increase in per capita foraging success in larger colonies under most circumstances, though there is an increase in its predictability. We then test the model with empirical data on the foraging behavior of the primitively eusocial wasp, Ropalidia marginata. The consistency between the data and the model suggests that foraging economics could provide a robust functional basis in explaining the relationship between colony size and productivity.     

Shiny cowbirds share foster mothers but not true mothers in multiply parasitized mockingbird nests

Obligate brood parasitic birds, such as cowbirds, evade parental care duties by laying their eggs in the nests of other species. Cowbirds are assumed to avoid laying repeatedly in the same nest so as to prevent intrabrood competition between their offspring. However, because searching for host nests requires time and energy, laying more than one egg per nest might be favoured where hosts are large and can readily rear multiple parasites per brood. Such ‘repeat parasitism’ by females would have important consequences for parasite evolution because young parasites would then incur indirect fitness costs from behaving selfishly. We investigated shiny cowbird (Molothrus bonariensis) parasitism of a large host, the chalk-browed mockingbird (Mimus saturninus), in a population where over 70 % of the parasitized mockingbird nests receive multiple cowbird eggs. We assessed egg maternity directly, using cameras at nests to film the laying of individually-marked females. We also supplemented video data with evidence from egg morphology, after confirming that each female lays eggs of a consistent appearance. From 133 eggs laid, we found that less than 5 % were followed by the same female visiting the nest to lay again or to puncture eggs. Multiple eggs in mockingbird nests were instead the result of different females, with up to eight individuals parasitizing a single brood. Thus, while cowbird chicks regularly share mockingbird nests with conspecifics, these are unlikely to be their maternal siblings. Our results are consistent with shiny cowbird females following a one-egg-per-nest rule, even where hosts can rear multiple parasitic young.     

Parental investment without kin recognition: simple conditional rules for parent–offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely genetic offspring can be mediated through a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to “recognize” one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Brood mate eviction or brood mate acceptance by brood parasitic nestlings? An experimental study with the non-evictor great spotted cuckoo and its magpie host

Some avian brood parasitic nestlings are highly virulent, destroying all host eggs or nestmates, while others accept growing up together with host nestmates. The traditional idea was that all brood parasitic nestlings would benefit from being alone in the host nest. Thus, why do nestlings of some brood parasitic species accept the company of host offspring in the nest? The trade-off hypothesis suggests that brood parasites must balance the costs and benefits of killing host young because of two major potential costs: risk of nest desertion and loss of begging assistance. Here, we test this hypothesis in a non-evictor cuckoo species, the great spotted cuckoo Clamator glandarius and its main host, the magpie Pica pica, by manipulating brood size (1–3 nestlings) and brood composition (only cuckoo, only magpie or mixed) during three consecutive breeding seasons. None of the broods were abandoned by host parents, and cuckoo nestlings alone in the nest tended to grow faster (i.e. wing length). Thus, none of the predictions of the two potential costs on which the trade-off hypothesis is based apply to the great spotted cuckoo–magpie system. Our experimental study could not directly test why chick killing has not evolved in great spotted cuckoos, but the results point in the direction of several possibilities. We suggest that chick killing in great spotted cuckoos may not be adaptive mainly because another, less costly strategy (i.e. outcompeting host nestmates for food), is efficient for successful parasitism of magpie hosts.     

Flight initiation distance decreases during social activity in lizards (<Emphasis Type="Italic">Sceloporus virgatus</Emphasis>)

Flight initiation distance, the predator–prey distance when escape begins, is predicted by escape theory to decrease if fleeing entails loss of benefits. Shortening of flight initiation distance during social interactions is known only in males and only in a few species. In a previous study, male, but not female, Sceloporus virgatus lizards had shorter flight initiation distance when interacting with tethered conspecifics. Females in that study were not gravid or close to ovulating. I predicted that flight initiation distance would be shorter in gravid females that perform sidle-hopping displays to reject courtship than in lone females. I tested this prediction and examined effects of social interactions by males with free-ranging conspecifics to ensure that previous findings were not artifacts of tethering and experimental introduction of conspecifics. Flight initiation distance was shorter in females when interacting with males than when alone; it was also shorter in males interacting with either sex. Thus, when beneficial for reproductive reasons, social interaction affects flight initiation distance in females, but at other times, it does not. Lesser shortening of flight initiation distance in females than males may be a consequence of greater social benefit to males and protection of reproductive investment by females.     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Parental investment without kin recognition: Simple conditional rules for parent-offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely offspring can be instantiated with a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to 'recognize' one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

The benefit of obligate versus facultative strategies in a shrimp–goby mutualism

Obligate mutualists are predicted to benefit more from mutualism than facultative mutualists and harbor traits that help derive this extra benefit. I tested these predictions with a shrimp–goby mutualism. Individual shrimp (Alpheus floridanus) construct burrows that are shared with individual gobiid fishes that warn shrimp when emergence from burrows is unsafe. The benefit to gobies is refuge from predation. I compared predator avoidance performance of obligate (Nes longus) and facultative (Ctenogobius saepepallens) shrimp-associated gobies with access to a shell, a shrimp burrow, or no shelter. The two gobies performed similarly with shells and no shelter, but N. longus outperformed C. saepepallens with shrimp burrows. Thus, N. longus benefits more from mutualism than C. saepepallens. Also, N. longus has four behavioral traits that likely allow it to benefit from mutualism more than does C. saepepallens: (1) diving into the nearest shrimp burrow when confronted with predators, (2) long flight initiation distance, (3) long time-until-re-emergence after taking refuge, and (4) remaining close to shelter. The latter three likely come with a cost to foraging or mate acquisition as they limit searching range outside the burrow and detract from time spent outside burrows. Thus, N. longus likely harbors traits that allow it to deal with these costs. I discuss a framework for determining whether traits crucial to obligate association with shrimp (both for reaping benefits and dealing with costs) are pre-adaptations or are created through coevolution with shrimp.