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At least 19 recordsLinked to original sources

A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA

DNA fingerprinting and mitochondrial DNA analyses have not been used in combination to study relatedness in natural populations. We present an approach that involves defining the mean fingerprint similarities among individuals thought to be unrelated because they have different mtDNA genotypes. Two classes of related individuals are identified by their distance in standard errors above this mean value. The number of standard errors is determined by analysis of the association between fingerprint similarity and relatedness in a population with a known genealogy. We apply this approach to gray wolf packs from Minnesota, Alaska, and the Northwest Territories. Our results show that: (1) wolf packs consist primarily of individuals that are closely related genetically, but some packs contain unrelated, non-reproducing individuals; (2) dispersal among packs within the same area is common; and (3) short-range dispersal appears more common for female than male wolves. The first two of these genetically-based observations are consistent with behavioral data on pack structure and dispersal in wolves, while the apparent sex bias in dispersal was not expected.

Behavioral Ecology and Sociobiology↗

Sex differences in the ontogeny of social behavior in pikas: possible relationships to dispersal and territoriality

A laboratory study on the ontogeny of social behavior in pikas (Ochotona princeps), an alpine lagomorph, was conducted to determine the role of early relationships between adult females and young and among siblings in the development of territorial and dispersal behaviors. Sex differences during development were examined because field studies have reported greater dispersal distances in young females than young males. At birth, females were significantly heavier than males. There were no sex differences in nursing frequency until after the 2nd week of age, when males initiated more nursing attempts than females. By the end of the weaning period (weeks 5 and 6), adult females became non-interactive with young, but aggression of young toward littermates and the mother increased until the eighth week. At this time, young males outweighed their sibling females. Young were dominant over their mothers by the age of 5 weeks, and young males were dominant over their sibling females. Sex differences were observed in aggression, scent-marking, exploratory activity, and submissive vocalizations, with higher rates in young males, except for submissive vocalizations, which were higher in females (Table 2). Vocalizations and scent-marking behavior increased over time, and were positively correlated with interaction rates. These data support the hypothesis that female young disperse farther than male young largely as the result of unsuccessful competition with male siblings for available territories close to the birthplace. A dispersal strategy for pikas is proposed.

Behavioral Ecology and Sociobiology↗

Pack size and wolf pup survival: their relationship under varying ecological conditions

The relationship between pack size and two parameters of reproductive success (litter size at 7–8 months and pup weights at 5–6 months) were determined for two wolf ( Canis lupus ) populations in northern Minnesota. Pup weights were not correlated with pack size for either population. Litter size, however, was correlated with pack size, but the direction of the relationship varied between the two study populations. In the superior National Forest, where prey were scarce and the wolf population was declining from high densities, litter size and pack size were inversely related. Pairs produced more surviving pups than did larger packs with one or more potential helpers. In the Beltrami Island State Forest, where prey were relatively abundant and the wolf population was increasing, pack size and litter size were positively correlated. The results suggest that ecological factors, such as prey availability, affected the ability or willingness of various pack members to provide food or other care for the pups. The lack of correlation between number of auxiliaries and number of pups in canid populations with low and declining prey densities may be explained on the basis of heterogeneous prey density resulting in drastic annual variation in litter production. No study to date has measured the actual benefit that pups derive from helping by auxiliaries, and the costs and benefits of it. The relationships discussed herein can be considered valid only after such research is completed.

Behavioral Ecology and Sociobiology↗

The relationship between parental age and reproductive effort in the California Gull (Larus californicus)

Reproductive effort was compared among California gulls ranging in age from 3 to 18 years old. Results indicated that reproductive effort increased with parental age. Older parents (11–18 years old) invested more in foraging effort and in defense of offspring than did younger parents (<10 years old), and did so over a greater period of time. Increases with age in the efficiency of parenting could not be demonstrated. Increased levels of reproductive effort, therefore, appeared to be the main factor responsible for age-related increases in breeding success. The higher levels of reproductive effort of old parents were associated with higher mortality. Thus, data support the prediction that a cost-benefit system of risk of mortality vs reproductive gain operates in this population. Higher levels of reproductive effort can increase an individual's reproductive success; however, it also increases the likelihood that it will die. Therefore, for the young parent, high risks of mortality involved in current production would be too costly (in terms of loss of future reproductive success) to justify the benefit of high current reproductive output. Parents achieve the highest likelihood of maximizing life-time reproductive output by increasing reproductive effort with age as the cost of mortality diminishes. Data testing the relationship between parental age and reproductive effort demonstrate the existence of age-related variation in reproductive investment. They also demonstrate that the concept of reproductive value proposed by Fisher (1930) has validity as a framework which supports much of our theory on life history evolution.

Behavioral Ecology and Sociobiology↗

Wolf pack spacing: Howling as a territory-independent spacing mechanism in a territorial population

Howling is a principle means of spacing in wolf populations. The relationship between a pack's responses to howling (replies, movements) and its location within its home range, was studied using human-simulated howling in a territorial population in northeastern Minnesota. The results indicated the responses were independent of the pack's location, or the locations of the pack and playback relation to the territory center. These results indicate that howling serves as a territory-independent spacing mechanism, that will result in the use of exclusive territories when coupled with strong, year-round site attachment, but with floating, exclusive, buffer-areas about migratory packs.

Behavioral Ecology and Sociobiology↗

Can non-breeding be a cost of breeding dispersal?

Breeding habitat selection and dispersal are crucial processes that affect many components of fitness. Breeding dispersal entails costs, one of which has been neglected: dispersing animals may miss breeding opportunities because breeding dispersal requires finding a new nesting site and mate, two time- and energy-consuming activities. Dispersers are expected to be prone to non-breeding. We used the kittiwake (Rissa tridactyla) to test whether breeding dispersal influences breeding probability. Breeding probability was associated with dispersal, in that both were negatively influenced by private information (previous individual reproductive success) and public information (average reproductive success of conspecifics) about patch quality. Furthermore, the probability of skipping breeding was 1.7 times higher in birds that settled in a new patch relative to those that remained on the same patch. Finally, non-breeders that resumed breeding were 4.4 times more likely to disperse than birds that bred in successive years. Although private information may influence breeding probability directly, the link between breeding probability and public information may be indirect, through the influence of public information on breeding dispersal, non-breeding thus being a cost of dispersal. These results support the hypothesis that dispersal may result in not being able to breed. More generally, non-breeding (which can be interpreted as an extreme form of breeding failure) may reveal costs of various previous activities. Because monitoring the non-breeding portion of a population is difficult, non-breeders have been neglected in many studies of reproduction trade-offs.

Behavioral Ecology and Sociobiology↗

Conflict of interest between a nematode and a trematode in an amphipod host: Test of the "sabotage" hypothesis

Microphallus papillorobustus is a manipulative trematode that induces strong behavioural alterations in the gamaridean amphipod Gammarus insensibilis , making the amphipod more vulnerable to predation by aquatic birds (definitive hosts). Conversely, the sympatric nematode Gammarinema gammari uses Gammarus insensibilis as a habitat and a source of nutrition. We investigated the conflict of interest between these two parasite species by studying the consequences of mixed infection on amphipod behaviour associated with the trematode. In the field, some amphipods infected by the trematode did not display the altered behaviour. These normal amphipods also had more nematodes, suggesting that the nematode overpowered the manipulation of the trematode, a strategy that would prolong the nematode's life. We hypothesize that sabotage of the trematode by the nematode would be an adaptive strategy for the nematode consistent with recent speculation about co-operation and conflict in manipulative parasites. A behavioural test conducted in the laboratory from naturally infected amphipods yielded the same result. However, exposing amphipods to nematodes did not negate or decrease the manipulation exerted by the trematode. Similarly, experimental elimination of nematodes from amphipods did not permit trematodes to manipulate behaviour. These experimental data do not support the hypothesis that the negative association between nematodes and manipulation by the trematode is a result of the "sabotage" hypothesis.

Behavioral Ecology and Sociobiology↗

Current versus future reproduction: an experimental test of parental investment decisions using nest desertion by mallards (Anas platyrhynchos)

Past investment in offspring may be important in determining a parent's ability to reproduce in the future and, hence, should affect the relative value of current offspring. However, there have been surprisingly few clear tests of whether animals actually adjust parental care in response to diminished opportunities for future reproduction. We modified the experimental protocol of Sargent and Gross [Behav Ecol Sociobiol (1985) 17:43–45] to examine offspring desertion by mallards ( Anas platyrhynchos ), and decoupled the influence of past investment from expected current benefits by controlling for the effect of offspring age on clutch value. Using 9 years of nest mortality data, we accounted for the increasing prospects of egg survival with clutch age by calculating clutch sizes throughout incubation with equivalent expected benefits. Applying this approach, we experimentally reduced 203 clutches at two different incubation stages such that they had equivalent expected benefits but differed in the amount of past investment. Nest desertion rates did not differ between early- and late-incubated clutches that had equivalent expected benefits. Rather, the probability of desertion increased with the severity of the clutch reduction treatment. These results suggest that female mallards adjust parental care according to the expected benefits of current offspring, rather than to diminished prospects for future reproduction due to past investment. We further examined whether females assessed expected benefits on the basis of clutch size alone or clutch size adjusted for the age of the clutch. Using Akaike's Information Criterion, the most parsimonious model to explain the probability of deserting an experimentally reduced clutch included both the proportion of the clutch remaining and clutch age. Thus, female mallards appear to fine-tune their level of parental care not only according to the relative number of offspring in the clutch, but also to the increased prospects for offspring survival as they age.

Behavioral Ecology and Sociobiology↗

Wildlife contact analysis: Emerging methods, questions, and challenges

Recent technological advances, such as proximity loggers, allow researchers to collect complete interaction histories, day and night, among sampled individuals over several months to years. Social network analyses are an obvious approach to analyzing interaction data because of their flexibility for fitting many different social structures as well as the ability to assess both direct contacts and indirect associations via intermediaries. For many network properties, however, it is not clear whether estimates based upon a sample of the network are reflective of the entire network. In wildlife applications, networks may be poorly sampled and boundary effects will be common. We present an alternative approach that utilizes a hierarchical modeling framework to assess the individual, dyadic, and environmental factors contributing to variation in the interaction rates and allows us to estimate the underlying process variation in each. In a disease control context, this approach will allow managers to focus efforts on those types of individuals and environments that contribute the most toward super-spreading events. We account for the sampling distribution of proximity loggers and the non-independence of contacts among groups by only using contact data within a group during days when the group membership of proximity loggers was known. This allows us to separate the two mechanisms responsible for a pair not contacting one another: they were not in the same group or they were in the same group but did not come within the specified contact distance. We illustrate our approach with an example dataset of female elk from northwestern Wyoming and conclude with a number of important future research directions.

Behavioral Ecology and Sociobiology↗

Sea lamprey orient toward a source of a synthesized pheromone using odor-conditioned rheotaxis

Characterization of vertebrate chemo-orientation strategies over long distances is difficult because it is often not feasible to conduct highly controlled hypothesis-based experiments in natural environments. To overcome the challenge, we couple in-stream behavioral observations of female sea lampreys ( Petromyzon marinus ) orienting to plumes of a synthesized mating pheromone, 7a,12a,24-trihydroxy-5a-cholan-3-one-24-sulfate (3kPZS), and engineering algorithms to systematically test chemo-orientation hypotheses. In-stream field observations and simulated movements of female sea lampreys according to control algorithms support that odor-conditioned rheotaxis is a component of the mechanism used to track plumes of 3kPZS over hundreds of meters in flowing water. Simulated movements of female sea lampreys do not support that rheotaxis or klinotaxis alone is sufficient to enable the movement patterns displayed by females in locating 3kPZS sources in the experimental stream. Odor-conditioned rheotaxis may not only be effective at small spatial scales as previous described in crustaceans, but may also be effectively used by fishes over hundreds of meters. These results may prove useful for developing management strategies for the control of invasive species that exploit the odor-conditioned tracking behavior and for developing biologically inspired navigation strategies for robotic fish.

Behavioral Ecology and Sociobiology↗

Factors influencing the movement biology of migrant songbirds confronted with an ecological barrier

Whether or not a migratory songbird embarks on a long-distance flight across an ecological barrier is likely a response to a number of endogenous and exogenous factors. During autumn 2008 and 2009, we used automated radio tracking to investigate how energetic condition, age, and weather influenced the departure timing and direction of Swainson&rsquo;s thrushes ( Catharus ustulatus ) during migratory stopover along the northern coast of the Gulf of Mexico. Most birds left within 1 h after sunset on the evening following capture. Those birds that departed later on the first night or remained longer than 1 day were lean. Birds that carried fat loads sufficient to cross the Gulf of Mexico generally departed in a seasonally appropriate southerly direction, whereas lean birds nearly always flew inland in a northerly direction. We did not detect an effect of age or weather on departures. The decision by lean birds to reorient movement inland may reflect the suitability of the coastal stopover site for deposition of fuel stores and the motivation to seek food among more extensive forested habitat away from the barrier.

Behavioral Ecology and Sociobiology↗

Inferring social structure and its drivers from refuge use in the desert tortoise, a relatively solitary species

For several species, refuges (such as burrows, dens, roosts, nests) are an essential resource for protection from predators and extreme environmental conditions. Refuges also serve as focal sites for social interactions, including mating, courtship, and aggression. Knowledge of refuge use patterns can therefore provide information about social structure, mating, and foraging success, as well as the robustness and health of wildlife populations, especially for species considered to be relatively solitary. In this study, we construct networks of burrow use to infer social associations in a threatened wildlife species typically considered solitary—the desert tortoise. We show that tortoise social networks are significantly different than null networks of random associations, and have moderate spatial constraints. We next use statistical models to identify major mechanisms behind individual-level variation in tortoise burrow use, popularity of burrows in desert tortoise habitat, and test for stressor-driven changes in refuge use patterns. We show that seasonal variation has a strong impact on tortoise burrow switching behavior. On the other hand, burrow age and topographical condition influence the number of tortoises visiting a burrow in desert tortoise habitat. Of three major population stressors affecting this species (translocation, drought, disease), translocation alters tortoise burrow switching behavior, with translocated animals visiting fewer unique burrows than residents. In a species that is not social, our study highlights the importance of leveraging refuge use behavior to study the presence of and mechanisms behind non-random social structure and individual-level variation. Our analysis of the impact of stressors on refuge-based social structure further emphasizes the potential of this method to detect environmental or anthropogenic disturbances.

Behavioral Ecology and Sociobiology↗

Increased pheromone signaling by small male sea lamprey has distinct effects on female mate search and courtship

Male body size affects access to mates in many animals. Attributes of sexual signals often correlate with body size due to physiological constraints on signal production. Larger males generally produce larger signals, but costs of being large or compensation by small males can result in smaller males producing signals of equal or greater magnitude. Female choice following multiple male traits with different relationships to size might further complicate the effect of male body size on access to mates. We report the relationship between male body size and pheromone signaling, and the effects on female mate search and courtship in the sea lamprey ( Petromyzon marinus ). We predicted that pheromone production in the liver and the liver mass to body mass ratio would remain constant across sizes, resulting in similar mass-adjusted pheromone release rates across sizes but a positive relationship between absolute pheromone release and body mass. Our results confirmed positive relationships between body mass and liver mass, and liver mass and the magnitude of the pheromone signal. Surprisingly, decreasing body mass was correlated with higher pheromone concentrations in the liver, liver mass to body mass ratios, and mass-adjusted pheromone release rates. In a natural stream, females more often entered nests treated with small versus large male odors. However, close-proximity courtship behaviors were similar in nests treated with small or large male odors. We conclude that small males exhibit increased release of the main pheromone component, but female discrimination of male pheromones follows several axes of variation with different relationships to size.

Behavioral Ecology and Sociobiology↗

Occurrence of quiescence in free-ranging migratory songbirds

Quiescence is a period of inactivity that occurs before the onset of migratory activity in nocturnally migrating birds. This behavior has been observed in captive birds in migratory disposition, but its occurrence in free-ranging migratory birds has been documented only anecdotally, and causal factors and function(s), if any, are unknown. In this study, we documented and characterized quiescence in three migratory songbird species (red-eyed vireo [ Vireo olivaceus ], Swainson’s thrush [ Catharus ustulatus ], and wood thrush [ Hylocichla mustelina ]) by measuring movement and proportion of time spent inactive prior to departure from a stopover site during fall migration. Individuals of each species displayed a period of inactivity prior to departure which varied from less than 30 min to over 90 min with red-eyed vireos engaged in the longest, most pronounced quiescence. We also examined how quiescence was related to intrinsic and extrinsic factors known to influence the departure of migrating birds, and found some evidence for an effect of age and departure time but no effect of a migrant’s energetic condition, departure direction, atmospheric conditions around departure, or day of year on quiescence. Our novel application of an automated radiotelemetry system yielded a large amount of data to characterize quiescence in free-ranging migratory birds, and we provide guidance for future studies to tease apart the various causal factors and function(s) of this migratory behavior.

Behavioral Ecology and Sociobiology↗

Testing four hypotheses to explain partial migration: Balancing reproductive benefits with limits to fasting endurance

Seasonal migration is ubiquitous in animals, and yet its underlying cause(s) remain poorly known. Species exhibiting short-distance altitudinal migration and intraspecific variation in migratory behavior (partial or differential migration) are ideal study systems for examining the selective pressures that affect individual migratory decisions. We used an individually marked population of yellow-eyed juncos, breeding along a 1000-m elevational gradient and migrating up and down that gradient, to examine the morphological, behavioral, and reproductive traits associated with migratory behavior. We tested the four most well-known hypotheses proposed to explain partial migration: the thermal tolerance, fasting endurance, dominance, and arrival time hypotheses. Our results indicate that: (1) limits to juncos’ fasting endurance constrain their ability to overwinter at high elevations, in support of the fasting endurance hypothesis, (2) differences in body size mediate fasting ability and are associated with variation in migratory behavior and overwinter apparent survival, (3) migratory behavior interacts with reproductive success, in partial support of the arrival time hypothesis, and (4) additional mechanisms that are not captured by the four well-known hypotheses might better explain individual variation in migratory behavior. Less migratory females achieved greater nesting success the following breeding season. Among males, nesting success influenced migratory tendency the following winter. Successful males may either migrate to a more benign winter climate without paying reproductive costs, or high levels of parental effort might physiologically constrain their ability to overwinter in harsh climates.

Behavioral Ecology and Sociobiology↗

Daily foraging activity of an imperiled ground squirrel: Effects of hibernation, thermal environment, body condition, and conspecific density

Food acquisition is among the most important tasks faced by free-ranging animals. Predation and thermal risks, however, can make foraging a costly endeavor and foraging can preclude other important activities. Moreover, seasonal life cycle events such as hibernation impose energetic thresholds and time constraints on foraging. These factors interact with an animal’s endogenous state to influence foraging behavior. We tested a suite of predictions based on foraging theory to explore the effects of thermal environment, body condition, and conspecific density on aboveground activity (which is primarily foraging activity) of the northern Idaho ground squirrel ( Urocitellus brunneus ), an imperiled rodent that hibernates for 9 months each year. We took advantage of the squirrels’ semi-fossorial lifestyle to document daily aboveground activity by attaching geolocators to squirrels. We modeled squirrel activity with generalized linear mixed-effects models to document the relative importance of thermal environment, body condition, and conspecific density for daily aboveground activity. Aboveground activity by northern Idaho ground squirrels increased throughout their active season and leaner squirrels increased their activity more than heavier squirrels as residual foraging opportunities diminished. Thermal conditions also influenced squirrel activity: squirrels spent less time above ground during extreme temperatures and on days with significant precipitation. Aboveground activity of northern Idaho ground squirrels largely adhered to predictions of risk-sensitive and state-dependent foraging theory. Management actions that enhance forage will likely improve the probability of recovery for this federally threatened species by minimizing trade-offs squirrels need to make to acquire sufficient food to survive hibernation and reproduce in subsequent years.

Behavioral Ecology and Sociobiology↗

Offspring sex ratios are male-biased reflecting sex-biased dispersal in Idaho, USA, wolves.

Offspring sex ratios can vary widely across species, and the reasons for such variation have long intrigued ecologists. For group-living animals, predicting offspring sex ratios as a function of group and environmental characteristics can be challenging. Additionally, mortality of group members can upend traditional theory used to explain offspring sex ratios observed in populations. Gray wolves ( Canis lupus ) in Idaho, USA, are an excellent study species for asking questions about offspring sex ratios given their group-living behavior and persistent exposure to human-caused mortality. I hypothesized that offspring sex ratios would be influenced by the characteristics of individuals, groups, and populations. I generated genotypes for 419 adult and 400 pup wolves during 2008–2018. There was a significant male-bias in litters of wolf pups with nearly 12% more male pups born than females. The individual, group, and population variables I considered did not have significant associations with offspring sex ratios. Local resource competition helped explain offspring sex ratios in wolves in my study system, but not local resource enhancement theory. Although female helpers have been shown to help slightly more than males, offspring sex ratios did not favor the helping sex suggesting that the overall benefit of female helpers may have been negligible in wolf groups during my study. Three wolf groups consistently overproduced males, the dispersing sex, suggesting that habitat quality was poor in their territories. The male-biased offspring sex ratios observed throughout this population reflect sex-biased dispersal in wolves in Idaho. Such a pattern suggests breeding females may be reducing local resource competition (e.g., mates and successful reproduction) by producing more males than females.

Idaho↗

Effects of phylogeny and habitat on social structure and behavior of two equid species

Horses ( Equus caballus ) and donkeys ( E. asinus ) are closely related species with different evolutionary histories: their ancestral forms have different social systems related to their adaptations to different ecosystems, exhibiting female defense and resource defense polygyny respectively. We explored any plasticity of response and possible effects of phylogeny and habitat on these species in a feral state, comparing phylogenetic and habitat effects on behavior. We examined two feral horse populations in similar habitat in western Utah, USA, and two feral donkey populations in different habitat types (central Utah Colorado Plateau and central Arizona Sonoran Desert, USA), over 4 years. Species differences were observed: all donkeys spent more time feeding, moving, and lying down than horses, with horses standing more. Donkeys in the Sonoran Desert spent more time feeding and moving, and less time standing and lying down than on the Colorado Plateau. Horses showed less agonistic and affiliative behavior than donkeys. Between species there was no difference in network degree between horses and donkeys, and donkeys had a more saturated network with higher eigenvector centrality but lower clustering coefficient and fewer connected components than horses. Within a species, we found differences between horse populations in network degree, components, and edge density but no difference between donkey populations. Flexibility in how behavior is expressed reflects an interaction between a species’ habitat and their behavioral response to that habitat. Moreover, phylogenetic history seemingly plays a strong role in the behavioral ecology of a species, even after they are domesticated and become feralized.

Arizona, Utah↗