Spatial clumping of food increases its monopolization and defense by convict cichlids, Cichlasoma nigrofasciatum

The hypothesis that resource monopolization and defense increaseas the spatial clumping of resources increases was tested usinggroups of three convict cichlids competing for 120 Daphnia magnaprey. Spatial clumping was manipulated by varying the distance(3, 20, or 40 cm) between three tubes through which the preyappeared. As predicted, monopolization of prey (percentage eatenby the dominant fish) and frequency of aggression (chases perminute) by dominant fish increased significantly as the distancebetween the tubes decreased. However, there was no evidenceof individual flexibility in the aggressiveness (percentageof conspecifics chased) of dominant fish across treatments.Differences among dominant fish in aggressiveness were positivelycorrelated with their ability to monopolize prey, but the strengthof the correlation decreased as the distance between the tubesincreased. Aggression appears to be a more effective mechanismof interference competition when resources are clumped thanwhen resources are dispersed.     

Dispersal polymorphism and physiological condition of males and females in the ant, Formica truncorum

This study deals with dispersal behavior of sexuals and intraspecificvariation in queen numbers. The specific questions are: (1)Is there an association between male and female dispersal behaviorand the number of queens in a colony? (2) Is there an associationbetween individual behavior and physiological condition? (3)Do males and females from monogyne (one queen per colony) andpolygyne (several functional queens per colony) colonies differwith respect to size, weight, and physiological condition? Theresults show that both males and females are more prone to dispersein monogyne than in polygyne colonies. Moreover, males and femalesof both monogyne and polygyne colonies show dispersal polymorphism,suggesting that an increased tendency of reproductive femalesto stay in the maternal colony may cause monogyne colonies toswitch to polygyny. The propensity to disperse is associatedwith the physiological condition of individuals. Larger andheavier females containing more fat and glycogen preferentiallydisperse, whereas smaller ones with less fat and glycogen moreeasily dealate and mate without a previous nuptial flight. Maledispersal correlates positively to larger size and higher levelsof glycogen; fat contents do not increase during maturation.The females produced in monogyne colonies are larger, heavier,and contain more fat and glycogen than those produced in polygynecolonies. The males produced in monogyne colonies have relativelylonger wings and are heavier than those produced in polygynecolonies. However, there are no differences in size and fatcontents between males from monogyne and polygyne colonies.     

Female mate copying in the guppy {Poecilia reticulata): age-dependent effects

Virtually all studies of mate choice to date have assumed thatfemales choose mates independent of one another. Social cues,however, such as the mate choice of conspecifics, may also playan important role in such decisions. Previous work has shownthat female guppies of similar age copy each other's choiceof mates. Here we examine the effect of relative age on matechoice copying in the guppy, Poecilia reticulata, and examinewhether younger individuals are more likely to copy the matechoice of older conspecifics than vice versa. Results indicatethat younger females copy the mate choice of older females,but older individuals do not appear to be influenced by themate choice of younger individuals.     

The influence of nest predation on mating strategies under polygyny

The mating status of an advertising male is one important factorfor female assessment. In species where already-mated malestry to attract additional females at some distance from theirmate, mating status may be difficult to assess. Limitationsin the time available to breed and a decrease in reproductivesuccess over the breeding season could make the costs outweighthe benefits of a time-consuming mate assessment process, whichautomatically postpones the onset of breeding. Time and energyconstraints will then limit the behavioral options for femalesin search of breeding opportunities, and females would suffera cost when assessing male mating status. Female choice maythen be better understood by analyzing the behavioral optionsfor females; the success of a strategy in which costly assessmentsare avoided and females choose mates indiscriminately, disregardingmating status ("fast" strategy), could be compared to the successof a strategy in which females carefully try to assess matingstatus ("coy" strategy). Nest predation should be included insuch an analysis because it influences the amount of male assistancefemales of polygynous males receive, as nest predation for onlyone of a polygynous male's two females leads to more male assistancefor the other female. In this study, I compare the outcome ofa "fast" strategy with that of a "coy" strategy under differentrates of nest predation. Although high probabilities of becominga mate of a polygynous male associated with a fitness reductiondecrease the relative success of a "fast" strategy, simulationsshow that high rates of nest predation considerably lower thedifference in the outcome of the two strategies. Therefore,the cost of assessing mating status must be relatively smallto be economical.     

Reproductive success of polygynous male corn buntings (Miliaria calandra) as confirmed by DNA fingerprinting

Despite a sex ratio approximating to unity, female corn buntingswere not equally distributed among males. In 1989 and 1990,41.2% of 50 males were monogamously paired, 29.4% were polygynous,and 23.5% were unpaired. Polygynous males usually paired withtwo females, although in 1990 three males were trigamous. Polygynousmales fledged more offspring from their territories than didmonogamous males, mainly because they had more mates. The fledgingsuccess per nesting female was slightly higher in territoriesof polygynous males, but not significantly so. DNA fingerprintingwas used to confirm the true paternity of 44 offspring from15 broods and the true maternity of 50 offspring from 16 broods.A further 12 offspring from three broods for which neither putativeparent was available were also fingerprinted. Actual reproductivesuccess of parents was close to that inferred from observationsof number of young raised. There was only one brood, containingtwo chicks (4.5% of offspring, or in 6.7% of broods), wherethe chicks were not fathered by the male defending the territory.However, this nest was close to the territory boundary, andthe defending male may have been assigned incorrectly. Therewere no cases of intraspecific brood parasitism (n = 16 broods).The copulation rate was low, and extrapair copulation attemptswere rare, probably because of the poor chances of sneakingonto a neighbor's territory undetected and the costs of leavinga territory unguarded.     

Area-restricted search by the plains pocket gopher (Geomys bursarius) in tallgrass prairie habitat

Because pocket gophers have the high energetic cost of excavatingburrows and an inability to detect distant food items throughthe soil, I hypothesized that individuals within establishedburrow systems would use area-restricted search as a foragingstrategy. To examine this hypothesis I compared gopher foragingeffort over a 10-month period between areas in which overallplant densities were experimentally varied. Gophers expendedapproximately 50% of their foraging effort in areas with thehighest plant density, even though these made up only 33% ofthe available area in experimental plots. In large, griddedareas sampled for an entire season as well as in small areasin which gophers foraged for less than 1 week, gopher foragingeffort was related to the density of a single leguminous plantspecies, Psoralea argophylla. In small plots where this plantspecies was at high density, gophers created more tunnel branches,thereby intensifying their search effort. Thus, area-restrictedsearch appears to increase the rate of encounter with the patchilydistributed Psoralea plants.     

Failure of simple optimal foraging models to predict residence time when patch quality is uncertain

Blue jays (Cyanocitta cristata) were presented with a foragingsituation in which half of the patches they encountered containedno prey and half contained a single prey item. Experimentallydetermined probability distributions controlled prey arrivaltimes in those patches that contained prey. Patch residencein empty patches was studied during four experiments. In thefirst, prey arrival was exponentially distributed. Residencetimes increased with travel time as predicted by a rate-maximizationmodel, but the bird stayed in empty patches much longer thanpredicted. During the second experiment, prey arrival was uniformlydistributed. The jays again stayed longer than optimal, andpatch residence times increased as travel time increased, althoughthe residence time that maximized rate of intake was independentof travel time under these conditions. In the third experiment,exponential and uniform patches were randomly intermixed. Thejays showed larger travel-time effects in the exponential thanin the uniform patch. However, the travel-time effect in theuniform patch was contrary to rate-maximization predictions,and the birds again overstayed in both patch types. In the fourthexperiment, prefeeding at the start of each foraging bout slightlyincreased overstaying rather than decreasing overstaying, aswould be expected if overstaying were due to underestimatingenvironmental quality. Consistent and dramatic overstaying anda travel-time effect under conditions where travel time hasno effect on optimal residence times suggest that the rate-maximizationapproach does not apply to foraging problems involving patchuncertainty.     

The definition of eusociality

We describe more precise definitions for the term "eusociality"and other social systems. Our criterion for eusociality is thepresence of castes, which are groups of individuals that becomeirreversibly behaviorally distinct at some point prior to reproductivematurity. Eusocial societies are characterized by two traits:(1) helping by individuals of the less-reproductive caste, and(2) either behavioral totipotency of only the more reproductivecaste (facultative eusociality) or totipotency of neither caste(obligate eusociality). We define "cooperative breeding" asalloparental care without castes. Cooperatively breeding societiesmay comprise two types, semisocial (distribution of lifetimereproductive success bimodal), and quasisocial (distributionof lifetime reproductive success unimodal), but this hypothesisrequires empirical analysis. Our definitions conceptually unifystudies of arthropod and vertebrate sociality.     

Visual information, resource value, and sequential assessment in convict cichlid (Cichlasoma nigrofasciatum) contests

We staged contests between convict cichlids (Cichlasoma nigrofasciatum)that were matched for size and gender to test the influenceof prior information and resource value on the duration andstructure of fights. The contestants were separated before thecontest by either clear or opaque dividers to allow or preventvisual assessment, respectively. Contests were shorter in the"clear" than in the "opaque" treatment, suggesting that visualassessment occurred. The duration of lateral display, a noncontactdisplay, was shorter in the clear than in the opaque treatment,but the treatments did not differ significantly in the durationof three contact displays (biting, mouth wrestling, and circling).These results are consistent with the hypothesis that lateraldisplay provides primarily visual information, probably aboutbody size, whereas the other behavior patterns provide primarilynonvisual information, probably about strength. Second contestsbetween the same pair of fish were shorter than first contests,suggesting that the information acquired during the first contestmade it easier to resolve the second. After the subordinatefish from the second contest was given access to a mate, itfought more persistently so that third contests were longerthan second contests. Our results support the predictions ofthe sequential assessment model.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.     

Divergence in male mating tactics between two populations of the soapberry bug: II. Genetic change and the evolution of a plastic reaction norm in a variable social environment

Many social behaviors are conditional, but behavioral comparisonsbetween populations do not normally distinguish genetic andenvironmental causation. As a result, the opportunity to testpredictions about the evolution of strategic conditionality(genotype x environment interaction) is lost. We apply theseconcepts in an examination of how interpopulation differencesin mean and variance of sex ratio have led to genetic differencesin the allocation of male effort to mate guarding versus nonguardingbetween genetically isolated populations of the soapberry bugin Oklahoma and Florida. We observed the mating behavior ofmales from the two populations at a series of experimental sexratios, and modeled their mating decisions as first-order Markovchains of independent mating states. Likelihood ratio testsof these behavioral sequences showed that the populations differedsignificantly in their response to sex ratio, and that onlymales from the variable environment (Oklahoma) altered theirbehavior in response to differences in female availability amongthe treatments. The flexible strategy of this population maybe adaptive and probably has evolved in response to sex ratiovariability.     

Energetic constraints and foraging efficiency

Previous research considers foraging options that differ interms of their gross rate of gain b and rate of energy expenditurec. This research argues that maximizing efficiency b/c willmaximize net energetic gain when there is an upper limit onthe amount of energy that can be assimilated. This analysisdoes not include the expenditure during the time for which theanimal is unable to forage because of this constraint. Whenthis expenditure is included, maximizing efficiency is no longeroptimal. Instead the best feeding option is the one with thehighest value of b/(c – c₁), where c, is the metabolicrate when the animal is not foraging.     

The cost of producing a sexual signal: testosterone increases the susceptibility of male lizards to ectoparasitic infestation

According to current evolutionary theory, advertising traitsthat honestly indicate an organism's genetic quality might becostly to produce or maintain, though the kind of costs involvedin this process are controversial. Recently the immunocompetencehypothesis has proposed that testosterone (T) stimulates theexpression of male sexually selected traits while decreasingimmunocompetence. Even though some recent studies have shownan effect of T on ectoparasite load, the dual effect of thehormone has not been addressed in free-living populations. Herewe report results of an experiment in a free-living populationof the lizard Psammodromus algirus during the mating season.Males implanted with T had larger patches of breeding colorand behaved more aggressively than control males. In T-implantedmales, the increase in number of ticks during the mating seasonwas significantly higher than in control males and this negativelyaffected several hematological parameters. T-males sufferedsignificantly higher mortality than control males during theexperiment The results from the manipulation of T are consistentwith the dual effect of this hormone.     

Environmental effects on prenatal growth rate in pronghorn and bighorn: further evidence for energy constraint on sex-biased maternal expenditure

Byers and Moodie (1990) proposed that high levels of maternalexpenditure in polygynous ungulates limit the ability of mothersto support elevated male fetal and neonatal growth rates. Thishypothesis assumes that females in high-expenditure speciesare at or near the maximum level possible and that females inlower-expenditure species are not. To test this assumption,we examined our long-term data on reproduction of pronghorn(Antilocapra americana) and bighorn (Ovis canadensis) at theNational Bison Range (western Montana, USA) and compared gestationlengths, birth weights, and prenatal growth rates of offspringborn following summers of low versus average or above averageprecipitation. In bighorn, these variables were unaffected bythe previous summer's rainfall, but in pronghorn, gestationlength and prenatal growth rate were significantly lower followingdry summers. Extended samples for both species confirmed earlierreports of sex-biased expenditure favoring males in bighornand the absence of sex-bias in pronghorn. Bighorn prenatal littergrowth rates and birth weights, corrected for maternal mass,are 63.5% and 38%, respectively, of pronghorn values. Thesedata support the Byers and Moodie (1990) contention that femalesof high-expenditure species do not show differential expenditureby offspring sex because they are at a reproductive expendituremaximum, whereas females of lower-expenditure species are ableto support excess expenditure in male offspring because optimalallocation to female offspring is farther from such a maximum.     

A prudent hoarder: effects of long-term hoarding in the European nuthatch, Sitta europaea

Although hoarding has been studied intensively for many decades,few studies have attempted to measure its actual fitness consequences.To fill this gap, we used ptilochronology, the growth of replacementfeathers as a measure of nutritional status, and thus a reflectionof starvation risk, of individual European nuthatches (Sittaeuropaea) during winter. We found that nuthatches are long-termhoarders, retrieving stored food up to at least 98 days afterstoring it. Long-term hoarding enhanced the nutritional statusof individual birds significantly because those individualsexperimentally given an opportunity to store seeds during autumnregrew plucked rectrices faster and larger than did controlbirds. Nuthatches used their stored seeds prudently by adjustingthe amounts they ate to meet their requirements, as determinedby ambient temperatures. Nuthatches refrained from using storedfood during periods of relatively benign conditions so theycould use the food during periods of more severe conditions.     

Suppressed breeding in the field vole (Microtus agrestis): an adaptation to cyclically fluctuating predation risk

The densities of microtine rodents and their main predators,small mustelids, fluctuate synchronously in 3–5-year cyclesin central and northern Fennoscandia. Predation by small mustelidshas been suggested as one of the driving forces in microtinecyclicity, causing deep synchronous declines of several volespecies. We studied experimentally the effects of small mustelidson mating behavior, foraging, and breeding in nonwintered fieldvoles (Microtus agrestis) originating from a cyclic population.By using mustelid odors, we simulated a crash phase environmentwith high predation risk for breeding pairs of voles. In ourexperiments, 87% of the female field voles suppressed breedingwhen exposed to mustelid odors. Both female and male behaviorchanged, and no mating behaviors were observed under the simulatedpredation risk. Weights of both sexes decreased when exposedto mustelid odor, probably due to decreased foraging; weightsof the control females increased due to pregnancy; and no weightchanges occurred in control males. Decreased breeding and foragingpossibilities under high predation pressure may form the basisfor the ultimate explanation for breeding suppression. Thereare at least two different mechanisms for breeding suppression:either mating does not take place or malnutrition in femalesdoes not' allow breeding to occur. Delayed breeding under highrisk of predation, for whatever reason, could increase the probabilityof individuals, especially that of the females, to survive overthe crash to the next, safer breeding season when their youngwould have better possibilities to survive.     

Conflict between sexes in the water strider, Gerris lacustris: a test of two hypotheses for male guarding behavior

We studied the effect of operational sex ratio on female reluctanceand male persistence to mate as well as on the length of copulationand postcopulatory guarding in Gerris lacustris by adding fivesurplus males or females to the basin with a pair in tandem.In the control treatment, a pair alone was tested. Accordingto the copulatory guarding hypothesis (CGH), males should prolongmating and guard females in the presence of surplus males. Accordingto the convenience polyandry hypothesis (CPH), females shouldshow lower levels of resistance to prolonged mating in the presenceof surplus males because the mating male protects the femaleagainst harassment from other males. As expected on the basisof both the CGH and CPH, mating (copulation + guarding) averagedlonger in the male-biased treatment. The behavior of males andfemales during mating suggested that both hypotheses hold true:females showed less resistance to prolonged mating (as predictedfrom CPH), and male behavior suggested stronger efforts to stayon the female when surplus males were present (as predictedfrom CGH). Comparisons of the treatment with surplus femaleswith the results from the mating pair without surplus individualssuggested that the capabilities of water striders in tandemto assess the sex of nearby nonmating striders are limited.     

Investigating structure and temporal scale in social organizations using identified individuals

Studies of individually identified animals can produce substantialdata sets containing information on the structure and temporalscale of social organizations. However, methods of analyzingsuch data are not well established. Important features of asocial organization are revealed by plotting the rate of persistenceof the associations between pairs of individuals over a rangeof time lags (lagged association rate). The consistency of long-termrelationships can be characterized using the rate of associationof pairs of individuals between their first and last observedassociations (intermediate association rate). A hierarchicalseries of models featuring exponentially decaying lagged associationrates may be fitted to these data. This technique retrievedthe essential parameters of five simulated social organizationsand, when used on real data, portrayed the essential featuresof the patterns of temporal change in relationships betweenanimals. The method should be especially useful for analyzingfissionfusion societies containing 10–10, 000 individuallyidentifiable animals.     

Dynamic discarding decisions: foraging theory for high-grading in a trawl fishery

The decision by fishermen to discard or retain fish of low valueto make room for more valuable fish in the hold of a boat (high-grading)is similar to diet choice problems faced by natural foragers.In our study, we apply the rationale of diet choice theory tohigh-grading behavior in the Oregon trawl fishery by treatingfishermen as foragers who must decide how much of each net'shaul to "ingest" before searching for more prey. We derive astate-dependent, temporal model of discarding behavior withina fishing trip. This optimization considers the availabilityof differently valued fish, trip quotas set by the regulatoryagency, and the risk of premature trip termination due to lossof gear or injury. The results indicate that those parametersaffect discarding behavior through their effect on the probabilityof exceeding the allowable catch, which we consider analogousto gut capacity. High-grading (partial prey consumption) occurredthroughout many simulated trips. The predictions were consistentwith the trends in discarding observed in the Oregon trawl fleet.Behavioral models such as ours can be useful to fishery managersby providing a means to explore the potential responses of fishermento new regulations before they are implemented.