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The North American clouded leopard (Neofelis nebulosa) population is far from self‐sustaining. Breeding success is poor and behavioral problems (i.e., fur‐plucking, tail‐chewing, excessive hiding or pacing, and intersexual aggression that results in mate killing) are common. This study was undertaken to investigate whether some of these problems may be indicators of chronic stress (as reflected by persistently elevated glucocorticoid levels) and whether they are associated with specific management factors. A fecal corticoid metabolite assay was validated to monitor adrenal activity in clouded leopards. Adrenocorticotropic hormone (ACTH) challenges conducted in four clouded leopards established the biological relevance of the assay system. Fecal corticoid concentrations increased 14‐fold above baseline within 24 hours after ACTH administration. Adrenal activity then was monitored in 72 (36 males; 36 females) clouded leopards (65% of the North American Species Survival Plan population) during a 6‐week period and compared to husbandry and behavior data. There was a significant (P < 0.01) gender difference in fecal corticoid concentrations, with females producing higher concentrations than males. Multiple regression analyses revealed negative associations (P < 0.01) between enclosure height, number of hours keepers spent with each animal per week, and corticoid concentrations. A positive correlation (P < 0.001) was found between the number of keepers caring for an individual and corticoid concentrations. Higher fecal corticoid concentrations (P ≤ 0.05) were measured in clouded leopards kept on public display or near potential predators compared to individuals maintained off exhibit or in the absence of predators. Individuals that performed self‐injuring behaviors also had elevated fecal corticoids (P < 0.01). Spearman‐rank correlation analysis of keeper ratings and hormone data revealed positive associations (P ≤ 0.05) between some behaviors (pace, sleep, hide, and fearful/tense) and fecal corticoid concentrations. Overall these results indicate that noninvasive fecal corticoid monitoring has enormous potential for investigating how management and behavioral problems are related to animal well‐being. If conducted under carefully controlled experimental paradigms, this technique could allow researchers and managers to identify problem areas of captive management for clouded leopards (e.g., enclosure height, keeper time) and evaluate the efficacy of strategies designed to promote animal welfare and increased reproductive success. Zoo Biol 21:77–98, 2002. © 2002 Wiley‐Liss, Inc.  相似文献   

3.

Rehabilitation and release are commonly used for confiscated, surrendered, and rescued primates. To improve release efficacy it is important to generate accurate behavioral profiles of release candidates. Research on primates traditionally uses observer ratings to measure individual differences. This method is easily implemented, but its validity has been questioned. We evaluated whether observer ratings reflect behavioral data indicating forest adaptation in 18 free-ranging rehabilitant orangutans (Pongo pygmaeus morio). In 2017, we used a species-specific questionnaire to measure how often orangutans engaged in behaviors linked to living successfully in the wild (e.g., nest building) and the extent to which they express personality traits that may influence forest adaptation. We collected 11 months of observational data on 17 of the orangutans concurrently to validate the questionnaire items, and collected further questionnaire data for 16 of the individuals in 2019. We used regularized exploratory factor analysis (REFA) and parallel analysis to condense the ratings and determine that two factors could be reliably extracted. We conducted another REFA using the observational data, and calculated factor congruence coefficients following procrustean rotation. The first of the two factors represented forest skills and human aversion, and was congruent with observational data. The second factor reflected boldness, sociability, and exploration, and was not congruent with observational data. Ratings correlated significantly with observations for all five questionnaire items reflecting adaptation to forest life, and for three of seven items reflecting personality traits. We conclude that ratings can be a valid approach to obtain individual-based behavioral information reflecting forest adaptation in free-ranging rehabilitant orangutans, and may be particularly useful in summarizing behaviors relevant to forest adaptation that are otherwise challenging to gather in primates.

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4.
Historically, the cheetah (Acinonyx jubatus) has been known for its poor reproductive performance in captivity. Although breeding success has improved over the past decade, the percentage of breeders in the captive population is still low and successful propagation unpredictable. Estrus in this species has been reported as “silent” by some, therefore contributing to breeding problems by making appropriate timing of pair introductions difficult. To investigate whether any observable behavioral changes may be associated with estrus, we carried out quantitative behavioral observations and concomitant noninvasive monitoring of estradiol metabolites excreted in feces of 14 captive female cheetahs for periods of 5–22 consecutive weeks. We found that changes in fecal estradiol concentrations correlated significantly with variation in the occurrence of several types of behaviors, including rolling, rubbing, sniffing, vocalizing, and urine spraying. However, the number and types of correlated behaviors varied across females, revealing no single behavior indicative of estrus, but rather a constellation of behaviors that increased in frequency when estradiol concentrations were elevated. There was no significant difference in the overall average estradiol concentrations or peak values of the females that had previously mated and conceived compared to those of the females that had failed to breed. Successful breeders appeared to show significantly higher rates of rubbing and rolling than nonbreeders. However, rates of rubbing, rolling, and urine spraying also were found to increase with age, and older individuals were more likely to have bred. The results of this study indicate that estrus in the cheetah cannot be regarded as “silent” since the frequencies of some behaviors appear to covary with fluctuating estradiol levels. However, behavioral monitoring of estrus may nevertheless be difficult and time-consuming due to individual variation and subtle changes in behavioral frequencies rather than changes in the types of behaviors displayed. Zoo Biol 17:193–209, 1998. © 1998 Wiley-Liss, Inc.  相似文献   

5.
The captive population of black rhinoceros (Diceros bicornis) is not self‐sustaining. The reasons for suboptimal reproduction and high mortality need to be investigated. This can only be achieved by cross‐institutional analyses of environments, behavior, and performance. In this study, we collected data on 23 zoos with black rhinoceros to compare zoo environments with reproductive success, mortality, and behavior. Institutional variation was characterized by enclosure area, percentage of walls around enclosure perimeter, percentage of public access along enclosure perimeter, climate, noise level, number of years zoo has maintained black rhinoceros, frequency of chlorine use, and number of male and female black rhinoceros at a zoo simultaneously. Birth and death rates for each institution were calculated from studbook records. We found that the breeding success of a zoo since 1973 correlated positively with enclosure area, and zoos with two or more females had a lower reproductive rate than zoos with only one female. Females residing during their pre‐reproductive years at a zoo with another reproductive female gave birth for the first time on average 3 years later than sole females. Mortality since 1973 correlated positively with percentage of public access. In Part I, we developed behavior profiles of 29.31 individual black rhinoceros from keeper ratings. Scores for males on the behavior trait Fear also correlated positively to percentage of public access, and we suggest that this aspect of black rhinoceros exhibits is a stressor for this species, especially the males. We found that different aspects of captive environments are associated with male and female black rhinoceros behavior. Male scores on the behavior trait dominant were higher in smaller enclosures, and female scores for a group of behaviors suggesting agitation (chasing/stereotypy/mouthing) were positively correlated with percentage of walls in their enclosure. These two behavior traits were found in Part I to be negatively correlated with the breeding success of an individual male or female. We re‐surveyed the behavior and husbandry of 29 black rhinoceros pairs in zoos 2 years after the original data were collected. The re‐survey confirmed that compatible black rhinoceros pairs are those with assertive females and submissive males, and that enclosure area and a low percentage of concrete walls around the enclosure are positive predictors of a pair's reproductive success. We conclude that temperament traits of individuals and characteristics of their captive environments both have an impact on a pair's breeding success. Our study demonstrates that cross‐institutional comparisons of zoo facilities, when integrated with behavioral assessments of individual animals, are a valuable tool for investigating potential causes of poor reproduction and well‐being in zoo animals. Zoo Biol 18:35–52, 1999. © 1999 Wiley‐Liss, Inc.  相似文献   

6.
It is widely expected that physiological and behavioral stress responses will be integrated within divergent stress‐coping styles (SCS) and that these may represent opposite ends of a continuously varying reactive–proactive axis. If such a model is valid, then stress response traits should be repeatable and physiological and behavioral responses should also change in an integrated manner along a major axis of among‐individual variation. While there is some evidence of association between endocrine and behavioral stress response traits, few studies incorporate repeated observations of both. To test this model, we use a multivariate, repeated measures approach in a captive‐bred population of Xiphophorus birchmanni. We quantify among‐individual variation in behavioral stress response to an open field trial (OFT) with simulated predator attack (SPA) and measure waterborne steroid hormone levels (cortisol, 11‐ketotestosterone) before and after exposure. Under the mild stress stimulus (OFT), (multivariate) behavioral variation among individuals was consistent with a strong axis of personality (shy–bold) or coping style (reactive–proactive) variation. However, behavioral responses to a moderate stressor (SPA) were less repeatable, and robust statistical support for repeatable endocrine state over the full sampling period was limited to 11‐ketotestosterone. Although post hoc analysis suggested cortisol expression was repeatable over short time periods, qualitative relationships between behavior and glucocorticoid levels were counter to our a priori expectations. Thus, while our results clearly show among‐individual differences in behavioral and endocrine traits associated with stress response, the correlation structure between these is not consistent with a simple proactive–reactive axis of integrated stress‐coping style. Additionally, the low repeatability of cortisol suggests caution is warranted if single observations (or indeed repeat measures over short sampling periods) of glucocorticoid traits are used in ecological or evolutionary studies focussed at the individual level.  相似文献   

7.
Anger-prone individuals are volatile and frequently dangerous. Accordingly, inferring the presence of this personality trait in others was important in ancestral human populations. This inference, made under uncertainty, can result in two types of errors: underestimation or overestimation of trait anger. Averaged over evolutionary time, underestimation will have been the more costly error, as the fitness decrements resulting from physical harm or death due to insufficient vigilance are greater than those resulting from lost social opportunities due to excessive caution. We therefore hypothesized that selection has favored an upwards bias in the estimation of others' trait anger relative to estimations of other traits not characterized by such an error asymmetry. Moreover, we hypothesized that additional attributes that i) make the actor more dangerous, or ii) make the observer more vulnerable increase the error asymmetry with regard to inferring anger-proneness, and should therefore correspondingly increase this overestimation bias. In Study 1 (N = 161), a fictitious individual portrayed in a vignette was judged to have higher trait anger than trait disgust, and trait anger ratings were more responsive than trait disgust ratings to behavioral cues of emotionality. In Study 2 (N = 335), participants viewed images of angry or fearful faces. The interaction of factors indicating target's formidability (male sex), target's intent to harm (direct gaze), and perceiver's vulnerability (female sex or high belief in a dangerous world) increased ratings of the target's trait anger but not trait fear.  相似文献   

8.
Although white rhinoceros (Ceratotherium simum) are common in captivity, few behavioral studies have been conducted and there is seemingly no research for immersive exhibits where potential for visitor effects is high. Moreover, little information exists on possible effects of weather and temperature on rhino outside their native range. Here we analyze 14,501 observations of rhino in a drive-through enclosure. Data were collected by researchers (n = 12,160 datapoints) and keepers (n = 2341 datapoints) over a 4-month period. We aimed to: (1) quantify behavior using detailed researcher-collected data and contemporaneous but ad hoc keeper-collected data; (2) compare datasets statistically; (3) establish effects of visitors, temperature, and weather on behavior; and (4) assess the influence of visitors on similarity of researcher/keeper datasets. Activity budgets were similar to the wild and the single previous study from a traditional (nondrive-through) enclosure. There was some discrepancy in activity budgets between researcher and keeper data due to significant differences in recorded frequency of two rare behaviors (horn rub; social interaction) and two behaviors that could be easily confused (grazing vs. standing with head-down): recording of other behaviors matched well. Weather and temperature affected behavior, with rhino becoming more sedentary (−locomotion, grazing; +resting, standing, and sedentary eating of hay) on hot/sunny days compared to cool/wet days. The number of visitor vehicles had a fairly negligible effect but resting was lower on busy days, possibly as vigilance increased. The match between researcher/keeper datasets was lowest when visitor numbers were high, suggesting visitors might affect keeper ability to accurately record behavior.  相似文献   

9.
The primary goal of captive breeding programmes for endangered species is to prevent extinction, a component of which includes the preservation of genetic diversity and avoidance of inbreeding. This is typically accomplished by minimizing mean kinship in the population, thereby maintaining equal representation of the genetic founders used to initiate the captive population. If errors in the pedigree do exist, such an approach becomes less effective for minimizing inbreeding depression. In this study, both pedigree‐ and DNA‐based methods were used to assess whether inbreeding depression existed in the captive population of the critically endangered Attwater's Prairie‐chicken (Tympanuchus cupido attwateri), a subspecies of prairie grouse that has experienced a significant decline in abundance and concurrent reduction in neutral genetic diversity. When examining the captive population for signs of inbreeding, variation in pedigree‐based inbreeding coefficients (fpedigree) was less than that obtained from DNA‐based methods (fDNA). Mortality of chicks and adults in captivity were also positively correlated with parental relatedness (rDNA) and fDNA, respectively, while no correlation was observed with pedigree‐based measures when controlling for additional variables such as age, breeding facility, gender and captive/release status. Further, individual homozygosity by loci (HL) and parental rDNA values were positively correlated with adult mortality in captivity and the occurrence of a lethal congenital defect in chicks, respectively, suggesting that inbreeding may be a contributing factor increasing the frequency of this condition among Attwater's Prairie‐chickens. This study highlights the importance of using DNA‐based methods to better inform management decisions when pedigrees are incomplete or errors may exist due to uncertainty in pairings.  相似文献   

10.
The welfare of nonhuman animals in captivity is widely dependent on the natural psychological, physical, and behavioral needs of the animals and how adequately these needs are met. Inability to engage in natural behaviors can lead to chronic stress and expression of stereotypic behavior. The majority of research on decreasing stereotypic behavior in captivity addresses problems at the group level and does not account for individual variability in each animal's needs, history, and preferences. This study combined physiological and behavioral measures of well being to comprehensively assess the unique needs of individually housed captive macaques (Macaca spp.) with the aim of developing tailored welfare strategies. Behavioral and hormonal data were collected under 2 conditions: baseline and individualized enrichment. The results showed a significant decrease in stereotypic behavior under the enrichment condition. Additionally, 7 out of 9 individuals showed a decrease in fecal glucocorticoid (stress hormone) levels, indicating a reduction in hypothalamic-pituitary-adrenal activity. Addressing welfare on an individual, rather than group, level allows for a better overall characterization of well being and maximizes the probability of improving the welfare of each animal.  相似文献   

11.
Objective: To determine whether Stunkard's Figure Rating Scale (FRS) is a valid and reliable index of weight status when an unbiased observer assigns the figure ratings of adult women viewed on videotape. Research Methods and Procedures: Seventy‐two women drawn from a community sample participated in a videotaped study in which height and weight were measured. The FRS is a rating scale displaying 9 silhouettes ranging from very thin to very obese. Women were assigned a figure rating “in‐person” by a research assistant (FRS used as a 17‐point scale) and by additional research assistants viewing women only on videotape (FRS used as both a 17‐ and 9‐point scale). Pearson's correlation coefficients were calculated for in‐person figure ratings, mean videotape figure ratings, and BMI. Results: BMI and in‐person figure ratings were highly correlated (r = 0.91), as were BMI and both mean 17‐point videotape figure ratings and mean 9‐point videotape figure ratings (r = 0.89 and 0.87, respectively). Inter‐rater agreement for in‐person figure ratings and mean 17‐point videotape figure ratings was 0.86, and agreement between in‐person figure ratings and mean 9‐point videotape figure ratings was 0.82. Discussion: The FRS can be used as an index of women's weight status by an unbiased observer, with subjects viewed in‐person or on videotape.  相似文献   

12.
It has been established that various species exhibit personality, defined as intra‐individual consistency and inter‐individual variation in behavioural phenotypes. For example, certain individuals may demonstrate consistently greater behavioural reactions and elevated stress responses. We conducted playback experiments and hormonal analyses on male chimpanzees (Pan troglodytes) in captivity to investigate the patterns and proximate mediators of individual variations in behavioural reactions. We found intra‐individual consistency and inter‐individual variation in behavioural reactions (intensive vigilance towards the direction of speakers) to vocalisations by unfamiliar chimpanzees. This behavioural reaction was positively correlated with changes in salivary cortisol concentration, suggesting that stress is a proximate factor mediating the variation in behavioural reactions. The males who were highly responsive to the conspecific vocalisation also exhibited high behavioural reactions towards the neutral broadcast stimulus (the jungle crow’s Corvus macrorhynchos ‘ka’ vocalisation). This observation is consistent with the notion that male chimpanzees vary in intrinsic behavioural tendency to different stimuli.  相似文献   

13.
Mechanisms affecting consistent interindividual behavioral variation (i.e., animal personality) are of wide scientific interest. In poikilotherms, ambient temperature is one of the most important environmental factors with a direct link to a variety of fitness‐related traits. Recent empirical evidence suggests that individual differences in boldness are linked to behavioral thermoregulation strategy in heliothermic species, as individuals are regularly exposed to predators during basking. Here, we tested for links between behavioral thermoregulation strategy, boldness, and individual state in adult males of the high‐mountain Carpetan rock lizard (Iberolacerta cyreni). Principal component analysis revealed the following latent links in our data: (i) a positive relationship of activity with relative limb length and color brightness (PC1, 23% variation explained), (ii) a negative relationship of thermoregulatory precision with parasite load and risk‐taking (PC2, 20.98% variation explained), and (iii) a negative relationship between preferred body temperature and relative limb length (PC3, 19.23% variation explained). We conclude that differences in boldness and behavioral thermoregulatory strategy could be explained by both stable and labile state variables. The moderate link between behavioral thermoregulatory strategy and risk‐taking personality in our system is plausibly the result of differences in reproductive state of individuals or variation in ecological conditions during the breeding season.  相似文献   

14.
《Animal behaviour》1986,34(4):1016-1025
Fourteen adult female domestic cats were watched by two observers for 3 months. Ratings of 18 aspects of each cat's behavioural style were obtained independently from each observer. Correlations between observers were statistically significant for 15 of the 18 aspects and seven of the correlation coefficients were greater than 0·7. The ratings were compared with results of direct recording methods, where equivalent measures were available and, in five out of six cases, the results of the ratings and direct methods were significantly correlated. The rating method is, therefore, generally reliable and can be adequately validated. Some assessments of observer ratings which are not obviously and easily related to direct recordings may prove particularly useful in developmental studies of alternative modes of behaviour and the origins of individual differences.  相似文献   

15.
In seasonal environments, parturition of most vertebrates generally occurs within a short time‐window each year. This synchrony is generally interpreted as being adaptive, as early born young survive better over the critical season than late born young. Among large herbivores, the factors involved in driving among‐ and within‐individual variation in parturition date are poorly understood. We explored this question by analyzing the relative importance of attributes linked to female quality (longevity, median adult body mass and cohort), time‐dependent attributes linked to female condition (reproductive success the previous year, relative annual body mass and offspring cohort (year)), and age in shaping observed variation in parturition date of roe deer. A measure of quality combining the effects of female longevity and median adult body mass accounted for 11% of the observed among‐individual variation in parturition date. Females of 2 yr old give birth 5 d later than older females. Our study demonstrates that high quality (heavy and long‐lived) females give birth earlier than low quality females. Temporally variable attributes linked to female condition, such as reproductive success in the previous year and relative annual body mass, had no detectable influence on parturition date. We conclude that parturition date, a crucial determinant of reproductive success, is shaped by attributes linked to female quality rather than by time‐dependent attributes linked to female condition in income breeders (individuals that rely on current resource intake rather than on accumulated body reserves to offset the increased energy requirements due to reproduction) such as roe deer.  相似文献   

16.
Ecologists are increasingly documenting individual variation in resource selection across populations in response to temporal or spatial environmental context. These behavioral patterns are assumed to be adaptive although previous studies have not linked them directly to survival and reproductive data to verify the assumed relationship between behavior and fitness. Recent work documented that higher density of secondary roads within home ranges of free‐ranging canids (wolves, coyotes and hybrids) increased mortality risk in the hybrid zone adjacent to Algonquin Park in Ontario, Canada. Here, we examine individual behavioral responses of canids to spatially varying levels of human‐disturbance and determine whether these responses to secondary roads resulted in differential mortality risk for canids across the hybrid zone. Specifically, we investigated resource selection within home ranges with GPS telemetry to determine whether canids selected roads more at night than during the day to minimize dangerous encounters with humans. Next, we modeled individual variation in spatial and temporal responses to secondary roads to evaluate the relative importance of intrinsic (Canis ancestry) and extrinsic (resource availability) influences on their behavior. Behavioral responses to roads were not well explained by variation in Canis ancestry. Instead, canids avoided roads more during day than at night as a non‐linear function of increased road availability. Furthermore, canids that survived exhibited a stronger relationship between day–night selection of roads and road availability than canids that died suggesting an adaptive nature of this behavior. By modifying their selection of roads between day and night, canids appear to be able to exploit the beneficial attributes of roads while mitigating human‐caused mortality risk. However, not all canids responded adaptively highlighting the importance of explicitly linking patterns of resource selection to components of fitness to accurately model and interpret individual variation in resource selection behavior of animals.  相似文献   

17.
Parents of many species provide multiple forms of care to their offspring. In many birds, parents often provision offspring with food and defend them from predators and/or nest‐site competitors. We tested how these two forms of parental care covary in a wild population of house sparrows (Passer domesticus). Using a behavioral reaction norm approach, we found that nestling provisioning exhibited between individual differences and positively covaried with a measure of nest defense: the propensity to attack a heterospecific nest box competitor, the European starling (Sturnis vulgaris). This result would seem to support parental investment theory and suggests that high‐provisioning parents have high‐value offspring, which they will defend more vigorously than low‐provisioning parents. In addition, we found that parents with nestlings that hatched earlier in the season and grew faster approached a model starling more frequently and tended to be more likely to strike the starling. However, we also found that although brood value explained significant variation in both nestling provisioning and nest defense, it did not eliminate the positive, between‐individual relationship between provisioning and defense. This suggests that some of the correlation between provisioning and defense is tied to individual identity and hence may be a behavioral syndrome in which differences between individuals in underlying attributes produce correlated behaviors.  相似文献   

18.
Differences in both stable and labile state variables are known to affect the emergence and maintenance of consistent interindividual behavioral variation (animal personality or behavioral syndrome), especially when experienced early in life. Variation in environmental conditions experienced by gestating mothers (viz. nongenetic maternal effects) is known to have significant impact on offspring condition and behavior; yet, their effect on behavioral consistency is not clear. Here, by applying an orthogonal experimental design, we aimed to study whether increased vitamin D3 content in maternal diet during gestation (vitamin‐supplemented vs. vitamin control treatments) combined with corticosterone treatment (corticosterone‐treated vs. corticosterone control treatments) applied on freshly hatched juveniles had an effect on individual state and behavioral consistency of juvenile Carpetan rock lizards (Iberolacerta cyreni). We tested the effect of our treatments on (a) climbing speed and the following levels of behavioral variation, (b) strength of animal personality (behavioral repeatability), (c) behavioral type (individual mean behavior), and (d) behavioral predictability (within‐individual behavioral variation unrelated to environmental change). We found higher locomotor performance of juveniles from the vitamin‐supplemented group (42.4% increase), irrespective of corticosterone treatment. While activity personality was present in all treatments, shelter use personality was present only in the vitamin‐supplemented × corticosterone‐treated treatment and risk‐taking personality was present in corticosterone control treatments. Contrary to our expectations, behavioral type was not affected by our treatments, indicating that individual quality can affect behavioral strategies without affecting group‐level mean behavior. Behavioral predictability decreased in individuals with low climbing speed, which could be interpreted as a form of antipredator strategy. Our results clearly demonstrate that maternal diet and corticosterone treatment have the potential to induce or hamper between‐individual variation in different components of boldness, often in interactions.  相似文献   

19.
Aggression control is becoming an important component in the management of animals in captivity, but rigorous quantification of aggressive behavior has heretofore been lacking. This study was done to assess the ability of melengestrol acetate (MGA) given with feed (1.54 mg/kg) to control aggression in a bachelor group of fringe‐eared oryx (Oryx gazella callotis). Systematic behavioral observations were conducted and fecal androgen content was measured for 42 and 90 days, respectively, before treatment, and during the 42 days of treatment. There was a significant reduction in concentrations of fecal androgen from 153 ± 6.0 to 95 ± 4.5 ng/g (T66 = 7, P < 0.0001). This reduction in androgen excretion was apparent after the first week of treatment. There was measurable MGA excreted in the feces during treatment. Although treatment did not arrest all aggressive behaviors among animals, the decline in androgens and increase in MGA was accompanied by a significant reduction in several measures of agonistic behavior. Posturing, aggressive contact, pursuit, and submission occurred significantly less frequently after treatment, and there was also a reduction in fighting‐intention movements. Thus, both ritualized and nonritualized aspects of aggression were affected. Reductions in hormones and aggressive behaviors coincided temporally, suggestive of a potential causal relationship. Consistent with this hypothesis is a strong positive correlation between fecal androgen and total aggressive acts. This effect was not the result of a single behavioral element but occurred across several categories of agonistic behavior. Zoo Biol 20:375–388, 2001. © 2001 Wiley‐Liss, Inc.  相似文献   

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