Habituation of Adult Magellanic Penguins to Human Visitation as Expressed through Behavior and Corticosterone Secretion

Abstract:  Ecotourism is increasing worldwide; hence, it is important to know how wildlife are affected behaviorally and physiologically by human visitation. We studied the effects of human visitation on the Magellanic Penguins ( Spheniscus magellanicus ) at Punta Tombo, Argentina, by monitoring changes in defensive head turns and plasma corticosterone (a hormone secreted in response to stress) for penguins with and without a history of tourist visitation. Habituation to human visitation was rapid. In penguins with no previous exposure to tourists, the number of defensive head turns and level of plasma corticosterone decreased significantly within 5 days of one 15-minute visit/day. Penguins living in tourist-visited and undisturbed areas secreted more corticosterone when captured and restrained than penguins visited by a person. Penguins in tourist areas, however, did not show as strong a corticosterone response to capture and restraint as did penguins in areas without tourists. This difference was due to a decreased capability of the adrenocortical tissue to secrete corticosterone in tourist-visited birds. Although our data show no direct negative effects of tourism on Magellanic Penguins at Punta Tombo, consequences of a modification of physiological capabilities (e.g., adrenocortical function) may not become apparent until much later in life. The physiological differences between tourist-visited and undisturbed groups of Magellanic Penguins emphasize the importance of monitoring the effects of anthropogenic disturbances on wildlife at multiple levels.     

Foraging behaviour and reproductive success in Magellanic penguins (Spheniscus magellanicus): a comparative study of two colonies in southern Chile

Marine Biology - During the breeding season 1996/97 we compared the foraging and diving behaviour of adult Magellanic penguins (Spheniscus magellanicus), growth rates of their chicks and their...     

Diving behaviour of Little Penguins from four colonies across their whole distribution range: bathymetry affecting diving effort and fledging success

Little Penguins, Eudyptula minor, breed in several small colonies in New Zealand and Australia. In this study, we compare the birds’ diving performances at different sites situated throughout their breeding range. Environmental conditions and breeding success vary drastically amongst colonies, but all birds feed on similar types of prey and face similar limitations on their foraging range. We examined several diving parameters and calculated the proportion of foraging zone available during breeding to examine whether oceanographic and geographic factors in the foraging zone can explain variations in diving behaviour and fledging success among the different colonies. In colonies with high fledging success, Penguin Island and Oamaru, penguins made shallow dives <50 m depth and had lower diving effort. More than 90% of the foraging zone was in waters <50 m depth in these colonies. Motuara Island also has shallow waters with 95% <50 m depth, but the fledging success was low. Phillip Island has only 42% of waters <50 m and comparatively low fledging success. Thus, penguins dived deeper and showed a higher diving effort in colonies with lower fledging success (Motuara Island and Phillip Island), indicating that they were disadvantaged compared to conspecifics from other colonies that dived shallower and with a lesser diving effort. We concluded that bathymetry is an important factor, but not the only one, which influences fledging success.     

Transition to independence and evidence of extended parental care in the gentoo penguin (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>)

We used radio telemetry and observations to study the activity patterns and behavior of gentoo penguin chicks at Admiralty Bay, King George Island, South Shetland Islands in 2005 during their “fledging period”; defined as the time between a chick’s first trip to sea and its final dispersal from the breeding colony. Gentoo penguins exhibited delayed dispersal of young and extended parental provisioning, behaviors not observed in other Pygoscelis species. Chicks took their first trip to sea at a mean age of 70 days of age, before finally departing the colony at a mean age of 82 days. During this fledging period, individual chicks made an average of five trips to sea. Trip duration increased significantly as chicks aged, with trips to sea becoming similar to literature values of adult foraging trips in both timing and duration. Behavioral observations and mass dynamics confirmed that many chicks were still being fed during this fledging period, with parental feeding behaviors most often observed in the late afternoon to evening hours. We hypothesize that these behaviors provide an opportunity for chicks to gain experience at sea prior to dispersal and might allow them to develop foraging skills before they are completely independent.     

Age,experience and reproductive performance in a long-lived bird: a hormonal perspective

The ultimate factors influencing age-specific reproductive performance in birds have been widely discussed, and several hypotheses have been suggested to explain why young/inexperienced breeders have lower reproductive success than older individuals. In comparison, proximate factors and, particularly, hormonal mechanisms influencing age-related reproductive performance have received lesser attention. In this paper, we examined how baseline levels of corticosterone and prolactin, two hormones involved in reproduction, changed with age and experience in a long-lived bird, the Black-browed albatross (Thallasarche melanophris) during the brooding stage. Corticosterone levels were not linked to age, whereas prolactin levels increased until individuals reached 15 years of age. First-time breeders had higher corticosterone levels and lower prolactin levels than experienced ones. Corticosterone levels were not correlated with breeding experience among experienced birds, whereas prolactin levels slightly increased with advancing experience. Among experienced breeders, there was no effect of individual quality on corticosterone and prolactin levels. Baseline corticosterone and prolactin levels were respectively, positively and negatively correlated to time spent fasting/brooding on the nest. Moreover, the probability of successfully fledging a chick was negatively related to corticosterone levels, but not to prolactin levels. Elevated corticosterone levels in first time breeders may serve as evidence for depleted body reserves resulting from lower foraging/brooding capabilities and therefore support the constraint hypothesis. Low prolactin levels in young/inexperienced birds may be interpreted either as evidence for their lower breeding capacities (constraint hypothesis) or for their limited breeding investment (restraint hypothesis). Finally, we report, for the first time, the hormonal changes associated with the onset of senescence. The very old and most experienced birds, which had the lowest probability of successfully fledging a young, displayed elevated corticosterone levels and low prolactin levels, possibly indicating a degradation of breeding skills and/or a disruption of the endocrine system in senescent birds.     

Value of Artificial Habitats for Amphibian Reproduction in Altered Landscapes

Abstract:  Installation and maintenance of stormwater ponds to detain and treat runoff from impervious surfaces is a common method of stormwater control in developed areas. That these ponds capture pollutants, however, is of concern for wildlife species that use the ponds, particularly pond-breeding amphibians. To assess the relative contribution of stormwater ponds to the persistence of amphibian populations in suburban landscapes, we compared amphibian use of stormwater ponds and other available wetlands in suburban and forested watersheds. We surveyed three suburban and three primarily forested first-order watersheds to identify all potential wetlands that might serve as breeding sites for pond-breeding amphibians. We performed call, egg-mass, and larval surveys to measure breeding effort at each wetland in spring and summer 2007 and 2008. In suburban watersheds most (89%) of the wetlands that had breeding activity were either stormwater ponds or otherwise artificial. This pattern was also evident in the forested watersheds, where amphibians were primarily found breeding in wetlands created by past human activity. Late-stage larvae were found only in anthropogenic wetlands in all study areas because the remaining natural wetlands did not hold water long enough for larvae to complete development. Our results suggest that in urban and suburban landscapes with naturally low densities of wetlands, wetlands created by current or historic land uses may be as important to amphibian conservation as natural wetlands or pools and that management strategies directed at urban and suburban landscapes should recognize and incorporate human-created wetlands.     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

A strategy for wildlife management in depopulating rural areas of Japan

Former ranges of wild animals have been reestablished in many developed countries. However, this reestablishment has led to increasing human–wildlife conflict in agroforest ecosystems. In Japan, human–wildlife conflict, such as crop raiding by and ecological impacts of wild ungulates and primates, is a serious problem in depopulated rural areas due to these animal range expansions and increased abundances. Japan's human population is predicted to decline by 24% by 2050, and approximately 20% of agricultural settlements will become completely depopulated. In this scenario, anthropogenic pressures on wildlife (e.g., hunting and habitat alteration) will continue to decrease and human–wildlife conflict will increase due to increasing wildlife recovery. Japan's local governments plan to slow range recovery, prevent species reestablishment, or remove recolonizing large mammals through lethal control. This strategy, however, is not cost-effective, and workforce shortages in depopulated communities make it infeasible. Moreover, the suppression of wildlife prevents the recovery of ecological functions and thus would degrade regional biodiversity. The declining pressure on wildlife that accompanies human depopulation will prevent the restoration of any past states of human–wildlife interaction. We suggest human-used areas in rural landscapes be aggregated in compact cities and that in transition zones between human settlements and depopulated lands that land-sharing approaches be applied. Concentrating management efforts in compact cities may effectively decrease human–wildlife conflict, rather than intensifying human pressures. Reforestation of depopulated lands may lead to recovery of wildlife habitats, their ecosystem functions, and regional biodiversity due to minimization of negative anthropogenic effects (land-sparing approach). Balancing resolution of human–wildlife conflict and ecological rewilding could become a new, challenging task for regional wildlife managers.     

Costs and benefits of brood desertion in female kentish plovers,Charadrius alexandrinus

Female kentish plovers Charadrius alexandrinus typically desert their broods after the chicks hatch, i.e. 1–4 weeks before the chicks fledge or become independent. In this paper we investigate the costs and benefits of desertion for females. Desertion incurs a cost for females: following desertion chick survival in broods is lower (0.95 +- 0.02 day^–1) than before the female deserts (0.98 +- 0.01 day^–1). We investigated several possible causes for reduced brood survival by comparing characteristics of broods before and after desertion (controlling for differences in brood age). After desertion males increased the time they spent foraging and they tended to reduce time spent brooding chicks. Increased mortality of chicks may occur in deserted broods because following desertion (1) males spend less time alert in vigilance behaviour than before desertion, (2) they attend the chicks from greater distances, and (3) they show greater distraction display distances (in response to human intruders). Growth or development of chicks, measured by weight gain and tarsus length, was not different before and after desertion. Females gain two potential benefits from desertion: (1) they may remate and produce a second brood within the same breeding season or (2) they may enhance their probability of surviving to breed in a subsequent season. At least 27% of female kentish plovers that deserted remated and renested in the same season in this study. In contrast, we found no evidence that brood desertion increased the survival of females: there was no difference in local survival rate (return rate) for females deserting before or after 6 days brood age. These results clearly demonstrate that female kentish plovers that desert their offspring prior to fledging incur costs, but we suggest that there is a trade-off with the potential benefits gained by remating and making a second breeding attempt in the same season.     

Effects of anthropogenic developments on common raven nesting biology in the West Mojave Desert.

Subsidized predators may affect prey abundance, distribution, and demography. Common Ravens (Corvus corax) are anthropogenically subsidized throughout their range and, in the Mojave Desert, have increased in number dramatically over the last 3-4 decades. Human-provided food resources are thought to be important drivers of raven population growth, but human developments add other features as well, such as nesting platforms. From 1996 to 2000, we examined the nesting ecology of ravens in the Mojave Desert, relative to anthropogenic developrhent. Ravens nested disproportionately near point sources of food and water subsidies (such as towns, landfills, and ponds) but not near roads (sources of road-killed carrion), even though both sources of subsidy enhanced fledging success. Initiation of breeding activity was more likely when a nest from the previous year was present at the start of a breeding season but was not affected by access to food. The relative effect of environmental modifications on fledging success varied from year to year, but the effect of access to human-provided resources was comparatively consistent, suggesting that humans provide consistently high-quality breeding habitat for ravens. Anthropogenic land cover types in the desert are expected to promote raven population growth and to allow ravens to occupy parts of the desert that otherwise would not support them. Predatory impacts of ravens in the Mojave Desert can therefore be considered indirect effects of anthropogenic development.     

Conservation outside Protected Areas and the Effect of Human‐Dominated Landscapes on Stress Hormones in Savannah Elephants

Biodiversity conservation strategies are increasingly focused on regions outside national protected areas, where animals face numerous anthropogenic threats and must coexist with human settlements, livestock, and agriculture. The effects of these potential threats are not always clear, but they could have profound implications for population viability. We used savannah elephants (Loxodonta africana) as a case study to assess the physiological stress associated with living in a human‐livestock‐dominated landscape. We collected samples over two 3‐month periods in 2007 and 2008. We used fecal DNA to identify 96 individual elephants in a community conservation area (CCA) and measured fecal glucocorticoid metabolite (FGM) concentrations as a proxy for stress. The CCA is community Maasai land managed for livestock and wildlife. We compared the FGM concentrations from the CCA to FGM concentrations of 40 elephants in Amboseli National Park and 32 elephants in the Maasai Mara National Reserve, where human settlements and intense livestock grazing were absent. In the CCA, we found no significant individual differences in FGM concentrations among the elephants in 2007 (p = 0.312) or 2008 (p = 0.412) and no difference between years (p = 0.616). The elephants in the CCA had similar FGM concentrations to the Maasai Mara population, but Amboseli elephants had significantly lower FGM concentrations than those in either Maasai Mara or the CCA (Tukey pairwise test, p < 0.001), due primarily to females excreting significantly lower FGM relative to males (p = 0.025). In the CCA, there was no relation among female group size, average pairwise group relatedness, and average group FGM concentration. We found no clear evidence of chronic stress in elephants living on CCA communal land, which is encouraging for conservation strategies promoting the protection of animals living outside protected areas. Conservación Fuera de Áreas Protegidas y el Efecto de Paisajes Dominados por Humanos sobre Hormonas del Estrés en Elefantes Africanos     

Augmenting Small Populations of Plovers: An Assessment of Cross-Fostering and Captive-Rearing

This study compared the growth and behavioral development of parent-reared, cross-fostered, and captive-reared Killdeer ( Charadrius vociferus ) chicks. Common species were used to test these techniques for possible application to rare endangered forms. Parent-reared chicks were raised naturally in the wild, cross-fostered chicks were raised by Spotted Sandpipers ( Actitis macularia ) in the wild, and captive-reared chicks were raised in captivity by humans. Both hatching and fledging success were significantly increased by captive-rearing, and cross-fostering produced approximately the same number of fledged young as natural parent-rearing. Captive-reared Killdeer chicks spent more time resting and less time feeding and stayed closer to siblings than cross-fostered or parent-reared chicks; these behavioral differences were not seen after release to the wild. Growth rates among the three groups were similar. All of the young Killdeer responded to Killdeer alarm calls. There was no evidence that captive-reared and cross-fostered Killdeer were negatively affected by their early experiences. Captive-rearing is a viable management technique for augmenting small populations of endangered shorebirds, such as the Piping Plover ( Charadrius melodus ). It is recommended over cross-fostering because captive-rearing is more flexible as a technique, produces more young, does not affect another species, and does not produce potential imprinting problems.     

Cuckoo growth performance in parasitized and unused hosts: not only host size matters

The quality and quantity of food delivered to young are among the major determinants of fitness. A parental provisioning capacity is known to increase with body size. Therefore, brood parasitism provides an opportunity to test the effects of varying provisioning abilities of different-sized hosts on parasitic chick growth and fledging success. Knowledge of growth patterns of common cuckoo, Cuculus canorus, chicks in nests of common hosts is very poor. Moreover, no study to date has focused on any currently unused hosts (i.e., suitable cuckoo host species in which parasitism is currently rare or absent). Here, I compare the growth performance of cuckoo chicks in nests of a common host (the reed warbler, Acrocephalus scirpaceus) and two unparasitized hosts (the song thrush, Turdus philomelos, and the blackbird, Turdus merula). Parasitic chicks were sole occupants of the observed nests, thus eliminating the confounding effect of competition with host chicks. Experiments revealed striking differences in parasitic chick growth in the two closely related Turdus hosts. Cuckoo chicks cross-fostered to song thrush nests grew much quicker and attained much higher mass at fledging than those in nests of their common reed warbler host. Alternatively, parasitic chicks in blackbird nests grew poorly and did not survive until fledging. I discuss these observations with respect to host selection by parasitic cuckoos.     

The effects of nesting stage,sex, and type of predator on parental defense by killdeer (Charadrius vociferous): testing models of avian parental defense

Summary Two models predicting the temporal patterns of parental investment in offspring defense over the nesting cycle were tested. The first is based on offspring age, the other on the vulnerability of offspring to predation. Both models make very similar predictions for altricial species after eggs have hatched, i.e., increases in intensity of parental defense until fledging. For precocial species, however, the post-hatching predictions of each model are different: the offspring age model predicts a continued increase in defense intensity, while the vulnerability model predicts a decline. I examined the temporal patterns of parental defense of a precocial shorebird, the killdeer (Charadrius vociferus), and determined which model was supported. Killdeer responses to human and natural predators were observed. Killdeer were less willing to leave the nest, responded most intensely, and displayed closest to a potential predator around hatching. Defense intensity increased from early to late incubation as predicted by the offspring age model. However, after hatching killdeer parental defense declined for both males and females, thus supporting the vulnerability model for this stage. Males and females responded significantly differently to all types of predators. Males took greater risks, remained on the nest longer, defended offspring more intensely, and displayed closer to the predator than females at the approach of a potential predator. Responses to natural predators depended on the type of predator and the approach made by the predator; a greater range of defense behavior was used for predators approaching on the ground compared to aerial predators. In general, killdeer responses to humans were more intense and less variable than their responses to natural predators. This was most likely because the human intruder approached nests and chicks more directly and closer than natural predators.     

Seasonal decline of offspring quality in the European starling Sturnus vulgaris: an immune challenge experiment

In seasonally fluctuating environments, timing of reproduction is a crucial determinant of fitness. Studies of birds show that late breeding attempts generally result in offspring of lower reproductive value, with lower recruitment and long-term survival prospects. Several proximate mechanisms, including a seasonal decline of immune system functioning, may lead to a seasonal decline of offspring fitness. We investigated seasonal variation in offspring quality by subjecting first- and second-brood chicks of a sexually size dimorphic species, the European starling Sturnus vulgaris, to an immune challenge with a bacterial endotoxin (LPS), and evaluated their growth and physiological response in terms of total plasma antioxidant capacity (TAC), concentration of reactive oxygen metabolites and hematocrit. LPS challenge did not affect chick growth or oxidative status. However, hematocrit of second-brood chicks was higher in LPS chicks compared to controls. Body mass halfway through the rearing period (days?8–9 post-hatching), TAC and hematocrit were lower among second- vs. first-brood chicks. Interestingly, sexual dimorphism in body mass at days?8–9 post-hatching markedly differed between broods, first-brood males being 4.7% and second-brood males 22.7% heavier than their sisters, respectively. Pre-fledging mortality occurred among second-brood chicks only and was strongly female-biased. Our findings suggest that starling chicks, even if in poor conditions, are little affected by a bacterial challenge, at least in the short-term. Moreover, our study indicates that sex differences in body size, possibly mediated by sex-specific maternal investment in egg size, may heavily impact on pre-fledging survival in a different way in the course of the breeding season, resulting in sex-specific seasonal decline of offspring fitness. Finally, we suggest that levels of circulating antioxidants should be regarded among the proximate causes of the association between timing of fledging and long-term survival in avian species.     

Stress and the city: urbanization and its effects on the stress physiology in European blackbirds

Animals colonizing cities are exposed to many novel and potentially stressful situations. There is evidence that chronic stress can cause deleterious effects. Hence, wild animals would suffer from city life unless they adjusted their stress response to the conditions in a city. Here we show that European Blackbirds born in a city have a lower stress response than their forest conspecifics. We hand-raised urban and forest-living individuals of that species under identical conditions and tested their corticosterone stress response at an age of 5, 8, and 11 months. The results suggest that the difference is genetically determined, although early developmental effects cannot be excluded. Either way, the results support the idea that urbanization creates a shift in coping styles by changing the stress physiology of animals. The reduced stress response could be ubiquitous and, presumably, necessary for all animals that thrive in ecosystems exposed to frequent anthropogenic disturbances, such as those in urban areas.     

Human disturbance and shifts in vertebrate community composition in a biodiversity hotspot

Understanding how human modification of the landscape shapes vertebrate community composition is vital to understanding the current status and future trajectory of wildlife. Using a participatory approach, we deployed the largest camera-trap network in Mesoamerica to date to investigate how anthropogenic disturbance shapes the occupancy and co-occurrence of terrestrial vertebrate species in a tropical biodiversity hotspot: the Osa Peninsula, Costa Rica. We estimated species richness in different categories of land protection with rarefaction analysis and estimated the expected occupancy with a joint species distribution model that included covariates for anthropogenic disturbance, land protection, habitat quality, and habitat availability. Areas with the most stringent land-use protections (e.g., Corcovado National Park, 24 species [95% CI 23–25]) harbored significantly more species than unprotected areas (20 species [19.7–20.3]), mainly due to a reduced presence of large-bodied species of conservation concern in unprotected areas (e.g., jaguar Panthera onca and white-lipped peccary Tayassu pecari). Small-bodied generalist species, such as opossums (Didelphidae) and armadillos (Dasypus novemcinctus), in contrast, were more common at disturbed sites, resulting in a significant difference in vertebrate community composition between sites with low and high disturbance. Co-occurrence of species was also mainly associated with response to disturbance. Similar responses to disturbance create two groups of species, those whose site-level occupancy usually increased as anthropogenic disturbance increased and those whose estimated occupancy decreased. The absence of large-bodied species entails an important loss of ecological function in disturbed areas and can hinder forest development and maintenance. Efforts to protect and restore forested landscapes are likely having a positive effect on the abundance of some threatened species. These efforts, however, must be sustained and expanded to increase connectivity and ensure the long-term viability of the wildlife community.     

Poaching Risks in Community‐Based Natural Resource Management

Poaching can disrupt wildlife‐management efforts in community‐based natural resource management systems. Monitoring, estimating, and acquiring data on poaching is difficult. We used local‐stakeholder knowledge and poaching records to rank and map the risk of poaching incidents in 2 areas where natural resources are managed by community members in Caprivi, Namibia. We mapped local stakeholder perceptions of the risk of poaching, risk of wildlife damage to livelihoods, and wildlife distribution and compared these maps with spatially explicit records of poaching events. Recorded poaching events and stakeholder perceptions of where poaching occurred were not spatially correlated. However, the locations of documented poaching events were spatially correlated with areas that stakeholders perceived wildlife as a threat to their livelihoods. This result suggests poaching occurred in response to wildlife damage occurred. Local stakeholders thought that wildlife populations were at high risk of being poached and that poaching occurred where there was abundant wildlife. These findings suggest stakeholders were concerned about wildlife resources in their community and indicate a need for integrated and continued monitoring of poaching activities and further interventions at the wildlife‐agricultural interface. Involving stakeholders in the assessment of poaching risks promotes their participation in local conservation efforts, a central tenet of community‐based management. We considered stakeholders poaching informants, rather than suspects, and our technique was spatially explicit. Different strategies to reduce poaching are likely needed in different areas. For example, interventions that reduce human‐wildlife conflict may be required in residential areas, and increased and targeted patrolling may be required in more remote areas. Stakeholder‐generated maps of human‐wildlife interactions may be a valuable enforcement and intervention support tool. Riesgos de Cacería Furtiva en el Manejo de Recursos Naturales Basado en Comunidades     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

Behavioral responses of bison and elk in Yellowstone to snowmobiles and snow coaches.

Managers of public lands are charged with protecting some of our most important natural resources and ecosystems, while providing for their use and enjoyment by visitors. Almost one million visitors entered Yellowstone National Park by motorized means on snowmobiles (87%) or snow coaches (13%) during 1992-2003. Most vehicles toured the central portion of the park where bison (Bison bison) and elk (Cervus elaphus) concentrate in geothermal areas. We sampled >6500 interactions between groups of these species and groups of snowmobiles and snow coaches (collectively, OSV, over-snow vehicles) during five winters (1999-2000, 2002-2004). Multinomial logits models were used to identify conditions leading to behavioral responses. Elk responded three times as often (52%) as bison (19%) during interactions with groups of snowmobiles and snow coaches due to increased vigilance responses (elk, 44%; bison, 10%). However, the frequency of higher-intensity movement responses by bison and elk were similar (6-7% travel, 1-2% flight, <1% defense) and relatively low compared to other studies of ungulates and snowmobile disturbance. The likelihood of active responses by bison and elk increased significantly if animals were on or near roads, groups were smaller, or humans approached. The likelihood of an active response by bison decreased within winters having the largest visitation, suggesting some habituation to snowmobiles and snow coaches. There was no evidence that snowmobile use during the past 35 years affected the population dynamics or demography of bison or elk. Thus, we suggest that regulations restricting levels and travel routes of over-snow vehicles (OSVs) were effective at reducing disturbances to bison and elk below a level that would cause measurable fitness effects. We recommend park managers consider maintaining OSV traffic levels at or below those observed during our study. Regardless, differing interpretations of the behavioral and physiological response data will continue to exist because of the diverse values and beliefs of the many constituencies of Yellowstone.