Response of Solidago canadensis clones to competition

Summary Transplants of ten Solidago canadensis clones were grown under high and low competition in the field to determine whether clones differed in survival, growth, and reproduction under natural conditions. Transplants had higher probability of survival and flowering and were larger in all measures of size when competition was experimentally reduced. Clones differed in almost all these measures of success, but only when variance among transplants within clones was reduced by excluding transplants that experienced heavy herbivore damage. Differences among clones were more apparent under low competition than under high competition, despite higher coefficients of variation within clones under low competition. Adjusting transplant size for initial size (parent ramet rhizome mass) did not change these results, although clones did differ in parent rhizome mass. All of these results suggest that there is little potential for selection to discriminate among these clones. Despite the strong differences in transplant performance between the competition treatments and among clones, the clones did not differ in competitive ability-almost none of the clone x competition interactions were significant. In addition, the measures of success of each clone were usually positively correlated between the high and low competition treatments, suggesting there were no tradeoffs between performance under high and low competition for these clones.     

Kin recognition and cannibalism in a subsocial spider

Evolution of cooperation and group living in spiders from subsocial family groups may be constrained by their cannibalistic nature. A tendency to avoid cannibalizing kin may facilitate tolerance among spiders and implies the ability to identify relatives. We investigated whether the subsocial spider Stegodyphus lineatus discriminates kin by recording cannibalism among juveniles in experiments during which amount of food and size difference among spiders in groups were varied. We hypothesized that family groups should be less cannibalistic than groups of mixed‐parental origin. Further, we tested whether food‐stress would influence cannibalism rates differently in kin and nonkin groups and the effect of relatedness on cannibalism within groups of spiders of variable size compared with those of homogenous size. In groups of six spiders, more spiders were cannibalized in nonsib groups than in sib groups under low food conditions. A tendency for nonkin biased cannibalism in starved spider pairs supported that kin recognition in S. lineatus is expressed when food is limited. Size variance of individuals within well‐fed groups of siblings and unrelated spiders had no influence on cannibalism rates. Apparently, both hunger and high density are important promoters of cannibalism. In addition to inclusive fitness benefits, we suggest that an ability to avoid cannibalizing kin will favour the evolution of cooperation and group living in phylogenetically pre‐adapted solitary species.     

Prey preferences ofPortia fimbriata,an araneophagic,web-building jumping spider (Araneae: Salticidae) from Queensland

Portia fimbriata from Queensland, a previously studied jumping spider (Salticidae), routinely includes web-building spiders and cursorial salticids in its diet, both of these types of prey being dangerous and unusual prey for a salticid. The present paper is the first detailed study ofP. fimbriata's prey preferences. Three basic types of tests of prey preference were used, providing evidence that (1)P. fimbriata males and females prefer spiders (both web-building spiders in webs and salticids away from webs) to insects; (2)P. fimbriata males and females prefer salticids to web-building spiders; (3)P. fimbriata males and females prefer larger spiders to smaller spiders; (4) there are intersexual differences in the preferences ofP. fimbriata for prey size, females preferring larger prey and males preferring smaller prey; and (5)P. fimbriata's prey preferences are not affected by a prior period without food of 2 weeks. When preferences were tested for by using both living, active prey and dead, motionless lures, the same preferences were expressed, indicating thatP. fimbriata can distinguish among different types of prey independent of the different movement patterns of different prey.     

Advantages of social foraging in crab spiders: Groups capture more and larger prey despite the absence of a web

Among group‐living spiders, subsocial representatives in the family of crab spiders (Thomisidae) are a special case, as they build protective communal leaf nests instead of extensive communal capture webs. It could thus be inferred that antipredator benefits (e.g., enhanced protection in larger nests) rather than foraging‐related advantages (e.g., capture of more and larger prey) promote sociality in this family. Nonetheless, subsocial crab spiders do share prey, and if this behaviour does not reflect mere food scramble but has a cooperative character, crab spiders may offer insights into the evolution of social foraging applicable to many other cooperative predators that hunt without traps. Here, we performed a comparative laboratory feeding experiment on three of the four subsocial crab spider species—Australomisidia ergandros, Australomisidia socialis and Xysticus bimaculatus—to determine if crab spiders derive advantages from foraging in groups. In particular, we tested artificially composed groups of five sibling spiderlings vs. single siblings in terms of prey capture success and prey size preference. Across species, groups had higher prey capture success (measured in terms of capture rates and capture latency) and were more likely to attack large, sharable prey—dynamics leading to reduced food competition among group members in favour of living and foraging in groups. Within groups, we further compared prey extraction efficiency among the three applied social foraging tactics: producing, scrounging and feeding alone. In A. ergandros, individuals were exceptionally efficient when using the non‐cooperative scrounger tactic, which entails feeding on the prey provided by others. Thus, our multispecies comparison confirms foraging advantages in maintaining a cooperative lifestyle for crab spiders, but also demonstrates the relevance of research into exploitation of cooperative foraging in this family.     

Effect of Group Size on Activity Budgets of Colobus vellerosus in Ghana

Group size influences foraging efficiency in several primates. We examined the activity budgets of 3 groups of Geoffroy's pied colobus (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana to determine whether larger group size induces scramble competition. We studied 2 groups (B1; N = 7-8 and WW; N = 31-33) occupying slightly overlapping home ranges from August to November 2000. We observed the third group, B2 (N = 15-16), comprising B1 and 7 male invaders in the same home range as B1 from August to November 2001. By comparing groups belonging to the same population and occupying sligthly overlapping or similar home ranges, we were able to control, to a certain extent, for differences in food distribution. We recorded a total of 3353 scans, yielding 14,886 activity records, over 73 days. As with other black-and-white colobus, resting was their most common activity (59%). Intergroup comparisons suggest that time spent feeding, resting and moving did not vary in relation to group size. However, intragroup comparisons between the sexes show that females in the large group spent more time feeding than males did, whereas this was not the case in the small group, which suggests that scramble competition may be occurring among female Colobus vellerosus at BFMS. It is also possible that this may be due to greater nutritional requirements because of a higher proportion of infants in the large group. In fact, the proportion is quite similar between the two groups, lending support to the idea that females in the two groups had comparable nutritional demands due to lactation. This suggests that increased feeding in females in the large group was partly an effect of scramble competition. Group size and group composition also influenced the frequency of social behavior. There was more grooming in the large group, and it was performed mostly by females. The distribution of activities throughout the day was similar to the pattern reported for other black-and-white colobus.     

Male-male relationships among wild bonobos (Pan paniscus) at Wamba,Republic of Zaire

Male-male relationships among wild bonobos (Pan paniscus) in two adjacent unitgroups (E1 and E2 groups), which were formed by division of the E group, were studied at Wamba, in the Central Zaire Basin, by analyzing the proximity and social interactions among males. Dominant-subordinate relationships between a male-male dyad were easily recognized from the directions of individual agonistic interactions. Male bonobos rarely joined forces in aggression. Clear differences in social status existed between adult and adolescent male bonobos in both groups, as reported in the case of chimpanzees (Pan troglodytes). The presence of mothers in the unit-group greatly influenced the dominant-subordinate relationships among males through strong mother-son bonds in both groups. However, the extent of the mother-son bonds differed between the groups. Males in the E2 group participated more frequently in agonistic or affinitive interactions than did males in the E1 group. Males in the E1 group were divided spatially into several clusters, while there were cohesive relationships among the adult males in the E2 group. The difference in intensities of mother-son bonds between the groups may be explained by the distribution of males at the time of the division of the E group. Differences in male-male relationships between bonobos and chimpanzees seem to be related to differences in intra- and inter-unit-group competition among males between the two species. Male chimpanzees may achieve coexistence by manipulating ambivalent relationships that are caused by intra- and inter-unit-group competition among them, while male bonobos may achieve coexistence by decreasing intra- and inter-unit-group competition among them.     

Effects of within‐colony competition on body size asymmetries and reproductive skew in a social spider

Reproductive partitioning is a key component of social organization in groups of cooperative organisms. In colonies of permanently social spiders of the genus Stegodyphus less than half of the females reproduce, while all females, including nonreproducers, perform suicidal allo‐maternal care. Some theoretical models suggest that reproductive skew is a result of contest competition within colonies, leading to size hierarchies where only the largest females become reproducers. We investigated the effect of competition on within‐group body size variation over six months in S. dumicola, by manipulating food level and colony size. We found no evidence that competition leads to increased size asymmetry within colonies, suggesting that contest competition may not be the proximate explanation for reproductive skew. Within‐colony body size variation was high already in the juvenile stage, and did not increase over the course of the experiment, suggesting that body size variation is shaped at an early stage. This might facilitate task specialization within colonies and ensure colony‐level reproductive output by early allocation of reproductive roles. We suggest that reproductive skew in social spiders may be an adaptation to sociality selected through inclusive fitness benefits of allo‐maternal care as well as colony‐level benefits maximizing colony survival and production.     

Colony size and parasitoid load in two species of colonial Metepeira spiders from Mexico (Araneae: Araneidae)

Summary While a number of advantages may result from group living, it may also lead to increased levels of attack by parasites because groups may be easier to find. This leads to the prediction that levels of parasitism should increase with colony size. We test this prediction by comparing colony size and parasitoid load for two species of colonial orb-weaving spiders from Mexico, Metepeira (undesc. sp., tentatively named atascadero) and Metepeira incrassata, which exhibit contrasting levels of social organization and utilize different habitats. For M. atascadero, which occurs solitarily or in small groups in desert/mesquite grassland habitat, rates of egg-sac parasitism fluctuate widely from year to year, and are closely tied to spider egg output. There is no relationship between colony size and rate of parasitism. For colonial M. incrassata, which occur in tropical rain forest/agricultural habitat, rates of parasitism are relatively constant from year to year. However, there is a positive relationship between colony size and rate of parasitism in this species. These differences are discussed with regard to the stability of the two habitats, prey availability, and the foraging behavior of the respective parasitoids.     

Feeding Competition and Group Size in Alouatta pigra

Researchers consider group size in primates to be determined by complex relationships among numerous ecological forces. Antipredator benefits and better resource defense are the primary pressures for large groups. Conversely, intragroup limited food availability, can result in greater intragroup feeding competition and individual energy expenditure in larger groups, creating energetic advantages for individuals in small groups and placing an upper limit group size. However, the extent to which food availability constrains group size remains unclear for many species, including black howlers (Alouatta pigra), which ubiquitously live in small social groups (≤10 individuals). We studied the relationship between group size and 2 key indices of feeding competition—day journey length and activity budgets—in 3 groups of wild Alouatta pigra at a hurricane-damaged site in Belize, Central America. We controlled for differences in food availability between home ranges (food tree density) and compared both indicators of feeding competition directly with temporal variation in food availability for each group. Our results show no consistent association between resource availability, group size, and either index of competition, indicating that feeding competition does not limit group size at the site—i.e., that larger groups can form without increased costs of feeding competition. The results support the search for other explanations, possibly social ones, for small group size in the primates, and we conclude with suggestions and evidence for such alternative explanations.     

Spacing in Colonial Spiders: Effects of Environment and Experience

Colonial orb-weaving spiders from Mexico (Metepeira spp.), show considerable geographic variation and temporal flexibility in group size and social spacing. A series of laboratory studies was conducted to test whether the variation observed in the field is the result of behavioral plasticity, or the result of genetic mechanisms inherent in different populations. Spiders from source populations in desert and moist tropical habitats were collected as eggs and raised in the laboratory under identical controlled conditions. To test for the effect of experience on conspecific tolerance, experiments were conducted rearing tropical and desert spiderlings in isolation and in communal groups. Measurements of spacing in laboratory colonies and observations of web building behavior show significant differences between populations, suggesting genetic differences between them. Desert spiders show an initial effect of isolation on tolerance of conspecifics — greater inter-individual distance that is eventually modified by communal adult experience. Tropical spiders put together after isolation show spacing patterns opposite those observed for desert spiders, and nearest neighbor distances similar to those seen in the communally reared (tropical) groups. These findings support the hypothesis that these populations represent separate species that differ with respect to their level of sociality.     

Relation of Intergroup Variation in Allogrooming to Group Social Structure and Ectoparasite Loads in Red Howlers (Alouatta seniculus)

We compare the allogrooming behavior of 5 troops (average size = 8.2) of red howlers (Alouatta seniculus) from the Venezuelan Llanos with that of other A. seniculus and Alouatta spp. of the genus. In 126.9 observation hr, we recorded 118 allogrooming events, with an average bout length of 109 sec. Females groomed more frequently than males did, but as groomees there is no significant differences between sexes. Adult males groomed adult females mostly in a sexual context (before copulation). Allogrooming rates differ significantly among groups. There are also significant differences among members of the same troop both as groomers and groomees, which we explain in the context of the social behavior and history of each troop. There is no significant correlation between weight of the groomee and duration of the grooming bout. However, the examination of grooming rates and ectoparasite load suggests that allogrooming may have hygienic consequences. Differences in allogrooming rates among species of Alouatta are related to differences in group kin structure and patterns of female competition, in particular, coalition formation. We conclude that the social structure and the degree of relatedness among individuals within a group (or among individuals in a population in interpopulation comparisons) is a more important determinant of allogrooming rate than body size or group size. Our results emphasize the importance of considering intergroup and interpopulational variation in behavior.     

Natal Dispersal Patterns of a Subsocial Spider Anelosimus cf. jucundus (Theridiidae)

Species that alternate periods of solitary and social living may provide clues to the conditions that favor sociality. Social spiders probably originated from subsocial‐like ancestors, species in which siblings remain together for part of their life cycle but disperse prior to mating. Exploring the factors that lead to dispersal in subsocial species, but allow the development of large multigenerational colonies in social species, may provide insight into this transition. We studied the natal dispersal patterns of a subsocial spider, Anelosimus cf. jucundus, in Southeastern Arizona. In this population, spiders disperse from their natal nests in their penultimate and antepenultimate instars over a 3‐mo period. We tracked the natal dispersal of marked spiders at sites with clustered vs. isolated nests. We found that most spiders initially dispersed less than 5 m from their natal nests. Males and females, and spiders in patches with different densities of nests, dispersed similar distances. The fact that both sexes in a group dispersed, the lack of a sex difference in dispersal distance, and the relatively short distances dispersed are consistent with the hypothesis that natal dispersal results from resource competition within the natal nest, rather than inbreeding avoidance in competition for mates. Additionally, an increase in the average distance dispersed with time and with the number of spiders leaving a nest suggests that competition for nest sites in the vicinity of the natal nest may affect dispersal distances. The similar distances dispersed in patches with isolated vs. clustered nests, in contrast, suggest that competition among dispersers from different nests may not affect dispersal distances.     

Intraspecific variation in group size in the blackbuck antelope: the roles of habitat structure and forage at different spatial scales

The main ecological factors that are hypothesized to explain the striking variation in the size of social groups among large herbivores are habitat structure, predation, and forage abundance and distribution; however, their relative roles in wild populations are not well understood. I combined analyses of ecological correlates of spatial variation in group size with analyses of individual behaviour in groups of different sizes to investigate factors maintaining variation in group size in an Indian antelope, the blackbuck Antilope cervicapra. I measured group size, habitat structure, forage, and the occurrence of predators in ten blackbuck populations, and, at a smaller spatial scale, within an intensively studied population. To examine the processes by which these ecological factors influence group size, I used behavioural observations and an experiment to estimate the shape of the relationship between group size and potential costs and benefits to individuals. Group size varied extensively both among and within populations. Analyses of spatial variation in group size suggested that both forage and habitat structure influence group size: large-scale, among-population variation in group size was primarily related to habitat structure, while small-scale, within-population variation was most closely related to forage abundance. Analyses of individual behaviour suggested that larger groups incur greater travel costs while foraging. However, individuals in larger groups appeared to experience greater benefits, namely the earlier detection of a “predator”, a reduction in vigilance, and an increase in the time spent feeding. Overall, these findings suggest that individuals in groups experience a trade-off between predation-related benefits and costs arising from feeding competition. Habitat structure and forage likely influence the nature of this trade-off; thus, variation in these ecological factors may maintain variation in group size. The role of predation pressure and other factors in explaining the remaining variation needs further exploration. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

近亲幽灵蛛用于兼性群居生活的时间消耗

所谓近亲幽灵蛛（Pholcus affinis）的兼性群居生活,是指他们可以单独生活,也可以几个个体共栖于一张网上成为一组。本文使用焦点动物追踪法,研究不同大小的组（指组内个体数量多少）、不同年龄阶段的蜘蛛在组内生活中用于“消耗性”行为（移动、织网、弹跳、拨丝或种内相互作用）的时间耗费。主要结论包括：蜘蛛的年龄和组的大小影响组内成员的付出。在不同大小的组里,2～3龄幼蛛（S型）、亚成体或成体（L型）用在“消耗性”行为上所用的时间百分率没有显著性差异,但4～5龄幼蛛（M型）随着组内成员的增多而要花费更多的时间。当组内个体数量超过3个,L型和M型蜘蛛之间有着更多的相互作用,而S型蜘蛛不论组内个体多少,其个体之间相互作用都很少。研究结果同时显示,近亲幽灵蛛在不同发育阶段有不同的群居生活策略。     

Competition for a limited space in kleptoparasitic Argyrodes spiders revealed by field experiments

Most kleptoparasitic Argyrodes spiders rely exclusively on host spider webs for obtaining their food. Because their densities occasionally reach high levels within a restricted area, competitive interactions may be important for determining the density of these unique spiders. Here I used two Argyrodes species commonly found on webs of the large orb-web spider Nephila clavata to clarify whether inter- and intraspecific competition influences abundance and within-web distribution by using observational data and field experiment. Removing Argyrodes flavescens from the host webs induced a remarkably high immigration of that species while density on control webs remained almost at the same level, which is evidence for strong intraspecific competition. Larger individuals of A. flavescens were located more frequently at the capture area of the host webs where it is easy to access prey ignored by the host spider, and spiders immigrating into webs from which that species had been removed were smaller in body size, suggesting interference competition for space among conspecific kleptoparasites. Argyrodes bonadea increased in number on webs from which A. flavescens had been removed, and the increase was correlated with the number of A. flavescens removed. This finding is evidence for interspecific competition that is rarely reported in spiders. A multiple regression model including numbers of a conspecific parasite as well as web and body sizes of the host spider could not detect competitive interactions between species, suggesting the importance of experimental approaches. Received: May 22, 2000 / Accepted: December 1, 2000     

Foraging behavior of the communal spider,Philoponella republicana (Araneae: Uloboridae)

The communal orb-weaving spider, Philoponella republicana,was observed in the subtropical moist forest of Southeast Peru. These spiders live in colonies of conspecifics whose individual orbs are connected by silk. The wrapping of a prey prior to feeding is a large component of the prey capture process because P. republicanahas no venom with which to kill an insect. Wrapping time was the only aspect of prey capture that was strongly correlated with the size of the insect captured. Occasionally we observed several individuals working together to wrap a prey item. These joint efforts were more frequent on prey larger than the capturing spider. Although group captures accounted for only 5.5% of captures, they represented 14.7% of the biomass obtained. A comparison of the relationship between wrapping time and prey size for solitary and group efforts suggested that, by working together, the spiders reduced their total handling time. In most cases only one spider fed on the captured prey.     

Survival benefits select for group living in a social spider despite reproductive costs

The evolution of cooperation requires benefits of group living to exceed costs. Hence, some components of fitness are expected to increase with increasing group size, whereas others may decrease because of competition among group members. The social spiders provide an excellent system to investigate the costs and benefits of group living: they occur in groups of various sizes and individuals are relatively short-lived, therefore life history traits and Lifetime Reproductive Success (LRS) can be estimated as a function of group size. Sociality in spiders has originated repeatedly in phylogenetically distant families and appears to be accompanied by a transition to a system of continuous intra-colony mating and extreme inbreeding. The benefits of group living in such systems should therefore be substantial. We investigated the effect of group size on fitness components of reproduction and survival in the social spider Stegodyphus dumicola in two populations in Namibia. In both populations, the major benefit of group living was improved survival of colonies and late-instar juveniles with increasing colony size. By contrast, female fecundity, female body size and early juvenile survival decreased with increasing group size. Mean individual fitness, estimated as LRS and calculated from five components of reproduction and survival, was maximized for intermediate- to large-sized colonies. Group living in these spiders thus entails a net reproductive cost, presumably because of an increase in intra-colony competition with group size. This cost is traded off against survival benefits at the colony level, which appear to be the major factor favouring group living. In the field, many colonies occur at smaller size than expected from the fitness curve, suggesting ecological or life history constraints on colony persistence which results in a transient population of relatively small colonies.     

Age and Sex-Based Differences in the Use of Prey Sensory Cues in Wolf Spiders (Araneae: Lycosidae)

Differences in foraging patterns mediated by sensory cues were examined between adult and juvenile male and female wolf spiders (Schizocosa rovneri; Lycosidae). Patch residence time for thirty-one spiders were tested among juveniles and adults in artificial foraging patches. Patches varied in sensory information provided by live prey (crickets) as follows: visual stimuli alone; vibratory stimuli alone; visual and vibratory stimuli together; and control (no stimuli). Spiders moved between patches for one hour, but could not feed. Adult Schizocosa rovneri use primarily visual information to determine patch residence time, but juveniles use vibratory cues as well. Significant age and sex-based differences in the use of sensory cues suggest that observed divergent foraging strategies are partly due to the use of different perceptual cues in prey detection.     

Scramble Competition Among <Emphasis Type="Italic">Colobus vellerosus</Emphasis> at Boabeng-Fiema,Ghana

We investigated the occurrence of scramble competition among Colobus vellerosus at Boabeng-Fiema, Ghana. If scramble competition had an impact on feeding efficiency among females, we expected a positive relationship between group size and the proportion of time spent feeding, day journey length, or home range size assuming resource availability is similar among the groups compared. We collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000–August 2001) on 2 study groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also collected ranging data on group movements at half-hour intervals. The large group (WW1) had a significantly longer day journey length than the small group (B1), and females in the large group spent a significantly greater proportion of time feeding in the wet season, a period of low food availability, which suggests it may be a bottleneck period when food resources are scarce and Colobus vellerosus is close to being energy limited. The proximity data suggested females may be able to reduce or adjust for competition by having fewer neighbors when they feed and by spreading out when in a larger group. However, we found no relationship between home range size and group size or that females spent a greater proportion of time feeding than adult males did. Our results highlight the need to factor in differences in food availability when investigating scramble competition. Though equivocal, our results suggest scramble competition occurs among Colobus vellerosus, leading us to suggest there was a match with the potential competitive regime, i.e., food distribution.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.