Predictors of helminth parasite infection in female chacma baboons (Papio ursinus).

Helminth parasite infection can impose major consequences on host fitness. Several factors, including individual characteristics of hosts, environmental conditions, and patterns of coinfection, are thought to drive variation in parasite risk. Here, we report on four key drivers of parasite infection-phase of reproduction, steroid hormone profiles, rainfall, and patterns of coinfection-in a population of wild female chacma baboons (Papio ursinus) in South Africa. We collected data on reproductive state and hormone profiles over a 3-year span, and quantified helminth parasite burdens in 2955 fecal samples from 24 female baboons. On a host level, we found that baboons are sensitive to parasite infection during the costliest phases of the reproductive cycle: pregnant females harbored higher intensities of Protospirura eggs than cycling and lactating females; lactating and cycling females had a higher probability of Oesophagostomum infection than pregnant females; and cycling females exhibited lower Trichuris egg counts than pregnant and lactating females. Steroid hormones were associated with both immunoenhancing and immunosuppressive properties: females with high glucocorticoid concentrations exhibited high intensities of Trichuris eggs but were at low risk of Oesophagostomum infection; females with high estrogen and progestagen concentrations exhibited high helminth parasite richness; and females with high progestagen concentrations were at high risk of Oesophagostomum infection but exhibited low Protospirura egg counts. We observed an interaction between host reproductive state and progestagen concentrations in infection intensity of Protospirura: pregnant females exhibited higher intensities and non-pregnant females exhibited lower intensities of Protospirura eggs with increasing progestagen concentrations. At a population level, rainfall patterns were dominant drivers of parasite risk. Lastly, helminth parasites exhibited positive covariance, suggesting that infection probability increases if a host already harbors one or more parasite taxa. Together, our results provide a holistic perspective of factors that shape variation in parasite risk in a wild population of animals.

Male rutting calls synchronize reproduction in Serengeti wildebeest.

Tightly synchronized reproduction in vast wildebeest herds underpins the keystone role this iconic species plays in the Serengeti. However, despite decades of study, the proximate synchronizing mechanism remains unknown. Combining a season-long field experiment with simple stochastic process models, we show that females exposed to playback of male rutting vocalizations are over three times more synchronous in their expected time to mating than a control group isolated from all male stimuli. Additionally, predictions of both mating and calving synchrony based on the playback group were highly consistent with independent data on wildebeest mating and calving synchrony, while control-based predictions were inconsistent with the data. Taken together, our results provide the first experimental evidence that male rutting vocalizations alone could account for the highly synchronized reproduction observed in Serengeti wildebeest. Given anthropogenically driven losses in many areas, a mechanistic understanding of synchrony can highlight additional risks declining wildebeest populations may face.

Social status and helminth infections in female forest guenons (Cercopithecus mitis).

OBJECTIVE: When resource competition within primate social groups is effective, high-ranking individuals generally gain fitness benefits. Contrary to expectations, female Cercopithecus mitis form linear dominance hierarchies without evidence for rank-related variation in fitness-relevant measures, raising questions about the evolution of guenon social structure. Here, we test whether social status predicts gastrointestinal helminth infections, known to influence health and morbidity in other mammalian hosts. In addition, we assess whether infections contribute to stress responses as indicated by fecal glucocorticoid (fGC) levels. METHODS: We quantified infections and hormone levels in 382 fecal samples from 11 adult female Sykes' monkeys (C. m. albogularis) over four months in one wild study group at Gede Ruins, Kenya. Using a generalized estimating equations technique, we modeled the odds of infection, relative infection intensity, and fGC variation. RESULTS: High-ranking females were less likely infected with Trichuris and Trichostrongylus, had lower fecal egg counts for both taxa, and overall lower helminth richness than low-ranking females. An inverse relationship between rank and Trichuris egg counts existed also in a study population of blue monkeys (C. m. stuhlmanni), where we collected comparable data over a shorter period. Regardless of rank, lactating females were more likely than non-lactating females to be infected with Trichuris, and had higher fecal egg counts for both Trichuris and Oesophagostomum. Lastly, we report evidence that Trichuris infections exacerbated energetic stress and that food supplementation by tourists increased infection levels. CONCLUSION: Our findings suggest that high-rank may provide long-term health and energetic benefits for female C. mitis, with potential fitness implications.

Mean fecal glucocorticoid metabolites are associated with vigilance, whereas immediate cortisol levels better reflect acute anti-predator responses in meerkats.

Adrenal hormones likely affect anti-predator behavior in animals. With experimental field studies, we first investigated associations between mean fecal glucocorticoid metabolite (fGC) excretion and vigilance and with behavioral responses to alarm call playbacks in free-ranging meerkats (Suricata suricatta). We then tested how vigilance and behavioral responses to alarm call playbacks were affected in individuals administered exogenous cortisol. We found a positive association between mean fGC concentrations and vigilance behavior, but no relationship with the intensity of behavioral responses to alarm calls. However, in response to alarm call playbacks, individuals administered cortisol took slightly longer to resume foraging than control individuals treated with saline solution. Vigilance behavior, which occurs in the presence and absence of dangerous stimuli, serves to detect and avoid potential dangers, whereas responses to alarm calls serve to avoid immediate predation. Our data show that mean fGC excretion in meerkats was associated with vigilance, as a re-occurring anti-predator behavior over long time periods, and experimentally induced elevations of plasma cortisol affected the response to immediate threats. Together, our results indicate an association between the two types of anti-predator behavior and glucocorticoids, but that the underlying mechanisms may differ. Our study emphasizes the need to consider appropriate measures of adrenal activity specific to different contexts when assessing links between stress physiology and different anti-predator behaviors.

"Mayday mayday mayday", the millennium ark is sinking!

Despite exceptional advances in ensuring the health and well-being of animals in human care, zoos of the twenty-first century are ill-prepared and overwhelmed by the sheer number of species requiring conservation support. Furthermore, small population management paradigms have failed to achieve the demographic and genetic targets required to sustain most endangered species in human care. Predictions made in the 1980s regarding the potential of a "millennium ark"-aided by the use of assisted reproductive technologies (ARTs)-for saving species have proven to be wildly over-optimistic. ARTs continue to be touted as a panacea for saving endangered species and even for resurrecting extinct ones. And yet, while the first successful interspecies embryo transfer in a wildlife species occurred 30 years ago, there still is not a single example of embryo-based technologies being used to consistently manage a conservation-reliant species. The limited contribution of ARTs to species conservation to date principally stems from the lack of knowledge of species biology, as well as inadequate facilities, space, expertise, and funding needed for their successful application. ARTs could and should be an important tool in our conservation toolbox, but we cannot fall into the trap of believing that we can "assist" or clone our way out of the present biodiversity crisis. Reproductive technologists overstate the potential of ARTs for saving endangered species, zoos overestimate their ability to sustain genetically and demographically viable captive populations with existing resources, and conservationists underestimate their need for zoos in the face of failing efforts to sustain species in nature. Unless all parties concerned-reproductive technologists, zoo biologists and conservationists-adopt parallel efforts to sustain wild populations and places, zoos risk becoming living museums exhibiting relic species that no longer exist in nature.

Oral and injectable synthetic progestagens effectively manipulate the estrous cycle in the Przewalski's horse (Equus ferus przewalskii).

To date, there has been limited research on manipulation of the estrous cycle in endangered equids. The objectives of this study were to assess the efficacy of using combinations of: (a) oral altrenogest and PGF2α, and (b) injectable altrenogest and PGF2α for manipulation of ovarian activity in Przewalski's mares. Reproductive cycles were monitored by assessing follicular changes with rectal ultrasound and changes in urinary steroid hormones. In Study 1, five cycling mares were treated with oral altrenogest (n=11 cycles) for 14 days. In Study 2, cycling mares were treated with oral altrenogest for 12 days (n=5 cycles; n=5 mares) or a single injection of biorelease altrenogest (n=10 cycles; n=6 mares). In all study groups, PGF2α was given 2 days before cessation of progestagen treatment. In Study 1, mares responded in six of 11 cycles (54%) where treatment occurred with normal ovarian follicular development post hormone therapy. In Study 2, mares responded in four of five (80%, oral altrenogest) and eight of 10 (80%, injectable altrenogest) cycles with the development of an ovulatory follicle. With the use of injectable altrenogest, there was an obvious suppression of urinary estrogens and progetsagens. These results indicate that manipulation of the estrous cycle of Przewalski's mares can be achieved by administering oral (12 days) or injectable form of altrenogest in conjunction with PGF2α. Findings in the present study may have long term application for the development of timed artificial insemination as a genetic management tool for this critically endangered equid.

Seasonal energetic stress in a tropical forest primate: proximate causes and evolutionary implications.

Animals facing seasonal variation in food availability experience selective pressures that favor behavioral adjustments such as migration, changes in activity, or shifts in diet. Eclectic omnivores such as many primates can process low-quality fallback food when preferred food is unavailable. Such dietary flexibility, however, may be insufficient to eliminate constraints on reproduction even for species that live in relatively permissive environments, such as moist tropical forests. Focusing on a forest-dwelling primate with a flexible diet (Cercopithecus mitis) we investigated whether females experience seasonal energetic stress and how it may relate to reproductive seasonality. We used fecal glucocorticoids (fGCs) as an indicator of energetic stress, controlling for the potentially confounding effects of social interactions and reproductive state. We modeled within-female fGC variation with General Linear Mixed Models, evaluating changes in feeding behavior and food availability as main effects. Regardless of reproductive state, fGCs increased when females shifted their diet towards fallback foods (mature leaves and other non-preferred items) and when they spent more time feeding, while fGCs decreased with feeding time on preferred items (insects, fruits, young leaves) and with the availability of young leaves. Changes in fruit availability had no general effects on fGCs, likely because fruits were sought out regardless of availability. As predicted, females in the energetically demanding stages of late pregnancy and early lactation showed greater increases in fGCs between periods of low versus high availability of fruits and young leaves than females in other reproductive states. Potential social stressors had no measurable effects on fGCs. Preliminary evidence suggests that seasonal energetic stress may affect the timing of infant independence from mothers and contribute to unusually long inter-birth intervals compared to closely related species of similar body size. Our findings highlight how the study of stress responses can provide insights into the proximate control of reproductive strategies.

Adrenal activity and anxiety-like behavior in fur-chewing chinchillas (Chinchilla lanigera).

Due to its complexity, in combination with a lack of scientific reports, fur-chewing became one of the most challenging behavioral problems common to captive chinchillas. In the last years, the hypothesis that fur-chewing is an abnormal repetitive behavior and that stress plays a role in its development and performance has arisen. Here, we investigated whether a relationship existed between the expression and intensity of fur-chewing behavior, elevated urinary cortisol excretion and anxiety-related behaviors. Specifically, we evaluated the following parameters in behaviorally normal and fur-chewing animals of both sexes: (1) mean concentrations of urinary cortisol metabolites and (2) anxiety-like behavior in an elevated plus-maze test. Urinary cortisol metabolites were higher only in females that expressed the most severe form of the fur-chewing behavior (P≤0.05). Likewise, only fur-chewing females exhibited increased (P≤0.05) anxiety-like behaviors associated with the elevated plus-maze test. Overall, these data provided additional evidence to support the concept that fur-chewing is a manifestation of physiological stress in chinchilla, and that a female sex bias exists in the development of this abnormal behavior.

Reproductive rate, not dominance status, affects fecal glucocorticoid levels in breeding female meerkats.

Glucocorticoid hormones (GCs) have been studied intensively to understand the associations between physiological stress and reproductive skew in animal societies. However, we have little appreciation of the range of either natural levels within and among individuals, or the associations among dominance status, reproductive rate and GCs levels during breeding. To address these shortcomings, we examined variation in fecal glucocorticoid metabolites (fGC) during breeding periods in free-ranging female meerkats (Suricata suricatta) over 11 years. The vast majority of variation in fGC levels was found within breeding events by the same female (~87%), with the remaining variation arising among breeding events and among females. Concentrations of fGC generally tripled as pregnancy progressed. However, females with a high reproductive rate, defined as those conceiving within a month following parturition (mean = 9 days postpartum), showed significant reductions in fGC in the final 2 weeks before parturition. Despite these reductions, females with a high reproductive rate had higher fGC levels at conception of the following litter than those breeding at a low rate. After controlling for the higher reproductive rate of dominants, we found no association between levels of fGC and either age or dominance status. Our results suggest that one should be cautious about interpreting associations between dominance status, reproductive skew and GCs levels, without knowledge of the natural variation in GCs levels within and among females.

Testes size, testosterone production and reproductive behaviour in a natural mammalian mating system.

1. Testosterone (T) is a key mediator in the expression of numerous morphological and behavioural traits in mammals, but the factors underlying individual variation in circulating T levels are poorly understood. 2. The intimate structural integration of sperm and T production within the testes, alongside the dependency of sperm production on high levels of T, suggests that T requirements for spermatogenesis could be an important driver of individual differences in T. 3. To test this hypothesis, we examine how male capacity for sperm production (as indicated by their testes size) is associated with T levels in a feral population of Soay sheep, resident on St. Kilda, Scotland, during their rutting season. 4. We found a strong positive relationship between an individual's testes size (as measured before their seasonal enlargement) and the levels of circulating T during their rut, suggesting that T requirements for spermatogenesis has a prominent influence on the production of this androgen. 5. In contrast, body condition and competitive ability did not independently predict T levels, findings that are inconsistent with conventional 'condition-dependent' and 'challenge' hypotheses of T production. 6. This influence of male's capacity for sperm production on T appeared to be substantial enough to be biologically relevant, as testes size also predicted male aggression and mate-seeking behaviour. 7. Our results suggest that a male's inherent capacity for sperm and T production is tightly phenotypically integrated, with potential consequences for a wide range of other T-mediated reproductive traits.

Abnormal reproductive patterns in Przewalski's mares are associated with a loss in gene diversity.

The ex situ population of the Przewalski's horse (Equus ferus przewalskii) is not self-sustaining (20% foaling rate), and the demography is skewed toward aging individuals with low gene diversity. We designed the present study to gain a better understanding of the reproductive biology of the Przewalski's mare and to determine whether age and gene diversity influenced reproductive function. Urine samples were collected 3-7 days/wk from 19 mares from May to September, and ultrasound examinations of follicular structures were performed 3 days/wk for 5 wk from May through July in nine individuals. A high proportion of mares exhibited abnormal (endocrine, 5 [26.3%] of 19; follicular, 2 [22.2%] of 9) or acyclic (endocrine, 4 [21.1%] of 19; follicular, 3 [33.3%] of 9) reproductive patterns. In four cyclic mares, estrous cycle length was 25.1 ± 1.2 days, with 12.2 ± 0.9 days of diestrus. Follicles in cyclic mares grew 1.2 ± 0.6 mm per day and ovulated after reaching 40.4 ± 8.9 mm. Mares with a high coefficient of inbreeding excreted reduced levels of mean urinary estrogens (r(2) = 0.476, P < 0.05), but age had no significant impact on reproductive patterns in this population. Overall, these data suggest that long-term genetic management of this population is necessary to maintain reproductive fitness.

Fecal steroid analysis for evaluating ovarian function in the greater kudu (Tragelaphus strepsiceros) and lesser kudu (Tragelaphus imberbis).

Seasonal reproductive-endocrine norms have not been described for the genus Tragelaphus, which consists of seven species of African antelope. Longitudinal patterns of progesterone metabolite excretion were assessed by radioimmunoassays in fecal samples collected noninvasively (three to seven samples per week) from greater kudu (Tragelaphus strepsiceros, n = 4) and lesser kudu (Tragelaphus imberbis, n = 4). Progesterone metabolite excretion patterns revealed seasonal estrous cycles in both species, and discrimination of pregnant versus nonpregnant females was achieved in lesser kudu. These data reveal the value of fecal progesterone metabolites for establishing reproductive-endocrine norms in both zoo-maintained and free-living antelopes of the genus Tragelaphus.

Rising fecal glucocorticoid concentrations track reproductive activity in the female giant panda (Ailuropoda melanoleuca).

To better understand the adaptive significance of adrenal glucocorticoid (GC) variation in the giant panda, we assessed patterns of fecal GC excretion over time as well as during estrus, parturient and non-parturient luteal phases, lactation and acyclicity in 17 adult females. Fecal estrogen and GC patterns were positively correlated (P<0.05) in four of five periestrual females (r = 0.57-0.92). Among all reproductive states, fecal GC was highest (P<0.05) during periestrus (non-parturient, 495.9 ± 100.7 ng/g [mean ± SE]; parturient, 654.1 ± 10 6.5 ng/g; P>0.05). Concentrations of GC metabolites were lower (P<0.05) during the later stage of the luteal phase in non-parturient (334.8 ± 24.8 ng/g) compared to parturient (470.4 ± 54.0 ng/g) females. Although fecal GC concentrations in cyclic, non-parturient females did not differ (P>0.05) across all seasons, there were seasonal variations (P<0.05) in females that were acyclic and non-lactational. However, the overall lack of difference (P>0.05) in GC values between reproductively cyclic and acyclic females did not support the hypothesis that ovarian acyclicity is due to increased adrenal activity (related or unrelated to physiological stress). Furthermore, GCs may play an important role in the normal endocrine milieu associated with sexual receptivity and late pregnancy. These data demonstrate that both reproductive status and seasonal factors are important modulators of adrenal function in this endangered species.

Social behavior, foraging strategies, and fecal glucocorticoids in female blue monkeys (Cercopithecus mitis): potential fitness benefits of high rank in a forest guenon.

Socioecological theory predicts that aggressive feeding competition is associated with linear dominance hierarchies and reproductive advantages for high-ranking females. Female blue monkeys contest fruits and have a linear dominance hierarchy, yet previous research has shown no evidence that high-ranking females benefit from greater feeding success or fertility. Here, we assess whether individuals differ in fecal glucocorticoid (fGC) excretion and examine proximate determinants of such differences to infer potential fitness correlates of rank, using data collected from two study groups in the Kakamega Forest, Kenya. We found that higher ranking females had preferential access to fruits in both groups, although the behavioral mechanisms leading to this effect varied between groups. Despite a consistent rank difference in feeding on fruits, an overall rank effect on fGCs emerged in only one group; females of this group spent comparatively more time feeding on fruits, fruits accounted for a greater proportion of the diet, and females engaged in more frequent food-related agonism. In addition, more females in this group were lactating during a period of low fruit availability, when rank effects on fGCs were particularly strong. Regardless of fruit availability, among lactating females of both groups higher rank was associated with lower fGC levels, indicating lower energetic stress in higher ranking females when energy demands were particularly high. Individual rates of agonism, a potential psychological stressor, were unrelated to fGCs at all times. After we accounted for rates of agonism and feeding on fruits, females of one group who groomed others more had lower fGCs, suggesting that variable social coping behavior can contribute to fGC variation in some groups. This study provides the first empirical evidence that high-ranking female blue monkeys may obtain fitness benefits from their social status, by gaining priority of access to fruits during critical times in the reproductive cycle.

Endocrine milieu of perioestrus in the giant panda (Ailuropoda melanoleuca), as determined by non-invasive hormone measures.

The aim of the present study was to determine the efficacy of faecal hormonal measures for evaluating ovarian activity in a significant sized cohort of giant pandas during the perioestrual period. Faecal excretion of oestrogen and progestagen metabolites corresponded with urinary patterns and receptive behaviours. Longitudinal assessment of 10 females revealed that, on average, faecal oestrogen concentrations started to rise (P < 0.05) above baseline (baseline mean +/- s.e.m.; 64.7 +/- 6.6 ng g(-1)) 5 days before the preovulatory oestrogen peak (484.6 +/- 126.8 ng g(-1)), which was followed by a gradual descent over 4 days to nadir. Mean faecal progestagen metabolite concentrations increased approximately twofold above baseline (from 186.2 +/- 37.7 to 347.2 +/- 75.7 ng g(-1); P < 0.05) during the 20-day interval after the preovulatory oestrogen surge. Variability within and among females precluded the use of a threshold of oestrogen or progestagen metabolites to predict reproductive status, yet faeces collected 2-3 days per week provided sufficient data to recognise that an individual was in the perioestrual period. Finally, in females that were examined for at least 3 consecutive years, there was an 18-53 day variation in the onset and an 8-13 day variation in the duration of perioestrual behaviour from year to year. In summary, these findings indicate that gonadal hormone profiles associated with the period immediately before, during and after oestrus are accurately revealed by analysis of the fibrous faeces of the giant panda. This approach has potential value for providing point-in-time information on the reproductive status of free-living individuals.

Fecal glucocorticoids as indicators of metabolic stress in female Sykes' monkeys (Cercopithecus mitis albogularis).

Because of their mediating role in the stress response and potential effects on fitness, glucocorticoid (GC) hormones are increasingly used to assess the physiological costs of environmental and behavioral variation among wild vertebrates. Identifying the proximate causes of GC variation, however, is complicated by simultaneous exposure to multiple potentially stressful stimuli. Here, we use data from a partially provisioned social group of Sykes' monkeys to evaluate the effects of potential psychological and metabolic stressors on temporal and individual variation in fecal GC (fGC) excretion among 11 adult females. Despite high rates of agonism over provisioned foods fGCs declined during periods of high provisioning frequency when fruit availability was dominated by neem (Azadirachta indica), an item requiring great feeding effort. Provisioned foods did not prevent fGC increases when availability of the most preferred main fruit item, tamarind (Tamarindus indica), declined drastically. Although rank-related differences in access to provisioned foods and rates of agonism did not lead to an overall effect of rank on fGCs, low-ranking females excreted more fGCs than high-ranking females during a period of high provisioning intensity and low fruit availability. The emergence of this rank effect was associated with elevated feeding effort in all females, a greater access to provisioned items by high-ranking females, and a higher proportion of time spent moving in low-ranking females. Our findings suggest that metabolic stressors were the primary determinants of both temporal and individual variation in fGCs, indicating potential fitness benefits for high-ranking females when food availability is limited.

Unique biphasic progestagen profile in parturient and non-parturient giant pandas (Ailuropoda melanoleuca) as determined by faecal hormone monitoring.

The luteal phase of the giant panda has been exclusively assessed by studying urinary hormone patterns in a very few individuals. To better understand hormonal dynamics of protracted progestagen excretion in this endangered species, we monitored hormonal metabolites in the fibrous faeces of multiple females in the USA and China. Giant pandas that were anoestrual during the breeding season excreted baseline progestagen throughout the year. In contrast, there were two distinctive periods when progestagen excretion increased in females that experienced behavioural oestrus, the first being modest, lasting for 61-122 days, and likely reflecting presumptive ovulation. This increase was far surpassed by a secondary rise in progestagen excretion associated with a rejuvenated luteal capacity or hormone production from an extra-gonadal source. The duration of this 'secondary' rise in progestagen excretion averaged approximately 45 days and terminated in a decline to baseline coincident with parturition or the end of a non-parturient luteal interval. Data revealed that, even with a complex, biphasic progestagen profile, the longitudinal patterns produced by giant pandas were relatively consistent among animals and across years within individuals. However, progestagen excretion patterns throughout this period could not be used to discriminate among non-pregnant, pregnant or pseudopregnant states.

Stress and the costs of extra-territorial movement in a social carnivore.

Costs associated with extra-territorial movement are believed to have favoured the evolution of delayed dispersal and sociality across a range of social vertebrates, but remain surprisingly poorly understood. Here we reveal a novel mechanism that may contribute substantially to the costs of extra-territorial movement: physiological stress. We show that subordinate male meerkats, Suricata suricatta, exhibit markedly elevated faecal glucocorticoid metabolite levels (a non-invasive measure of hypothalamic-pituitary-adrenal axis activity) while conducting extra-territorial prospecting forays. While brief increases in glucocorticoid levels are unlikely to be costly, chronic elevations, arising from prolonged and/or frequent forays, are expected to compromise fitness through their diverse negative effects on health. Our findings strongly suggest that prolonged extra-territorial movements do result in chronic stress, as the high glucocorticoid levels of prospectors do not diminish on longer forays and are no lower among males with greater prospecting experience. A generalized 'stress' of extra-territorial movement may therefore have strengthened selection for delayed dispersal and sociality in this and other species, and favoured the conduct of brief forays over extended periods of floating. Our findings have implications too for understanding the rank-related distribution of physiological stress in animal societies, as extra-territorial movements are often conducted solely by subordinates.

The causes of physiological suppression among female meerkats: a role for subordinate restraint due to the threat of infanticide?

In many animal societies, subordinates exhibit down-regulated reproductive endocrine axes relative to those of dominants, but whether this 'physiological suppression' arises from active interference by dominants or subordinate self-restraint is a matter of debate. Here we investigate the roles that these processes play in precipitating physiological suppression among subordinate female meerkats, Suricata suricatta. We show that, while subordinate females are known to suffer stress-related physiological suppression during periodic temporary evictions by the dominant female, their low estrogen levels while within their groups cannot be readily attributed to chronic stress, as their fecal glucocorticoid metabolite levels during this time are comparable to those of dominants. The low estrogen levels of subordinate females also cannot be explained simply by self-restraint due to factors that could reduce their payoff from maintaining their fertility regardless of the presence of the dominant female (young age, a lack of unrelated mates, poor body condition and limited breeding experience), as substantial rank-related differences in fecal total-estrogen metabolite levels remain when such factors are controlled. We suggest that this residual difference in estrogen levels may reflect a degree of subordinate restraint due in part to the dominant female's ability to kill their young. Accordingly, subordinate female estrogen levels vary in association with temporal variation in the likelihood of infanticide by the dominant. Attempts to identify the causes of physiological suppression should be cautious if rejecting any role for dominant interference in favor of subordinate restraint, as the dominant's capacity to interfere may often be the reason why subordinates exercise restraint.

Fecal steroid monitoring for assessing gonadal and adrenal activity in the golden eagle and peregrine falcon.

We examined the efficacy of noninvasive monitoring of endocrine function via fecal steroid immunoassays in the golden eagle and peregrine falcon. High-pressure liquid chromatography analyses of fecal glucocorticoid metabolites (fGCM) revealed that minor percentages of immunoreactive fGCM co-eluted with [(3)H]corticosterone in both sexes of the eagle (2.5-2.7%) and falcon (7.5-11.9%). In contrast, most fecal estrogen metabolites in eagle and falcon females co-eluted with radiolabeled estradiol-17beta ([(3)H]; 57.6, 64.6%, respectively) or estrone ([(3)H]; 26.9, 4.1%, respectively). Most fecal progestin metabolite immunoreactivity in the female eagle (24.8%) and falcon (21.7%) co-eluted with progesterone ([(14)C]). Most fecal androgen metabolite immunoreactivity in eagle (55.8%) and falcon (63.7%) males co-eluted with testosterone ([(14)C]). Exogenous adrenocorticotropin hormone induced increased fGCM excretion above pre-treatment in both species, but only significantly (P < 0.05) in the eagle. Both species showed increased fGCM after saline administration, suggesting the detection of 'handling stress.' Both species exhibited enterohepatic and renal recirculation of administered steroids as demonstrated by biphasic and triphasic excretion patterns. Thus, noninvasive fecal hormone monitoring is a valid and promising tool for assessing gonadal and adrenal status in rare and threatened birds-of-prey.