Fighting without weaponry: a review of male-male contest competition in butterflies

The males of many butterfly species compete via pair-wise intrasexual contests, yet lack any obvious morphological traits conventionally associated with animal aggression. The evolution of contest behavior in this group is therefore interesting because the means by which individuals are able to inflict costs upon each other during combat are unclear. Here we review the relevant empirical information within a game-theoretical context in order to analyze contest-related costs and mechanisms of dispute settlement. Territorial butterfly contests carry obvious role asymmetries, and individuals in the 'resident' role are more likely to win across all studied species. We use this phenomenon as a framework for gaining further insight into the evolution of contest settlement in this group. Four principal hypotheses are evaluated: (1) that residency is used as an arbitrary convention, (2) that residents stand to gain a greater payoff if successful, (3) that residency bestows higher resource-holding potential (RHP) on males in that role, and (4) that residents are males of intrinsically higher RHP. We found most support for the latter explanation (the 'superior-competitor' hypothesis), partly because the alternatives require a restrictive set of ecological conditions, but also due to positive support for the idea of intrinsic RHP. The strongest evidence is provided by species in which repeated interactions (between the same individuals) always have the same outcome, regardless of changes in residency status. However, we also found a consistent effect on contest duration due to apparent 'confusion over residency,' which suggests a conventional element to contest behavior. Although butterfly contests must be costly in some way, the literature provides limited insight into how costs accrue, and hence the determinants of contest RHP remain obscure. We suggest that careful experimentation, using 'natural' manipulations where possible, should prove most beneficial in identifying costs and exploring schedules of contest settlement in this group. Moreover, future investigations should be sensitive to the fact that butterfly territoriality and associated male-male interactions are seated within a broader ecological and life-historical context.     

Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition

The effect of aggressive competition over food resources on energy intake rate is analyzed for individuals of three groups of 25–35 white-faced capuchin monkeys, Cebus capucinus, living in and near Lomas Barbudal Biological Reserve, Costa Rica. An individuals energy intake rate on a given food species was affected by its rank and the number of agonistic interactions within the feeding tree. Dominant group members had higher energy intake rates relative to subordinate group members whether or not there was agonism within the feeding tree. Low- and mid-ranked individuals had lower energy intake rates in trees with higher amounts of aggression, while energy intake rate of high-ranked individuals was not affected by the amount of aggression in the feeding tree. Energy intake was not influenced by the sex of the individual when rank was held constant statistically. Energy intake was positively correlated with total crown energy (measured in kilojoules) within the feeding tree for two of three study groups. This difference may be explained by the quality of each groups territory. Finally, high-ranked individuals are responsible for the majority of agonism within feeding trees and target middle- and low-ranked individuals equally. These findings fit the predictions of current socioecological models for within-group contest competition over food resources. The results of this study suggest that within-group competition affects energy intake rate in white-faced capuchin monkeys.     

Proximate mechanisms of contest competition among female Bwindi mountain gorillas (Gorilla beringei beringei)

Socioecological models provide a framework for predicting how animals respond competitively to the abundance and distribution of food resources. Testing predictions of socioecological models requires analysis of relationships among food resource characteristics, competitive behaviors, and measures of rank-related skew in energy balance or reproductive success. A positive relationship between dominance rank and energy balance has been observed among female mountain gorillas in Bwindi Impenetrable National Park, Uganda. This study examines the proximate mechanisms underlying feeding competition among those females. To assess the contestability of food resources, we measured the time a female spent feeding at a food site (food site residence time). We also examined the relationship between dominance rank and the access to resources, as well as the rate, context, and direction of aggression, and the number of neighbors in close proximity. As predicted, females had longer food site residence times and higher aggression rates with fruit and decaying wood than with herbaceous vegetation, suggesting that those resources may be contestable. Aggression was predominantly directed down the dominance hierarchy, although against expectation, rank was not significantly correlated with aggression rates or the time spent feeding on contestable foods. Higher-ranking females had significantly fewer neighbors, suggesting that lower-ranking females avoid higher-ranking ones. This study provides additional support for the claim that there is variability in how primates respond to the quality and distribution of food resources and that avoidance as a strategy to cope with feeding competition may result in similar skew in energy balance as rank-related aggression.     

白洋淀沉积物中磷的存在形态及垂直变化规律研究

湖泊沉积物中磷的含量及其形态分布是影响湖泊营养化进程的极为重要因素,对研究湖泊营养化等环境问题具有重要意义。采用化学连续提取分析法,对白洋淀环淀中村水域不同地点的沉积物中磷的形态及其在垂直方向上的变化进行了研究。结果表明,研究区域内,环淀中村水域表层沉积物总磷（TP）质量分数在544.424～608.197mg·kg-1之间,无机磷（IP）是沉积物中磷的主要形态,有机磷（OP）质量分数较小,约占10%～27%,无机磷中钙磷（Ca-P）占的比例相对较大,在质量分数上约占75.8%～94.1%。从各形态磷质量分数的变化范围来看,Ca-P〉OP〉Fe/Al-P;人类活动对湖泊沉积物中磷的形态及其水平和垂向变化影响很大。     

The roles of competition and environmental heterogeneity in the maintenance of behavioral variation and covariation

Many models of selection predict that populations will lose variation in traits that affect fitness. Nonetheless, phenotypic variation is commonly observed in natural populations. We tested the influences of competition and spatial heterogeneity on behavioral variation within and among populations of Merriam's kangaroo rats (Dipodomys merriami) and tested for the differential expression of trait correlations. We found that populations of D. merriami exhibited more aggression at sites with more competition. Contrary to theoretical predictions and empirical results in other systems, the sites with the greatest spatial heterogeneity and highest levels of competition did not exhibit the most behavioral variation among individuals. However, the greatest within-individual behavioral variability in boldness (response to cues of predator presence) was exhibited where spatial heterogeneity was highest. Aggression and boldness of D. merriami were highly repeatable, that is, individuals behaved in a consistent manner over time, and the two behaviors were also highly correlated. Interestingly, the strength of this correlation was greatest where the competitive community was least diverse. These findings add to increasing evidence that natural populations of animals exhibit patterns of behavioral covariance, or personality structure, and suggest that competitive variation may act to erode personality structure.     

天目山自然保护区柳杉种群种内和种间竞争

调查了天目山自然保护区老殿、大树王和禅源寺3个柳杉典型样地,以研究竞争在柳杉种群衰退中的作用。结果表明,柳杉基本上是柳杉群落的单一优势种,相对优势度占58.79%～93.34%;单位面积内柳杉胸高截面积达59.96m2·hm-2,材积达904.9m3·hm-2。柳杉群落存在较严重的竞争,单木竞争指数以老殿最大,平均达4.35;大树王最小,平均为0.58。以竞争指数测度的竞争强度表明,种内竞争大于种间竞争。柳杉单株竞争强度与胸径成反比。认为种内竞争是导致柳杉种群衰退的重要原因之一。     

Between-group contest competition for food in a highly folivorous population of black and white colobus monkeys (Colobus guereza)

In theory, between-group contest (BGC) competition for food can greatly influence female social relationships and reproductive success in primates, but few studies have investigated whether such effects occur and, if so, under what ecological conditions. There is evidence that adult male black and white colobus monkeys (Colobus guereza, “guerezas”) defend the food their mates need against other groups, suggesting that BGC competition is important in this species. Using data on feeding, ranging, vegetation patterns, and intergroup encounters between six neighboring guereza groups, I provide evidence that the highly folivorous guerezas at Kanyawara, Kibale National Park, Uganda, engaged in BGC competition over unevenly dispersed, relatively high-quality feeding areas or “core areas”. Intergroup aggression was common, and groups’ home ranges overlapped. Groups were more likely to initiate high-level aggression if they encountered another group within or near their core area, and groups that initiated and won encounters often fed in the same areas in which losing groups had fed. Guerezas fed selectively on species with contagious (clumped) distributions and concentrated their feeding efforts in areas of the forest that contained the most food (core areas). Groups could be ranked in a linear dominance hierarchy, and group rank number was inversely related to the quantity and quality of food in groups’ core areas. This study not only provides good evidence that BGC competition occurs in primates but it also reinforces the idea that folivore food resources may be worth defending.     

Male choice and competition in Tetraopes tetraophthalmus: effects of local sex ratio variation

Summary Male milkweed beetles (Tetraopes tetraophthalmus) are shown to exhibit choice between potential mates. Males, however, also exhibit direct competition for access to potential mates. These differences in male behavior are dependent on the local sex ratio, with males becoming increasingly choosy when the sex ratio is female-biased, and more competitive when the sex ratio is malebiased. Females show neither choice nor competition. In both the laboratory and in the field, body size is important in determining patterns of competition and choice. Larger males win most aggressive encounters, and males generally prefer large females as mates. The combination in a single sex, of competition for mates and mate choice is not predicted by classical sexual selection theory. Male choice and competition may have evolved in T. tetraophthalmus due to the combination of (1) the mating system (2) the pattern of variation in female fecundity, and (3) the spatially and temporally variable operational sex ratios. This explanation is not dependent on high heritability of body size.     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

Interannual variation of ichthyoplankton density and species composition in the waters off northeastern Taiwan

The ichthyoplankton in the waters of northeastern Taiwan have been studied since 1989. Based on collections in August 1989 to 1991, this report treats primarily the annual variation of ichthyoplankton density, species composition and assemblage diversity among the waters of the East China Sea, the Kuroshio and the mixing zone. The ranges of average densities found for fish eggs were 7 to 3897 ind 1000 m^-3 and 0.02 to 1.43 g 1000 m^-3, for ichthyoplankton 154 to 1364 ind 1000 m^-3 and 0.33 to 3.23 g 1000 m^-3, and for zooplankton 52 to 532 g 1000 m^-3. An interannual comparison indicates a tendency towards a decrease in ichthyoplankton density from 1989 to 1991. In total, 5803 individuals assigned to at least 91 families and 254 taxa at species level were collected. Lantern fish form the most common species group (Benthosema pterotum: 12.1%; Diaphus pacificus: 4.2%). The other species in order of frequency are: gobies (5.8%); Callionymus sp. (3.4%); Auxis spp. (3.1%); Trachinocephalus myops (3.0%); Nibea sp. (2.8%); Synagrops philippinensis (2.3%); Priacanthus macracanthus (2.2%); and percophid (2.0%). The relative frequency of the ten most abundant taxa fluctuated annually. Neither interannual distinction nor interannual correlation was confirmed by statistical analysis, but intra-annual rank correlations were established among the water masses. A weak tendency towards increasing diversity from 1989 to 1990 and little diversity in the mixing zone were observed. We conclude that there is a complex inter-dependence of assemblages from the waters of northeastern Taiwan.     

Partitioning the factors of spatial variation in regeneration density of shade-tolerant tree species

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.     

Planktonic bacteria in the Humber Estuary; Seasonal variation in population density and heterotrophic activity

Seasonal variation in population density and heterotrophic activity (as maximum potential rate, V _max, for glucose mineralization) of bacteria attached to suspended solids, of free bacteria and of total bacteria was examined in the water of the Humber Estuary, north-east England. At the principal site, which was studied over 2 yr, values of density and activity of attached and total bacteria showed marked seasonal periodicity, being low in summer and high from autumn to spring. This was a consequence of density of attached bacteria being dependent on the concentration of suspended solids, which was high in winter and low during summer. Values of density and activity of free bacteria, which were less than those of attached bacteria, showed no seasonal pattern but fluctuated irregularly and were independent of concentration of suspended solids. At two further sites, which were studied over 1 yr only, the density of bacteria varied seasonally in a similar manner, but the activity of attached and total bacteria showed no distinct seasonal pattern — this may, however, be due to limitations of the data rather than to fundamental differences between the sites studied.     

Unraveling the complexities of variation in female mate preference for vertical bars in the swordtail, Xiphophorus cortezi

Investigations into the nature of mate choice suggest that variation in female mate preferences is often context dependent, varying in response to genetic and environmental influences on female condition as well as to external environmental stimuli. Determining whether variation in female mate preference is adaptive requires understanding the variables involved that produce this variation and how they interact. Comparative, multivariate studies of wild-caught adult females can be used as initial assessments of variation in female mate preferences, providing valuable insights into the parameters that influence female preferences under natural conditions. We examined variation in female preferences for the pigment pattern vertical bars across five populations of the swordtail, Xiphophorus cortezi. The populations we examined are genetically differentiated and varied in the frequency of males and females with bars. We also considered a variable indicative of within-individual variation (size, as influenced by age) and a variable that varies across individuals (genotype for vertical bars: barred or barless). Using Akaike information criterion, all candidate models explaining variation in strength of preference included female bar state, female size, and population. We suggest that a combination of genetic (bar state) and environmental (female size) conditions influenced how they responded to experience with both male phenotypes in X. cortezi. Future studies should examine the possibility that barred and barless females respond differently over an environmental gradient.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Constraints on group size in primates and carnivores: population density and day-range as assays of exploitation competition

Summary We hypothesise that foraging group size (FGS) and population group size (PGS) in primates and carnivores are related to quantifiable variables indexing the intensity of exploitative competition. Group size is predicted to increase with both food density and travel capabilities, as estimated by the constraint-free day-range (DR_s), i.e. the day-range of a solitary individual uninfluenced by competition from conspecifics. We test this exploitation competition hypothesis among primates and carnivores, using data on populations, species and genera. Food density was indexed by population density. Where DR_s could be estimated by regression it was found to be correlated with observed day-range (DR_m). DR_s was therefore indexed by DR_m in all species. Population density and DR_m were associated with each other, but in a multiple regression each variable contributed independently to explaining variation in group size. PGS was predicted better than FGS, but regressions involving either measure of group size were significant in all analyses. The multiple regression analyses were validated by the method of linear contrasts, which accounts for possible lack of independence among taxa. We conclude that species differences in group size are influenced by both food density and DR_s. This suggests that variation in the intensity of exploitation competition is partly responsible for differences in group size.Correspondence to: R.W. Wrangham     

Short-term effect of temperature variation on the competition between PAOs and GAOs during acclimation period of an EBPR system

Sequencing batch reactor (SBR) for enhanced biological phosphorus removal (EBPR) processes was used to investigate the impact of the temperature shock on the competition between phosphorus-accumulating organisms (PAOs) and glycogen accumulating organisms (GAOs) in start-up stage. During the 34 days operation, SBR was set with temperature variation(0–5 d, 22±1°C; 6–13 d, 29±1°C; 14–34 d, 14±1°C). PAOs and GAOs were analyzed by fluorescent in situ hybridization (FISH), and intracellular polyphosphate granules were stained by Neisser-stain. The results showed that the influence of temperature shock on PAOs’ abundance was more serious than that on GAOs in the enriching process. Under sudden and substantially temperature variation, from 22±1°C to 29±1°C and then to 14±1°C, the domination of PAOs was deteriorated. After temperature shock, PAOs’ competitive advantages at low temperature that concluded in other study did not appear in our study. As mesophilic, GAOs (indicated by Alphaproteobacteria and Gammaproteobacteria) were more temperature adaptive and better grew and took the domination at 14±1°C in the end. In the competition process, organisms of tetrad forming organisms (TFOs)-like shape which were considered as typical GAOs, were observed. With the evidence of poly-P granules containing by Neisser-straining and result of FISH, these organisms of TFOs-like shape were better to be assumed as adaption state or a special self-protecting shape of PAOs.     

Synergistic influences of phase, density, and climatic variation on the dynamics of fluctuating populations

Although ecologists have long recognized that certain mammalian species exhibit high-amplitude, often multiannual, fluctuations in abundance, their causes have remained poorly understood and the subject of intense debate. A key contention has been the relative role of density-dependent and density-independent processes in governing population dynamics. We applied capture-mark-recapture analysis to 25 years of monthly trapping data from a fluctuating prairie vole Microtus ochrogaster population in Illinois, USA, to estimate realized population growth rates and associated vital rates (survival and recruitment) and modeled them as a function of vole density and density-independent climatic variation. We also tested for phase dependence and seasonality in the effects of the above processes. Variation in the realized population growth rate was best explained by phase-specific changes in vole density lagged by one month and mean monthly temperatures with no time lags. The underlying vital rates, survival and recruitment, were influenced by the additive and interactive effects of phase, vole density, and mean monthly temperatures. Our results are consistent with the observation that large-scale population fluctuations are characterized by phase-specific changes in demographic and physiological characteristics. Our findings also support the growing realization that the interaction between climatic variables and density-dependent factors may be a widespread phenomenon, and they suggest that the direction and magnitude of such interactive effects may be phase specific. We conclude that density-dependent and density-independent climatic variables work in tandem during each phase of density fluctuations to drive the dynamics of fluctuating populations.