Reproductive Consequences of Aggression in a Territorial Songbird

Territorial aggression can influence males’ ability to obtain high‐quality resources and access to mates; however, in many species, the reproductive consequences of variation in aggression are unknown. In this study, we investigated how individual variation in aggressive behavior relates to reproductive success in socially monogamous, genetically polygynous song sparrows (Melospiza melodia). Prior research in this species shows that male song sparrows differ in their willingness to engage in agonistic interactions with territorial intruders and that individual variation in aggression appears to have functional significance. Aggressive males have been shown to obtain territories where females produce larger clutch sizes, suggesting that individuals who display high levels of territorial aggression are defending high‐quality territories or females. Further, aggressive males are considered a greater threat to territory‐holding males than less aggressive males. In this study, we ask whether individual differences in aggression are linked to differences in extra‐pair reproductive success, annual reproductive success, and offspring quality. We did not uncover a relationship between aggression and annual reproductive success or patterns of extra‐pair paternity. However, we found that the nestlings of aggressive males grew at a faster rate than the nestlings of less aggressive males. Future studies should attempt to identify mechanisms to explain the relationship between offspring growth rate and male aggression and investigate whether faster offspring growth rates translate to greater survival and recruitment of offspring.     

Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

What makes a nest-building male successful? Male behavior and female care in penduline tits

Why do females increase parental effort when caring for theoffspring of attractive males? First, attractive males may bepoor fathers so that their females are compelled to increasetheir own contribution in order to fledge some young (the partner-compensationhypothesis). Second, females mated to attractive males may bewilling to increase their parental effort to reap high indirectbenefits for their offspring, and in turn males can decreasetheir own contribution (the differential allocation hypothesis[DAH]). We investigated these hypotheses in the penduline titRemiz pendulinus, a small passerine bird that has sequentialpolygamy by both sexes and strict uniparental care either bythe male or the female. We focused on two sexually selectedmale traits: nest size and nest-building behavior. We show thatmale care is unrelated to nest-building behavior, whereas femalesare more likely to care for the offspring of those males thatspend more time nest building. Females also more likely carefor the offspring of males that build large nests. Consequently,the reproductive success of males increases with nest size andnest-building behavior. Our results are consistent with theDAH and suggest that nest-building behavior and nest size areunder postmating sexual selection in penduline tits.     

Two Sides of the Same Coin? Consistency in Aggression to Conspecifics and Predators in a Female Songbird

Different forms of aggression have traditionally been treated separately according to function or context (e.g. aggression towards a conspecific versus a predator). However, recent work on individual consistency in behavior predicts that different forms of aggression may be correlated across contexts, suggesting a lack of independence. For nesting birds, aggression towards both conspecifics and nest predators can affect reproductive success, yet the relationship between these behaviors, especially in females, is not known. Here we examine free-living female dark-eyed juncos (Junco hyemalis) and compare their aggressive responses towards three types of simulated intruders near the nest: a same-sex conspecific, an opposite-sex conspecific, and a nest predator. We also examine differences in the strength of response that might relate to the immediacy of the perceived threat the intruder poses for the female or her offspring. We found greater aggression directed towards a predator than a same-sex intruder, and towards a same-sex than an opposite-sex intruder, consistent with a predator being a more immediate threat than a same-sex intruder, followed by an opposite-sex intruder. We also found positive relationships across individuals between responses to a same-sex intruder and a simulated predator, and between responses to a same-sex and an opposite-sex intruder, indicating that individual females are consistent in their relative level of aggression across contexts. If correlated behaviors are mediated by related mechanisms, then different forms of aggression may be expressions of the same behavioral tendency and constrained from evolving independently.     

Sex-specific consequences of life in the city

The vast majority of species are excluded from human-dominatedlandscapes, but some species persist and appear to thrive, despitefacing challenges they would not encounter in less disturbedhabitat. Corticosteroid (cort) secretion may play a significantrole in enabling an animal to cope with urban breeding habitatbecause it helps mediate physiological and behavioral responsesto environmental challenges. Here we present tests of 3 hypothesesrelating to cort secretion in urban animals: 1) environmentalchallenges vary between urban and rural habitat, as reflectedin differing levels of cort secretion in animals breeding inthese habitat types, 2) there are fitness correlates associatedwith variation in cort levels within populations, and 3) parasiteload is one of the mechanistic links between variation in cortlevels and reproductive success. Male white-crowned sparrows(Zonotrichia leucophrys) in urban habitat had significantlyhigher baseline corticosterone levels than males in rural habitat,whereas female Z. leucophrys' hormone levels did not differbetween the 2 habitat types. To assess the fitness correlatesof variation in hormone levels, we monitored seasonal reproductivesuccess in one urban population and found that baseline corticosteronelevels were predictive of female, but not male, reproductivesuccess: females with higher corticosterone fledged fewer offspring.Data do not suggest a direct relationship between corticosteronelevels and fitness and instead indicate an indirect relationshipbetween hormone secretion, territory quality, body condition,and reproductive success. Females with blood parasites had lowerheterophil:lymphocyte ratios, lower body condition scores, fewermates, and fledged fewer offspring than females without parasites.In contrast, parasites did not negatively affect reproductivesuccess in males but were associated with reduced body condition.Corticosterone levels did not differ between birds with andwithout parasites. In tests of all 3 of our hypotheses, we foundsex-specific effects, highlighting the importance of consideringsex in investigation of physiological responses to disturbance.     

Lateralization of Aggression during Reproduction in Male Siamese Fighting Fish

A preference for the left‐eye use during aggressive interactions has been widely reported in the literature, even though in some cases the direction of lateralization varies among individuals within populations. Laterality of aggression in male Siamese fighting fish has been described in a number of studies, yet very little is known about lateralization of aggression during reproduction and/or parental care in fish. Here, we aimed to investigate the relationship between the different reproductive phases and lateralization in eye use during aggressive interactions in males of Siamese fighting fish. Lateralization in eye use is influenced during the early reproductive state, before and after the bubble nest construction stages. We found that nest‐holding males preferentially used the right eye before and after bubble nest construction independent of the sex of the intruder. During the later reproductive phases, aggressiveness increased whereby the direction of lateralization rather than the degree was influenced supporting the hypothesis that reproductive state influences behavioral consistency in Siamese fighting fish.     

Consistent individual variations in aggressiveness and a behavioral syndrome across breeding contexts in different environments in the Black-tailed Gull

Individual behaviors of animals do not evolve separately; they do so in association with other behaviors caused by single shared genetic or physiological constraints and/or favored by selection. Thus, measuring behavioral syndromes—suites of correlated behaviors across different contexts—leads to a better understanding of the adaptive significance of variations in behaviors. However, relatively few studies have examined behavioral syndromes in wild animal populations in changing environments. We investigated a potential behavioral syndrome across antipredator nest defense, territorial defense, chick provisioning, and mating behaviors of male Black-tailed Gulls Larus crassirostris in two successive years under different conspecific territorial intrusion risks and food conditions. Males that presented high levels of antipredator nest defense (aggressive antipredator defenders) against a crow decoy (crows are egg predators) defended their territories against conspecific intruders more frequently than did other males (nonaggressive antipredator defenders), independent of the risk of intrusion. Aggressive antipredator defenders also fed their chicks more frequently than nonaggressive males, but only in a year of low food availability. Taken together, this indicates that males show consistent aggressiveness regardless of breeding context (antipredator and territorial defense), but can regulate food provisioning according to food availability.     

Defending the Weak: Parental Defense Peaks When Chick Vulnerability is Greatest in the Herring Gull (Larus argentatus)

Successful reproduction often depends upon parents providing offspring with resources and protection. In birds, reproductive success can often be enhanced by parents engaging in antipredator behaviors, but these behaviors can be costly. Theoretically, individuals should temporally modify the intensity of nest defense behavior to balance the costs and benefits of current and future reproductive success. More specifically, nest defense should vary throughout a nesting attempt to maximize fitness of the adults. Here, we consider the relationship between nest defense behavior and chick vulnerability in the herring gull (Larus argentatus), where chicks are under high predation risk. We estimated chick vulnerability by quantifying survival probabilities at different periods of the nestling stage. Simultaneously, we quantified changes in parental aggression throughout the nesting cycle by simulating predation attempts using a human predator model. We found that chick survival probabilities were lowest (i.e., vulnerability was highest) and parental aggression in nest defense was greatest during the first 10 days after hatching. Thus, we show that parents are most defensive when chicks are most vulnerable and that adults optimize nest defense behaviors in a way that maximizes their fitness.     

The assessment of reproductive potential in Micronesian kingfisher pairs

The Micronesian kingfisher Halcyon cinnamomina cinnamomina is now extinct in the wild and survives only in U.S. zoos. Although pairs have reproduced in captivity, the captive breeding program has been plagued with inconsistencies in reproductive success. One problem has been determining whether birds housed as reproductive pairs have really established pairbonds. The behavior of newly paired birds may provide information that could be used to predict future reproductive success. Observations conducted for 90 min immediately after introduction of five new pairs at three institutions suggest that the specific behavioral combination of high nest activity and low aggression may indicate reproductive compatibility. Pairs that ultimately produced surviving offspring were less aggressive and engaged in more nest activity than pairs that were not productive. Observations conducted on four of these pairs for an additional month after introduction failed to provide additional information relevant to the assessment of pair status. Although long-term behavioral observations are also desirable, quick assessments such as these may be useful for determining the reproductive potential of more pairs than would be possible using traditional methods. The identification of sexually compatible pairs is a critical step toward the ultimate goal of re-establishing this species in the wild. Zoo Biol 17:425–432, 1998. © 1998 Wiley-Liss, Inc.     

Paternal care as a conditional strategy: distinct reproductive tactics associated with elaboration of plumage ornamentation in the house finch

When individuals in a population differ in physiological conditionand residual reproductive value, selection should favor phenotypicplasticity in reproductive investment such that individualsare able to adopt the reproductive tactic that results in thehighest fitness under given conditions. Here we examined reproductivetactics in relation to the elaboration of condition-dependentsexual ornamentation (carotenoid breast coloration) in a Montanapopulation of the house finches (Carpodacus mexicanus). Malesused distinct reproductive tactics depending on elaborationof their sexual ornamentation. Males with red pigmentation (maximum ornament elaboration) paired with females that nestedearlier, but these males did little provisioning of incubatingfemales and nestlings. In contrast, males with yellow colorationpaired with females that nested later, but these males fedfemale and nestlings more. Consequently, for red males offspringrecruitment was primarily affected by earlier nest initiation, whereas in yellow males it was affected most by male provisioning.In males with intermediate plumage coloration, all measuredcomponents, nest initiation, provisioning of incubating female,and nestling feeding, strongly contributed to offspring recruitment.The fitness consequences of alternative reproductive tacticsof males were influenced by breeding experience and fidelityof their mates. Among first-time breeders, red males achievedthe highest fecundity because of the advantage gained throughearly nesting and pairing with more experienced females andbecause of compensation by their mates for low male provisioningof nestlings. Among experienced breeders, males with intermediateplumage coloration achieved the highest fecundity because ofthe combined benefits of relatively early pairing and high parental care. High variation in sexual ornamentation in a Montana populationof house finches may favor distinct associations of sexualdisplays with a particular set of reproductive behaviors.     

Form,function and consequences of density dependence in a long‐distance migratory bird

The density dependence of demographic parameters and its implications for population regulation have long been recognized. Recent work has revealed potential effects of density on mating systems and sexual selection, but few studies concurrently assess the consequences of density on both demography and sexual selection. Such an approach is important because population processes and individual behaviors can interact to influence population growth and evolutionary trajectories. In this study, we tested the density dependence of breeding success, extra‐pair paternity, and the opportunity for sexual selection in a population of American redstarts Setophaga ruticilla using two different measures of density. To evaluate temporal patterns, we analyzed annual territory density, based on the total number of territories at our study site each year. To evaluate spatial patterns, we analyzed local territory density within years, based on the number of territories surrounding a focal territory. Greater annual density was associated with fewer offspring fledged per female, a reduced mean population rate of fledging success, and a lower relative contribution of extra‐pair paternity to male fitness. Greater local density was associated with fewer offspring fledged, reduced fledgling success, higher rates of nest loss, and higher rates of paternity loss on focal territories. Interestingly, greater local density was also associated with greater nestling mass on focal territories, which could imply that more densely‐packed territories contain superior resources. Overall, our results suggest that the effects of crowding via greater territory density reduce fecundity through increased nest predation, rather than reduced food availability, and increase rates of extra‐pair paternity. Thus, the selective pressures faced by individuals and their reproductive behaviors are likely to differ based on the annual and local density they experience, which may then feed back into population demography.     

Females decide whether size matters: plastic mate preferences tuned to the intensity of male-male competition

Recent studies have indicated that mating success of large malesmay improve under increasing levels of mating competition. Thisoutcome is explained 1) if male mating competition is overridingfemale preferences for male traits that are unrelated to, ornegatively correlated with, male size and dominance and, inso doing, dictates the distribution of matings or 2) if femalesalter their preferences with respect to large males when male–malecompetition is intense. Under both hypotheses, one could expectlarge, dominant males to be more successful under intense competitionthan under weak competition. However, only the first explanationpredicts that male mating success under intense competitionshould be determined by dominance; traits that are unrelatedto male dominance should be uncorrelated to mating success.In contrast, if females change their preferences (explanation2), males with traits beneficial to females independent of thecompetitive environment can maintain a high mating success underall levels of male–male competition. We tested these alternativesusing a small marine fish, the sand goby, Pomatoschistus minutus.The mating success of large males increased under conditionsallowing intense male competition, whereas females showed apreference for good nest building independent of the level ofcompetition. These findings suggest that females are in controlof their choice by altering their preference for male size inresponse to the intensity of male–male competition ratherthan female preference being overshadowed by male dominance.This plasticity of preferences implies that the strength ofsexual selection is not constant at the population level.     

Parental provisioning behaviour plays a key role in linking personality with reproductive success

Repeatable behavioural traits (‘personality’) have been shown to covary with fitness, but it remains poorly understood how such behaviour–fitness relationships come about. We applied a multivariate approach to reveal the mechanistic pathways by which variation in exploratory and aggressive behaviour is translated into variation in reproductive success in a natural population of blue tits, Cyanistes caeruleus. Using path analysis, we demonstrate a key role for provisioning behaviour in mediating the link between personality and reproductive success (number of fledged offspring). Aggressive males fed their nestlings at lower rates than less aggressive individuals. At the same time, their low parental investment was associated with increased female effort, thereby positively affecting fledgling production. Whereas male exploratory behaviour was unrelated to provisioning behaviour and reproductive success, fast-exploring females fed their offspring at higher rates and initiated breeding earlier, thus increasing reproductive success. Our findings provide strong support for specific mechanistic pathways linking components of behavioural syndromes to reproductive success. Importantly, relationships between behavioural phenotypes and reproductive success were obscured when considering simple bivariate relationships, underlining the importance of adopting multivariate views and statistical tools as path analysis to the study of behavioural evolution.     

Behavioral syndromes influence mating systems: floater pairs of a lizard have heavier offspring

Variation in correlated behaviors or behavioral syndromes couldhave interesting effects on mating systems, especially if thevariation in syndrome exists in both sexes. Both males and femalesof the lizard Eulamprus heatwolei display two behavioral typesof a behavioral syndrome, defined by correlations between territorial,exploratory, and predator avoidance behaviors. We tested howthis variation in behavioral syndrome influences reproductivesuccess, pairing patterns, and offspring weight. We used spatialbehavior and residency in the field to identify territorialand floater individuals. Females were relocated to the laboratoryto give birth, and all offspring, dams, and potential sireswere genotyped to determine offspring paternity. During fieldsurveys, 164 lizards were caught of which 27.5% were territorialand the rest were floaters. Paternity was assigned to 66% ofthe 104 offspring produced by 33 dams. Territorial sires fathereda greater proportion of the offspring of territorial dams thanfloater sires. Larger territorial males were more likely tosire the entire clutch or share paternity with fewer additionalsires than smaller territorial males. Floater sire size, however,did not influence the number of fathers per litter. Floaterfemales produced heavier offspring than their territorial counterparts,and offspring fathered by floaters were heavier than maternalhalf-sibs fathered by territorial males. We speculate that differencesin offspring weight may be the result of differences in yolkprovisioning by females and parent genetic compatibility.     

Rapid change in nest size of a bird related to change in a secondary sexual character

Among bird species in which males contribute to nest building,sexual selection has favored larger nests. I investigated determinantsof nest size in the barn swallow Hirundo rustica and how nestsize changed during the period 1977–2003, when tail length(a male secondary sexual character) increased by more than 1.2standard deviations. Males with short tails contributed moreto nest building than long-tailed males, signaling their futureinvestment in food provisioning of offspring. Pairs of barnswallows were consistent in nest size when build ing new neststhe same or different years, and level of phenotypic plasticityin nest size was small and could not account for temporal patternsin nest size. Offspring resembled their parents with respectto nest size, indicating a significant heritability of nestsize, independent of whether offspring were reared by theirparents or by foster parents, and there was a significant negativegenetic correlation between male tail length and outer nestvolume and amount of nest material. The temporal increase inmale tail length was associated with a decrease in nest size,with the amount of nest material in 2003 on average being lessthan a third of the amount used in 1977. Temporal change innest size could be accounted for by indirect selection on taillength causing change in nest size to match that predicted fromchange in tail length and the genetic correlation between maletail length and nest size.     

Temporal Patterns of Territorial Behavior and Circulating Testosterone in the Lapland Longspur and Other Arctic Passerines

We studied territorial aggression in relation to circulatingtestosterone levels in free-living birds of four species innorthern Alaska. The Lapland longspur, Calcarius lapponicus,is an abundant breeding passerine on the arctic tundra. Unlikemany passerines at lower latitudes, male Lapland longspurs donot defend a "multiple-purpose territory" that serves to providenest sites, food and shelter. Rather, after arrival on the breedinggrounds, they perform aerial display flights over a looselydenned "nest area" for a very brief period of two days or so,showing tolerance of other males. This song display may be involvedin courtship. During this phase, male longspurs show a briefand pronounced peak in circulating testosterone levels, andare not aggressive toward simulated territorial intrusions (STIs).Males then "guard" their sexually receptive mates for aboutten days, during which they are highly aggressive toward STIs,but do not sing as much. During the next phase, incubation,the males become very tolerant of conspecific males. Their circulatingtestosterone levels decline to baseline levels, and they generallydo not sing or display aggression in response to STIs. Threeother passerines, the white-crowned sparrow, Zonotrichia leucophrysgambelii, American tree sparrow, Spizella arborea, and savannahsparrow, Passerculus sandwichensis, show patterns of territorialaggression typical of species studied at lower latitudes. Welldefinedterritories are defended for several weeks, during which thereis a prolonged peak in plasma concentrations of testosterone.These three species continue to sing and display aggressioneven late in the season, unlike the longspurs. The peak of testosteronein the longspurs occurs simultaneously with the peak in songdisplay, while in mid-latitude species it occurs with the peakin reproductive aggression. These data suggest that the interrelationshipof testosterone and aggression in Lapland longspurs may be differentfrom that of passerines with multiple-purpose territories, andmay be related to the constraints of breeding in the open arctictundra.     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.     

The kinds of traits involved in male--male competition: a comparison of plumage, behavior, and body size in quail

I compared the role of ornate plumage, behavior, and body sizeduring male—male competition in two species of New Worldquail. Gambel's quail (Callipepla gambelii) is a highly ornateand dichromatic species, whereas scaled quail (C. squamata)is unornamented and monochromatic. During paired contests betweenunfamiliar males, high rates of testosterone-mediated behaviors(tidbitting, calling) and large body size (mass, tarsus, andtail length) corresponded to winners. In the highly ornate Gambel's quail, male head plumes also influenced the outcomeof contests. Plume enhancement made Gambel's quail more likelyto win contests, whereas plume removal made males more likelyto lose. Plume position also reflected male status. Winningmales erected plumes, whereas losers frequently flattened them.Some plumage ornaments, such as belly patches, did not playa primary role during male contests. Unlike static ornaments,head plumes are highly modifiable and likely signal immediateinformation regarding a male's intent, similar to a coverablebadge. Combined, intrasexual selection favored dynamic traits(fast display rates, modifiable ornaments) and static traits(body size) as indicators of male condition or motivation.In scaled quail only, male size was favored both by male—malecompetition and female choice. Accordingly, the degree of sizedimorphism (tarsus length) is greater in scaled than in Gambel'squail. The frequency of overt aggression (chases, pecks, displacement)also differed between species. Gambel's quail were very aggressive,and subordinates often challenged their opponents. In contrast, scaled quail were less aggressive, and subordinates rarely disputedrank. Interspecific comparison indicated differences in themaintenance of male status and possibly in the honesty of signaling.Both appear to be related to differences in social system.     

Sexual selection in a moth: effect of symmetry on male mating success in the wild

Sexual selection is generally caused by female choice and male–malecompetition. In female choice process, female preference isfavored indirectly and/or directly by sexual selection. In indirectselection, females expressing the preference might gain indirectgenetic benefits. In direct selection, females expressing thepreference might gain direct benefits or avoid male-imposedcosts. The white-tailed zygaenid moth Elcysma westwoodii ismonandrous, and males often gather around a female to mate withher, suggesting a high opportunity for sexual selection on maletraits. We quantified phenotypic selection on male morphologyin this species in the field. The morphological characters analyzedincluded body weight, antenna length, forewing length, hindwing length, hind wing tail length, genital clasper length,and the fluctuating asymmetry (FA) of these bilateral traits.In E. westwoodii, selection favored males with more symmetricgenital claspers, as well as longer and more symmetrical hindwings and antennae. Negative correlations between FA and sizewere also detected in the clasper and the antenna. Our resultssuggest that FAs of male traits, in particular the genital clasper,may have indirect and direct influences on mating success. Duringa copulatory attempt, an E. westwoodii male will try to graspthe female's abdominal tip with his claspers but often failto do so because of the female's reluctance to mate. The femaleabdominal tips are smooth and strongly sclerotized and couldthus be difficult for males to grasp. We hypothesize that moresymmetrical male claspers are more efficient in overcoming femalereluctance.     

Circulating androgens are influenced by parental nest defense in a wild teleost fish

While social interactions influence vertebrate endocrine regulation, the dynamics of regulation in relation to specific behaviors have not been clearly elucidated. In the current study, we investigated whether androgens (testosterone) or glucocorticoids (cortisol) play a functional role in aggressive offspring defense behavior in wild smallmouth bass (Micropterus dolomieu), a teleost fish with sole paternal care. We measured circulating testosterone and cortisol concentrations in plasma samples taken from parental males following a simulated nest intrusion by a common nest predator, the bluegill sunfish (Lepomis macrochirus). To understand whether endocrine regulation changes across the parental care period, we looked both at males guarding fresh eggs and at males guarding hatched embryos. Plasma testosterone levels increased in males subjected to a simulated nest intrusion when compared to sham controls. Testosterone concentrations in males guarding embryos were lower than in males guarding fresh eggs, but circulating testosterone was positively correlated with the level of aggression towards the nest predator at both offspring development stages. However, there was no increase in cortisol levels following a simulated nest intrusion, and no relationship between cortisol and any measured parameter. These results suggest that androgens play an important role in promoting aggressive nest defense behavior in teleost fish.