Egg presence, egg loss, and female mate preferences in the sand goby (Pomatoschistus minutus)

We studied the effect of egg presence on female mate choicein a fish with paternal care. Females who were allowed a freechoice between two males mated within a shorter time than femaleswho were randomly assigned to a particular male. When a secondfemale was allowed to choose among the males, she preferredthe same male as the previous female. This result shows thatfemales are concordant in their mate choice. When the initialfemale was randomly assigned to mate with one of two males (forcedchoice), the second female mated randomly with respect to thefirst one. Thus females do not prefer males with eggs. If theinitial female was given a free choice, but the eggs were removedfrom the chosen male, the test female mated randomly. When boththe males initially had mated but one randomly determined male'seggs were removed, the test female preferred the male who wasstill guarding eggs. These experiments show that females avoidspawning in unsuccessful nests. When the females in the freechoice/egg removal experiment mated with the unsuccessful malethere was a considerably bigger size difference in favor ofthis male than when the females mated with the other male. Weconclude that female sand gobies show clear mate preferences,but that they do not prefer males with eggs over males withouteggs. They do, however, avoid mating with males guarding unsuccessfulnests. We therefore suggest that egg loss could be an importantfactor selecting for egg preference.     

Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea)

The degree and direction of sexual dimorphism varies widely,but in several taxa of orb-weaving spiders, including Nephila,males may be less than one-tenth the size of females. This differenceis commonly attributed to selection through precopulation sexualcannibalism: females may either fail to detect very small males,or ignore them as potential prey items. However, there is oftenthe potential for male-male competition in these species becauseseveral males can be found on the web of a single female. Weinvestigated experimentally the effects of sexual cannibalismand male-male competition on male body size and hence sexualdimorphism in the Australian golden orb-weaver (Nephila plumipes).Small males were less likely to be detected and cannibalizedthan larger males. However, larger males excluded small malesfrom the central hub of the web, where mating takes place. Theconflicting effects of sexual cannibalism and male-male competitionmay be responsible for the relatively large variation in malebody size in this species.     

Temperature affects male and female potential reproductive rates differently in the sex-role reversed pipefish, Syngnathus typhle

The differences in potential reproductive rate between the sexescan be used to predict the operational sex ratio and the patternsand intensity of mating competition and hence sexual selectionin a population. This article describes how one environmentalcomponent, temperature, affects potential reproductive ratesof the two sexes in the paternally brooding, sex-role reversedpipefish (Syngnathus typhle). Males brooded embryos much longer(on average 58 days) in cold water (about 10°C) than inwarmer water (35 days at about 15°C). As a consequence,the potential reproductive rate (number of eggs brooded perday) of males was significantly higher in warm water. In females,however, potential reproductive rate, i.e., number of eggs producedper day given an unlimited access to mates, was not significantlydifferent between temperatures. In both sexes, potential reproductiverate was positively related to body size. At both temperatures,females had the potential to reproduce faster than males. Asa result, the operational sex ratio will become female biasedand sex-roles reversed, as is the case in this species. Sincetemperature differently influenced the potential reproductiverates of males and females, with the sexual difference largerat lower temperatures, more intense female-female competitionis predicted at low temperatures.     

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.     

Sexual selection and extrapair fertilization in a socially monogamous passerine, the zebra finch (Taeniopygia gullata)

We investigated paternal exclusion rate (the percentage of youngreared by a male that were not his genetic offspring), and behavioraland reproductive variables influencing this rate, in a freelybreeding laboratory population of zebra finches (Tae-niopygiaguttala castanotis), a socially monogamous grassfinch. Priorto the experiment, each male founder was fitted with eithertwo red bands (creating a phenotype previously demonstratedto be attractive to females) or two green bands (unattractiveto females) as part of a unique combination of four leg bands.The overall paternal exclusion rate was 28%, as determined bymultilocus, minisatellite DNA fingerprinting of 278 offspringreared by 26 males and their mates. Mean exclusion rates were16% and 40% for red- and green-banded males, respectively. Exclusionrates were directly proportional to rates of female participationin unforced extrapair copulations (UEPCs) with red-banded malesthat occurred when females were fertile. Rates of fertile, forcedextrapair copulations (FEPCs) and fertile UEPCs involving green-bandedmales either failed to influence exclusion rate or varied inverselywith exclusion rate, indicating that extrapair fertilization(EPF) is under female control. Effort devoted by males to seekingEPFs increased exclusion rate. Results suggest that males placegreater effort into seeking fertile versus infertile EPCs andthat unattractive males accrue fitness gains through high parentalinvestment (PI), whereas attractive males benefit through decreasedPI and increased allocation to EPF.     

Mate-choice copying under predation risk in the Trinigadian guppy (Poecilia reticulata)

Although females of numerous species possess genetically-basedpreferences for certain male trails and male preferentiallywith males possessing these traits, recent theoretical and experimentalevidence indicates that they may also copy (imitate) the matechoices of other females under certain circumstances. Such mate-choicecopying is expected to be most prevalent when females have theopportunity to observe the mate choices of others and when matechoice is costly to females. One potential direct fitness costof mate choice is increased individual risk of mortality dueto predation. Here, we investigate for the first time the effectof increasing the apparent risk of predation on the tendencyof females to copy the experimentally staged mate choice ofanother female. Using adult female guppies (Poecilia reticulata)originating from a Trinidadian population that experiences arelatively high fish predation pressure, we first establishthat they possess a preference for the more brighdy coloredof two simultaneously presented males in the absence of bothmate-copying opportunity and an immediate threat of predation.However, most females reversed their initial mate preferencewhen given an opportunity to copy the mate preference of anotherfemale in the absence of predation threat The proportion offemales reversing their preference when given the opportunityto do so was not affected by increasing the apparent risk of(fish) predation. This result may be owing either to femaleguppies tending to copy the mate choice of others whenever theopportunity arises because the benefits of doing so accrue irrespectiveof the ambient risk of predation or to females choosing randomlybetween males with respect to their color pattern in the presenceof the predator irrespective of mate-copying opportunity. Thesetwo explanations for the apparent lack of an effect of predationrisk on mate-choice copying per se are both plausible but unfortunatelycould not be easily distinguished here. It may thus be possible,and interesting, that individual female guppies chose randomlybetween the available males in the presence of the predatorbut otherwise copied the choice of others when given the opportunityto do so.     

Sexual ornamentation and immunocompetence in the barn swallow

The handicap hypothesis of honest signaling suggests that secondarysexual characters reliably reflect phenotypic or genotypic qualityof signalers. This hypothesis is based on the assumptions thatsignals are costly to produce and/or maintain and the cost ofa given level of signaling is higher for low quality than forhigh quality signalers. We tested these assumptions in a fieldexperiment in which the size of a secondary sexual character[tail length in male barn swallows (Hirundo rustica)] was experimentallymanipulated. Males were randomly assigned to tail elongation,tail shortening, or two control treatments (tail manipulation,or just capture, ringing, and handling). Male barn swallowswere challenged with an injection of sheep red blood cells,and blood was sampled on the day of first capture and after3 to 4 weeks for determination of concentrations of gamma-globulins.Tail-elongated males did not increase levels of gamma-globulinswhile males of the other three groups demonstrated increases.Analyses of variation in gamma-globulins within treatment groupsrevealed a positive correlation between gamma-globulins andoriginal tail length among males with elongated tails. Theseresults suggest that tail length imposes an immu-nocompetencecost on males, and that males with naturally long tails aredifferentially better able to cope with this cost.     

Unifying cornerstones of sexual selection: operational sex ratio,Bateman gradient and the scope for competitive investment

What explains variation in the strength of sexual selection across species, populations or differences between the sexes? Here, we show that unifying two well‐known lines of thinking provides the necessary conceptual framework to account for variation in sexual selection. The Bateman gradient and the operational sex ratio (OSR) are incomplete in complementary ways: the former describes the fitness gain per mating and the latter the potential difficulty of achieving it. We combine this insight with an analysis of the scope for sexually selected traits to spread despite naturally selected costs. We explain why the OSR sometimes does not affect the strength of sexual selection. An explanation of sexual selection becomes more logical when a long ‘dry time’ (‘time out’, recovery after mating due to e.g. parental care) is understood to reduce the expected time to the next mating when in the mating pool (i.e. available to mate again). This implies weaker selection to shorten the wait. An integrative view of sexual selection combines an understanding of the origin of OSR biases with how they are reflected in the Bateman gradient, and how this can produce selection for mate acquisition traits despite naturally selected costs.     

The function of long tails in female barn swallows (Hirundo rustica): an experimental study

The outermost tail feathers in male barn swallows (Hirundo rustica)are the target of a strong directional female mate preference.The tail ornament is also expressed in females, since femaleshave considerably longer tails than juveniles, either due to(1) a strong genetic correlation between the characters in thetwo sexes, or (2) direct sexual selection on females. To discriminatebetween these two hypotheses, we manipulated the length of theoutermost tail feathers in female barn swallows shortly afterarrival by either shortening or elongating the outermost tailfeathers, or maintaining their length among control individuals.Start of laying of the first clutch, reproductive performance,or provisioning of offspring did not show any significant differencesamong treatments. Original female tail length before manipulationwas unrelated to reproductive performance, while male tail lengthexplained some variation in the number of clutches and, to someextent, the total number of eggs laid per year. Females withlonger tails arrived earlier at the breeding grounds. Manipulatedfemale tail length was positively correlated to the tail lengthof their mates. Our results support the correlated responsehypothesis but do not support the sexual selection explanationfor the existence of exaggerated tail feathers in female barnswallows.     

Predation, sensitivity, and sex: why female black rhinoceroses outlive males

Among sexually dimorphic, polygynous mammals, adult femalestend to outlive males and respond more strongly to predatorsthan males. We asked whether a monomorphic, polygynous speciesvirtually immune to predation due to large size (black rhinoceros,Diceros bicornis) conforms to this pattern. Data on 193 interactionswith lions (Panthera leo) and spotted hyenas (Crocuta crocuta)in two nonhabituated populations in Namibia studied from 1991to 1993 revealed that: (1) females were more vigilant or aggressivethan males to either of the potential predators; and (2) whethersolitary or with calves, females attacked more often than males.Although solitary females tended to be more aggressive to lionsthan to hyenas, neither females with calves or males seemedto discriminate between the two carnivores. We also simulatedthe behavior of human predators (poachers) during 69 encounterswith rhinoceroses. While both sexes abandoned local sites becauseof our presence, females ran farther than males, covering upto 40 km in a day. These findings implicate a behavioral mechanismto explain why secondary sex ratios favor females - males aremore prone to human predation, a prediction consistent withdata from 12 populations throughout Africa. Black rhinocerosesappear to be an unanticipated exception to the well-establishedpattern of male-biased mortality in polygynous mammals; in theabsence of intense human predation (a recent event), male mortalityfails to exceed that of females, suggesting that intrasexualcompetition in a polygynous mammal: may not be the primary causeof unbalanced secondary sex ratios. Our results on the causesof sex differences in mortality and in responsiveness to differentpredators reinforce the relevance of behavioral ecology to conservation;such information is necessary for planning how best to minimizenegative human influences on the few remaining wild Africanrhinos.     

Testosterone effects on the immune system and parasite infestations in the barn swallow (Hirundo rustica): an experimental test of the immunocompetence hypothesis

The immunocompetence hypothesis predicts that testosterone (T)enhances the expression of male secondary sexual characterswhile exerting a suppressive effect on the immune system therebyexposing hosts to higher intensities of parasite infestations.In a natural population of barn swallow (Hirundo rustica) males,the intensity of infestation by some ectoparasites was negativelycorrelated with tail length and was positively correlated withimmunoglobulin levels, but no clear relationship was observedbetween immune responses (leukocyte counts, immunoglobulins)and tail length. Males implanted with T had higher intensitiesof parasite infestations at the time of recapture than controlmales, and T-implanted males experienced an increase in countsof eosinophils. In T-implanted males, immunoglobulin levelsinitially decreased and then increased as time from implantationelapsed. Among T-implanted males, those with longer tails hada smaller increase in eosinophil counts, tended to experiencea smaller increase of parasite infestations, and were more likelyto survive until the following breeding season than those withshorter tails. The relationships between parasite burden, immunesystem, and exaggeration of tail length in the natural populationof males are consistent with some aspects of the immunocompetencehandicap hypothesis. The results from the manipulation of Tplasma levels are also partly consistent with the hypothesis,since T-implantation resulted in higher levels of parasite infestations,but contradict the assumption of an obligatory immunosuppressiveeffect of T. Higher activation of the immune system of T-implantedmales indicate that high T plasma levels imposed a two-foldcost because of the effects on parasites and the immune responseto parasites, and this suggests that the effect of T on parasitesmight not be mediated by the immune system of the host. Theresults of the manipulation of T plasma levels support the handicapversion of the immunocompetence hypothesis since high quality,long-tailed males paid less in terms of activation of the immunesystem, change in parasite infestations, and chances of survivalthan low-quality, short-tailed males.     

Correlates of male quality in the field cricket, Gryllus campestris L.: age, size, and symmetry determine pairing success in field populations

Sexual selection in field crickets (Gryllidae) is well documented.Several studies have identified male traits, such as age andbody size, that influence pairing success in the field. HereI show how these traits covary with male quality by examiningthe degree of fluctuating asymmetry in male Gryllus campestris.Older males were both larger and more symmetrical. Principalcomponents analysis suggests that these three variables wereessentially measures of the same trait, male quality. A comparisonof paired and calling males showed that males of high qualitywere more successful in obtaining mates; paired males were older,larger, and more symmetrical. The area of the harp covariedwith morphological traits and determined the carrier frequencyof the male's call. Information related to male quality wastherefore available for female discrimination.     

Fluctuating asymmetries and advertisement call variation in the cricket frog, Acris crepitans

We used an anuran acoustic communication system to test a predictionof the "fluctuating asymmetries/good genes" hypothesis thatfemales prefer more symmetric mates because symmetry indicatesgenetic quality. Mate preferences of female cricket frogs (Acriscrepitans) can be influenced by three call characters: dominantfrequency, numbers of pulses per call, and number of pulse groupsper call. We tested the hypothesis that these preferences resultin females preferring more symmetric males. We measured fluctuatingasymmetries of characters not involved with the communicationsystem (head and tibia), and those involved in signal production(laryngeal characters) and signal reception (aural characters).We determined whether the asymmetries in these characters wererelated to the three variables that enhance call attractiveness.Most of the multiple regression models showed no significantassociation between the fluctuating asymmetries of charactersand any of the calls. The regression of head and tibia fluctuatingasymmetry on pulse number was significant, but partial regressioncoefficients revealed that more pulses were associated witha more symmetric head length and a less symmetric tibia length.Our findings provide little or no support for the fluctuatingasymmetries/good genes hypothesis. We emphasize, however, thatthis hypothesis should not be abandoned based on negative resultsof a single study, but deserves further scrutiny.     

A shift in female social organization independent of relatedness: an experimental study on the field vole (Microtus agrestis)

Home range size, overlap between home ranges, movement patternsand social behavior in two generations of female field voleswere examined by radio tracking unrelated individuals in enclosedareas. In addition, potential effects of increased relatednesswere examined by comparing setups with groups of sisters andunrelated females. Overwintered females showed less overlapbetween ranges and moved shorter and more irregular distancescompared with females of the year. Based on responses to introduced,unfamiliar conspecifics it was concluded that the overdisperseddistribution of overwintered females was maintained primarilyby avoidance behavior and that infanticidal behavior was equallyfrequent in the two generations. Observed differences betweengenerations could not be ascribed to differences in relatedness,and experimental increase of the degree of relatedness had negligibleeffects. The overdispersion and restricted movements shown byoverwintered females are instead suggested to be caused by acombination of low food availability and exposure to intensepredation in spring/early summer, whereas spacing and movementpatterns of females of the year are less influenced by thesefactors. As aggressive interactions are rare and populationdensity is usually low in spring, it is unlikely that breedingdensity and population growth is restrained by the social behaviorof overwintered females.     

Early arrival, initiation of nesting, and social status: an experimental study of breeding female red-winged blackbirds

Red-winged blackbirds (Agelaius phoeniceus) arrive at breedingsites several months before nests are constructed. Males inthis highly polygynous species presumably return early to defenddesirable territories. Females, however, also begin to arrivealmost two months before nesting is attempted. Early returnto breeding sites could enhance reproductive success by increasinga female's social status and thereby allowing earlier nesting.I measured the effect of experimentally delayed arrival on thetiming of nesting, reproductive success, and social status offemales. Birds were captured as they arrived in early springand detained in an aviary. These experimental females were laterreleased at their capture sites before control females beganconstruction of nests; controls had arrived during the sameperiod as experimentals but were not detained. Experimentalfemales nested, on average, more than a week later than controls,although I could detect no effect of timing on reproductivesuccess during this study. By manipulating arrival date, butnot covariates of arrival time such as age or experience, thisresult indicates that timing of arrival directly influencedlaying date. Delayed females were subordinate to control femaleswith the same mate. This decline in social status may have beenresponsible for the delay in nesting and could have fitnessconsequences for females in some years, because the offspringof earlier nesting, dominant birds are more likely to receivemale parental care in this population.     

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.     

Parental effort of kestrels (Falco tinnunculus) in nest defense: effects of laying time, brood size, and varying survival prospects of offspring

We studied the nest defense behavior of Eurasian kestrels (Falcotinnunculus) towards a stuffed pine marten (Martes martes) througha 3-year vole cycle (1990–92) in western Finland. Survivalprobability of offspring decreases with a later start of breeding,and, therefore, early breeders should protect their offspringmore than late ones. We found this true for males during theincubation period, but not for females. In addition, we expectedthe nest defense intensity to increase with offspring number.During the incubation period, this was true for females, butnot for males. During the nestling phase, parents did not adjusttheir defense effort to natural or manipulated (by one to twoyoung) brood size. Survival prospects of kestrel offspring werehighest in the increasing vole year 1991 and lowest in the decreasingvole year 1992, and, therefore, we expected the defense activityof kestrels to follow the same trend. However, the oppositeresult appeared true for females with a similar tendency formales. Most hypotheses predicting avian nest defense behaviorwere not supported by our data. Temporally heterogeneous environmentand low degree of nest-site tenacity of migratory kestrels maymake them unfamiliar with environmental variation and survivalprospects of their offspring. Therefore, fitness benefits ofparental care are not predictable, and kestrels may thus adjusttheir parental effort to their own future reproductive potential(i.e., number of future breeding attempts), rather than to somecurrent investment indicator, like offspring age and number.