Social Cooperative Behavior Research Articles

Evolutionary dynamics of public goods games with diverse contributions in finite populations

The public goods game is a powerful metaphor for exploring the maintenance of social cooperative behavior in a group of interactional selfish players. Here we study the emergence of cooperation in the public goods games with diverse contributions in finite populations. The theory of stochastic process is innovatively adopted to investigate the evolutionary dynamics of the public goods games involving a diversity of contributions. In the limit of rare mutations, the general stationary distribution of this stochastic process can be analytically approximated by means of diffusion theory. Moreover, we demonstrate that increasing the diversity of contributions greatly reduces the probability of finding the population in a homogeneous state full of defectors. This increase also raises the expectation of the total contribution in the entire population and thus promotes social cooperation. Furthermore, by investigating the evolutionary dynamics of optional public goods games with diverse contributions, we find that nonparticipation can assist players who contribute more in resisting invasion and taking over individuals who contribute less. In addition, numerical simulations are performed to confirm our analytical results. Our results may provide insight into the effect of diverse contributions on cooperative behaviors in the real world.

Quorum Sensing Inhibition Selects for Virulence and Cooperation in Pseudomonas aeruginosa

With the rising development of bacterial resistance the search for new medical treatments beyond conventional antimicrobials has become a key aim of public health research. Possible innovative strategies include the inhibition of bacterial virulence. However, consideration must be given to the evolutionary and environmental consequences of such new interventions. Virulence and cooperative social behaviour of the bacterium Pseudomonas aeruginosa rely on the quorum-sensing (QS) controlled production of extracellular products (public goods). Hence QS is an attractive target for anti-virulence interventions. During colonization, non-cooperating (and hence less virulent) P. aeruginosa QS-mutants, benefiting from public goods provided by wild type isolates, naturally increase in frequency providing a relative protection from invasive infection. We hypothesized that inhibition of QS-mediated gene expression removes this growth advantage and selection of less virulent QS-mutants, and maintains the predominance of more virulent QS-wild type bacteria. We addressed this possibility in a placebo-controlled trial investigating the anti-QS properties of azithromycin, a macrolide antibiotic devoid of bactericidal activity on P. aeruginosa, but interfering with QS, in intubated patients colonized by P. aeruginosa. In the absence of azithromycin, non-cooperating (and hence less virulent) lasR (QS)-mutants increased in frequency over time. Azithromycin significantly reduced QS-gene expression measured directly in tracheal aspirates. Concomitantly the advantage of lasR-mutants was lost and virulent wild-type isolates predominated during azithromycin treatment. We confirmed these results in vitro with fitness and invasion experiments. Azithromycin reduced growth rate of the wild-type, but not of the lasR-mutant. Furthermore, the lasR-mutant efficiently invaded wild-type populations in the absence, but not in the presence of azithromycin. These in vivo and in vitro results demonstrate that anti-virulence interventions based on QS-blockade diminish natural selection towards reduced virulence and therefore may increase the prevalence of more virulent genotypes in the Hospital environment. More generally, the impact of intervention on the evolution of virulence of pathogenic bacteria should be assessed. Trial Registration: ClinicalTrials.gov NCT00610623

Beliefs, guilt, and the brain: Neural mechanisms underlying social cooperative behavior

Beliefs, guilt, and the brain: Neural mechanisms underlying social cooperative behavior

Human uniqueness-self-interest and social cooperation

Humans are unique among all species of terrestrial history in both ecological dominance and individual properties. Many, or perhaps all, of the unique elements of this nonpareil status can be plausibly interpreted as evolutionary and strategic elements and consequences of the unprecedented intensity and scale of our social cooperation. Convincing explanation of this unique human social adaptation remains a central, unmet challenge to the scientific enterprise. We develop a hypothesis for the ancestral origin of expanded cooperative social behavior. Specifically, we present a game theoretic analysis demonstrating that a specific pattern of expanded social cooperation between conspecific individuals with conflicts of interest (including non-kin) can be strategically viable, but only in animals that possess a highly unusual capacity for conspecific violence (credible threat) having very specific properties that dramatically reduce the costs of coercive violence. The resulting reduced costs allow preemptive or compensated coercion to be an instantaneously self-interested behavior under diverse circumstances rather than in rare, idiosyncratic circumstances as in actors (animals) who do not have access to inexpensive coercive threat. Humans are apparently unique among terrestrial organisms in having evolved conspecific coercive capabilities that fulfill these stringent requirements. Thus, our results support the proposal that access to a novel capacity for projection of coercive threat might represent the essential initiating event for the evolution of a human-like pattern of social cooperation and the subsequent evolution of the diverse features of human uniqueness. Empirical evidence indicates that these constraints were, in fact, met only in our evolutionary lineage. The logic for the emergence of uniquely human cooperation suggested by our analysis apparently accounts simply for the human fossil record.

Emotional Understanding, Cooperation, and Social Behavior in High-Functioning Children with Autism

In contrast to typically developing children, children with autism rarely exhibit cooperative social behavior. To examine whether this problem reflects global developmental delays or autism-specific deficits, the present study compared cooperation, emotional understanding, personality characteristics, and social behavior of 10 children with autism who had average IQ to those of 16 children with Attention-Deficit/Hyperactivity Disorder (ADHD) and Oppositional Defiant Disorder (ODD) and 10 typically developing children. In cooperative behavior, level of emotional understanding, and aloof behavior, the autism group outperformed the ADHD/ODD group and did not differ significantly from typically developing children. However, the autism group showed worse emotion recognition and more active-but-odd behavior than the other groups. The results indicate that high-functioning children with autism can develop cooperative social behavior and advanced theory of mind abilities, but continue to show deficits in identifying emotions and displaying socially appropriate behavior.

Social neuroscience, empathy, brain integration, and neurodevelopmental disorders

Paul MacLean has investigated integrated brain functioning through selected brain lesions in animals that disturb circuits necessary for complex behaviors, such as social displays. MacLean is unique in his comparative neurobehavioral approach that emphasizes the evolutionary origins of parenting and social behaviors and the implications of brain changes in the evolution from reptiles (social displays) to mammals (nursing, audiovocal communication, play) to man (self-awareness, intentionality, social context) that link affect and cognition. Subjectively, how “looking with feeling toward others,” the basic element in empathy, evolved has been a central concern of his. Neuroimaging studies of social cognition, mother–infant communication, moral behavior, forgiveness, and trust are consistent with particular brain systems being activated in cooperative social behaviors. The identification of mirror neurons is pertinent to MacLean's model of isopraxis and studies of thalamocortical resonances may be pertinent to his neurobehavioral models. Studies of behavioral phenotypes in human neurodevelopmental disorders are consistent with MacLean's model of brain circuits being linked to complex behaviors during development. In autistic disorder, the behavioral phenotype involves disrupted social communication, deviant imaginative play, and motor stereotypies. In Lesch–Nyhan syndrome (LNS), self-injury occurs in individuals with normal sensory systems intact who require and request physical restraint to prevent self-injury; they ask for assistance from others to prevent them from harming themselves. Autism involves the lack of subjective awareness of others intentions and LNS involves a failure in self-regulation and self-control of self-injurious behavior. MacLean's models laid the groundwork for studies focused on understanding brain functioning in these conditions.

Developing a techno entrepreneurial workforce for effective technological innovation: its necessity for the sustainable development of SSEs in India

Due to the application of new technologies, changes in tasks and organisational structures, a shorter product lifetime cycle, and changing customer requirements of product and services, as well as liberalisation and globalisation, entrepreneurs in the small-scale sector are confronted with new qualification demands. The typical technical/scientific qualification has to be extended by interdisciplinary knowledge and skills, which enables creative problem solutions, cooperative social behaviour with leadership qualities and effective communication competencies, as well as a holistic view of technical problems.

A Neural Basis for Social Cooperation

Cooperation based on reciprocal altruism has evolved in only a small number of species, yet it constitutes the core behavioral principle of human social life. The iterated Prisoner's Dilemma Game has been used to model this form of cooperation. We used fMRI to scan 36 women as they played an iterated Prisoner's Dilemma Game with another woman to investigate the neurobiological basis of cooperative social behavior. Mutual cooperation was associated with consistent activation in brain areas that have been linked with reward processing: nucleus accumbens, the caudate nucleus, ventromedial frontal/orbitofrontal cortex, and rostral anterior cingulate cortex. We propose that activation of this neural network positively reinforces reciprocal altruism, thereby motivating subjects to resist the temptation to selfishly accept but not reciprocate favors.

On the relative importance of haplodiploidy, assortative mating and social synergy on the evolutionary emergence of social behavior.

Advances in multiagent simulation techniques make it possible to study more realistic dynamics of complex systems and allow evolutionary theories to be tested. Here I use simulations to assess the relative importance of reproductive systems (haplodiploidy vs. diploidy), mate selection (assortative mating vs. random mating) and social economics (pay-off matrices of evolutionary games) in the evolutionary dynamics leading to the emergence of social cooperation in the provision of parental care. The simulations confirm that haplo-diploid organisms and organisms mating assortatively have a higher probability for fixing alleles and require less favorable conditions for their fixation, than diploids or organisms mating randomly. The simulations showed that social behavior was most likely to emerge a) when the cost for parental investment was much lower than the benefits to the offspring, b) when cooperation improved synergistically the fitness of offspring compared to the corresponding egoistic behavior and c) when alleles coding for altruistic or social behavior could be rapidly fixed in the population, thanks to mechanisms such as haplo-diploidy and/or assortative mating. Cooperative social behavior always appeared if sociality conferred much higher fitness gains compared to non cooperative alternatives suggesting that the most important factors for the emergence and maintenance of social behavior are those based on energetic or efficiency considerations. The simulations, in congruence with the scant experimental evidence available, suggest that economic considerations rather than genetic ones are critical in explaining the emergence and maintenance of sociality.

The significance of relatedness and gene flow on population genetic structure in the subsocial spider Eresus cinnaberinus (Araneae: Eresidae)

Interdemic selection, inbreeding and highly structured populations have been invoked to explain the evolution of cooperative social behaviour in the otherwise solitary and cannibalistic spiders. The family Eresidae consists of species ranging from solitary and intermediate subsocial to species exhibiting fully cooperative social behaviour. In this study we, in a hierarchical analysis, investigated relatedness of putative family clusters, inbreeding and population genetic structure of the subsocial spider Eresus cinnaberinus. Five hierarchical levels of investigation ranging from large scale genetic structure (distances of 250 and 50 km level 1 and 2) over microgeographic structure (20 km2 and 4 km2, level 3 and 4) to a single hill transect of 200 m (level 5) were performed. The purpose of level 5 was two-fold: (1) to investigate the relatedness of putative family groups, and (2) to evaluate the influence of both family living and sampling design on higher level estimates. Relatedness estimates of putative family groups showed an average relatedness of R=0.26. There was no indication of inbreeding. In contrast to social spiders, genetic variation was abundant, Heapproximately0.10. The population genetic structure was intermediate between social and asocial spiders. Genetic variance increased continually across hierarchical levels. Family structured neighbourhoods biased differentiation estimates among level 5 samples (FST=0.04) and level 3 and 4 samples (0.07<FST<0.18), and apparent inbreeding among level 3 and 4 samples, FIS>0, was caused by disjunct sampling from separate neighbourhoods. Larger scale samples were highly differentiated 0.12<FST<0.26, depending on level and sampling design. Due to a distance effect family living did not influence estimates of the higher level 1. Although the dispersing sex among social spiders and the subsocial E. cinnebarinus differ, females versus males, female behaviour of both sociality classes lead to high genetic variance.Copyright 1998 The Linnean Society of London

The significance of relatedness and gene flow on population genetic structure in the subsocial spiderEresus cinnaberinus(Araneae: Eresidae)

Interdemic selection, inbreeding and highly structured populations have been invoked to explain the evolution of cooperative social behaviour in the otherwise solitary and cannibalistic spiders. The family Eresidae consists of species ranging from solitary and intermediate subsocial to species exhibiting fully cooperative social behaviour. In this study we, in a hierarchical analysis, investigated relatedness of putative family clusters, inbreeding and population genetic structure of the subsocial spiderEresus cinnaberinus. Five hierarchical levels of investigation ranging from large scale genetic structure (distances of 250 and 50 km level 1 and 2) over microgeographic structure (20 km2and 4 km2, level 3 and 4) to a single hill transect of 200 m (level 5) were performed. The purpose of level 5 was two-fold: (1) to investigate the relatedness of putative family groups, and (2) to evaluate the influence of both family living and sampling design on higher level estimates. Relatedness estimates of putative family groups showed an average relatedness ofR=0.26. There was no indication of inbreeding. In contrast to social spiders, genetic variation was abundant,He≈0.10. The population genetic structure was intermediate between social and asocial spiders. Genetic variance increased continually across hierarchical levels. Family structured neighbourhoods biased differentiation estimates among level 5 samples (FST=0.04) and level 3 and 4 samples (0.07<FST<0.18), and apparent inbreeding among level 3 and 4 samples,FIS>0, was caused by disjunct sampling from separate neighbourhoods. Larger scale samples were highly differentiated 0.12<FST<0.26, depending on level and sampling design. Due to a distance effect family living did not influence estimates of the higher level 1. Although the dispersing sex among social spiders and the subsocialE. cinnebarinusdiffer, females versus males, female behaviour of both sociality classes lead to high genetic variance.

Ecological constraints drive social evolution in the African mole–rats

The African mole-rats (family Bathyergidae) are subterranean hystricomorph rodents occurring in a variety of habitats and displaying levels of sociality which range from solitary to eusocial, making them a unique mammalian taxonomic group to test ecological influences on sociality. Here, we use an extensive DNA-based phylogeny and comparative analysis to investigate the relationship between ecology, sociality and evolution within the family. Mitochondrial cytochrome-b and 12s rRNA trees reveal that the solitary species are monophyletic when compared to the social species. The naked mole-rat (Heterocephalus glaber) is ancestral and divergent from the Damaraland mole-rat (Cryptomys damarensis), supporting previous findings that have suggested the multiple evolution of eusociality within the family. The Cryptomys genus is species-rich and contains taxa exhibiting different levels of sociality, which can be divided into two distinct clades. A total of seven independent comparisons were generated within the phylogeny, and three ecological variables were significantly correlated with social group size: geophyte density (p < 0.05), mean months per year of rainfall greater than 25 mm (p < 0.001), and the coefficient of rainfall variation (p = 0.001). These results support the food-aridity hypothesis for the evolution of highly social cooperative behaviour in the Bathyergidae, and are consistent with the current theoretical framework for skew theory.

Low roost-site fidelity in pallid bats: associated factors and effect on group stability

Despite potential costs of changing roost or densites, many animals frequently move between roosts or dens. Pallid bats (Antrozous pallidus) change diurnal roost sites frequently and also reportedly have a variety of cooperative social behaviors, many of which are associated with the care of developing offspring. Roost switching is likely to increase the costs of maintaining the group stability expected with social cooperation. Pallid bats roosting in rock crevices in central Oregon were studied with radiotelemetry to (1) examine characteristics of day roosts, (2) determine what ecological factors were correlated with low roost fidelity, and (3) examine the temporal stability of roosting groups of pregnant and lactating bats. Pallid bats changed roosts an average of once every 1.4 days throughout the summer. The bats exhibited seasonal shifts in roost use, occupying roosts behind thin slabs of rock in cool weather and roosts in deep rock crevices in warm weather. Roost switching was not correlated with daily variations in weather conditions or with structural characteristics of the diurnal roosts, although switching may have allowed bats to maintain familiarity with several roosts that vary in microclimate. Roost switching was positively correlated with ectoparasite load. High ectoparasite levels were correlated with lower body weights in lactating females (Fig. 3), suggesting that parasites may be costly to the bats. Roost switching may be a strategy to decrease ectoparasite loads by interrupting the reproductive cycles of those parasites that spend at least part of their life cycle on the walls of the roost. Both pregnant and lactating pallid bats frequently changed their diurnal roost location, but lactating bats tended to travel shorter distances between consecutive roosts. Lactating bats were more likely to continue to associate with particular roostmates despite changes in the location of the diurnal roost (Fig. 4) and were less likely to roost alone. Although the stability of groups of lactating bats was not absolute, evidence supported the prediction that such groups are more cohesive than are those of pregnant bats.

Psychopharmacology of Central Serotonergic Systems

Serotonin neurons in the rostral and caudal brainstem raphe nuclear groups give rise to collateralized ascending and descending projections which provide modulatory input into most networks throughout the entire neuraxis. The rostral raphe system is interconnected with target forebrain areas through reciprocal limbic-midbrain loops, which suggests that serotonin has a role in the regulation of complex intelligent adaptive behavior. Serotonergic pathways sensitize brainstem and spinal cord central rhythmic pattern generators which organize repetitive autonomic and motor activities, e.g. oral-buccal and nutritive behaviors, facilitate tonically active motor neurons innervating antigravity muscles, and disfacilitate somatosensory information processing. Serotonin effects are mediated by multiple receptor subtypes with distinct pre- and postsynaptic localization and regional distribution pattern. They belong to the G protein superfamily, coupling to adenylate cyclase (5-HT1,4,5,6,7) or phospholipase C (5-HT2), and to the ligand-gated ion channel superfamily (5-HT3). Drugs acting at these receptors are known to modulate various aspects of cooperative social behavior and responding latency, i.e. impulsivity, in a variety of experimental models of anxiety and depression. The clinical efficacy of the so-called selective serotonin reuptake inhibitors (SSRIs) in disorders characterized by poor impulse control, e.g. bulimia nervosa, obsessive-compulsive disorder (OCD) and violent suicidal or homicidal behavior, may likewise be due to improved responding latency.

Effects of 1988 Fires on Ecology of Coyotes in Yellowstone National Park: Baseline Preceding Possible Wolf Recovery

Sixty healthy coyotes Canis latrans and 53, 8-12 week old pups captured at dens were radio­tagged in the Lamar Valley and Blacktail Plateau areas of the northern range of Yellowstone National Park. Adults range in age from 1 to 12 years and averaged 3.3 years old. Territorial packs in both study area are adjacent, non-overlapping, contiguous, and averaged 15 km2. Based on information the last four winters and data collected from 1946 to 1949, territorial areas are traditional and have changed little in the last 45 years. We estimate that 85 to 90% of coyotes on the northern range belong to packs. A territorial group or pack during the winter consists of 2 alpha individuals, 2 or 3 beta adults, and 2 or 3 adult-sized pups. Average pack size was 6.3 for Lamar Valley and 4.6 for Blacktail Plateau. Mean litter size for 1990 through 1994 was 4.1, 5.7, 6.5, 3.3, and 2.3 for those five years. Initial density estimates are 1.4 coyotes per square mile. Preliminary scat analysis suggests that small mammals, especially voles, dominate the diet with ungulate remains becoming important in May through July (presumably elk calves) and late winter (mostly scavenging). Graduate students Eric Gese, Kezha Hatier, and Scott Grothe have finished their data collection and are analyzing data and preparing manuscripts. More than 2500 hours of foraging observations were conducted from January 1991 through June 1993 resulting in data collection on more than 4400 predation attempts on small mammals. Eight hundred and fifty hours of den observations were completed during 1992 and 1993. Beta pack members were observed to bring food to pups and protect den sites from intruders. Coyote behavior and ungulate mortality data were collected on 80 carcasses found or translocated during the 1992-93 and 1993-94 winters. Coyotes were observed at these carcasses for 484 hours. Alpha males usually feed first at carcasses followed by alpha females, beta individuals and pups. Those first to feed typically eat the internal organs and muscle tissue first. Betas and pups are left to feed mostly on bones and hide. Five successful and 4 unsuccessful predations by coyotes on ungulates have been seen. Coyotes appear to impact ungulate numbers in 3 ways: predation on calves and fawns shortly after birth (up to 8 weeks), predation on short-yearlings and adults during winter, and indirect impact from harassment of other predators at ungulate-kills. Coyotes may be the major ungulate predator on the northern range due to cooperative social and foraging behavior, their ability to take advantage of vulnerable ungulates, and their high population levels. Wolf extirpation has probably resulted in high coyote population densities and coyotes have, at least, partially slid into this vacant niche.

Effects of 1988 Fires on Ecology of Coyotes in Yellowstone National Park: Baseline Preceding Possible Wolf Recovery

Sixty healthy adult coyote Canis latrans and 53, 8-12 week old pups captured at dens were radio­ tagged in the Lamar Valley and Blacktail Plateau areas of the northern range of Yellowstone National Park. Adults range in age from 1 to 12 years and average nearly 3.3 years old. Territorial packs in both study areas are adjacent, non-overlapping, contiguous, and average 15 km2. Based on information from the last four winters and data collected from 1946 to 1949, territorial areas are traditional and have changed little in the last 45 years. We estimate that 85 to 90% of coyotes on the northern range belong to packs. A territorial group or pack during the winter consists of 2 alpha individuals, 2 or 3 beta adults, and 2 or 3 adult-sized pups (average pack size = 7). Fifteen marked coyotes have died since November of 1992. Initial density estimates are 1.4 coyotes per square mile. Preliminary scat analysis suggests that small mammals, especially voles, dominate the diet with ungulate remains becoming important in May through July (presumably elk calves) and late winter (mostly scavenging). Two graduate students have finished their data collection and another student is currently working on his last field season. More than 2500 hours of foraging observations were conducted from January 1991 through June 1993 resulting in data collection on more than 4400 predation attempts on small mammals. Eight hundred and fifty hours of den observations were completed during 1992 and 1993. Beta pack members were observed to bring food to pups and protect den sites from intruders. Coyote behavior and ungulate mortality data were collected on sixty-one carcasses found during the 1992-1993 winter. Five successful and 4 unsuccessful predations by coyotes on ungulates have been seen. Coyotes appear to impact ungulate numbers in 3 ways: predation on calves and fawns shortly after birth (up to 8 weeks), predation on short-yearlings and adults during winter, and indirect impact from harassment of other predators at ungulate-kills. Coyotes may be the major ungulate predator on the northern range due to their cooperative social and foraging behavior, ability to take advantage of vulnerable ungulates, and high population levels. Wolf extirpation has probably resulted in high coyote population densities and coyotes have, at least partially, slid into this vacant niche.

Effects of 1988 Fires on Ecology of Coyotes in Yellowstone National Park: Baseline Preceding Possible Wolf Recovery

Sixty-five healthy coyote (Canis latrans) adults and 53, 8-12 week old pups captured at dens were radio-tagged in the Lamar Valley and Blacktail Plateau areas of the northern range of Yellowstone National Park. Adults range in age from 1 to 11 years and averaged nearly 3 years old. Territorial packs in both study areas are adjacent, non­overlapping, contiguous, and averaged 15 km2. Based on information the last three winters and data collected from 1946 to 1949, territorial areas are traditional and have changed little in the last 45 years. We estimate that 85 to 90% of coyotes on the northern range belong to packs. A territorial group or pack during the winter consists of 2 alpha individuals, 2 or 3 beta adults, and 2 or 3 adult-sized pups (average pack size = 7). Only one radioed adult coyote has died since May of 1992. Twenty­ four of 53 pups have died between the ages of 3 and 9 months old. Population productivity ranges from 1.8 to 2.5 pups recruited per territory. The reproductive failure rate among breeding groups averaged 15% during 1990 and 1991. Initial density estimates are 1.4 coyotes per square mile. Intensive foraging observations were conducted from January through June 1991 (353 hours) and from November 1991 through April 1992 (1100+hours). Focal observations collected from January-June 1991 resulted in 427 capture attempts on small mammal prey with 162 (38%) successful. Habitat type played a key role in the success rate. Preliminary analysis of the November 1991 to April1992 data indicated a substantial reduction in prey attempts and prey success. This reduction was mostly a function of harsh snow conditions in early winter and abundlint elk carrion in late winter. Over eighty ungulate carcass were located this winter in the 2 study areas. However, small mammals, especially voles, dominate the diet with ungulate remains becoming important in May through July (presumably elk calves) and late winter (mostly scavenging). We have documented numerous successful and unsuccessful predation attempts on ungulates in our study areas. Coyotes appear to impact ungulate numbers in 3 ways: predation on calves and fawns shortly after birth (up to 8 weeks), predation on short-yearlings and adults during winter, and indirect impact from harassment of other predators at ungulate-kills. Coyotes may be the major ungulate predator on the northern range due to cooperative social and foraging behavior, their ability to take advantage of vulnerable ungulates, and their high population levels. Wolf extirpation has probably resulted in high coyote population densities and coyotes have, at least, partially slid into this vacant niche.

Identity and cooperative social behavior: Pseudospeciation or human integration?

(1991). Identity and cooperative social behavior: Pseudospeciation or human integration? World Futures: Vol. 31, Cooperation: Toward a Post-Modern Ethic, pp. 95-106.

Effects of 1988 Fires on Ecology of Coyotes in Yellowstone National Park: Baseline Preceding Possible Wolf Recovery

Fifty-four healthy coyotes (Canis latrans) and 32, 8-12 week old pups captured at dens were radio­tagged in the Lamar Valley and Blacktail Plateau areas of the northern range of Yellowstone National Park. Seven of the 40 captured in the fall were 6 month-old pups which suggest slow population productivity. Adults range in age from 1 to 12 years and averaged nearly 4 years old. Territorial packs in both study areas are adjacent, non-overlapping, contiguous, and averaged 15 km2/ We estimate that 85 to 90% of coyotes on the northern range belong to packs. A territorial group or pack during the winter consists of 2 alpha individuals, 2 or 3 beta adults, and 2 or 3 adult-sized pups (average pack size = 7). Nine adults were killed (2 mountain lion [Felis concolar], 2 road-kill, 2 shot, and 3 unknown) which equates to a 15% annual mortality rate. Eleven of 36 pups have died between the ages of 3 and 9 months old. Population productivity ranges from 2.0 to 2. 7 pups recruited per territory. The reproductive failure rate among breeding groups averaged 15% during 1990 and 1991. Initial coyote density estimates are 0.09 per km2. Intensive foraging observations were conducted from January through June 1991. In 353 hours of focal observations 427 capture attempts were made on small mammal prey with 162 (38%) successful. Habitat type played a key role in the success rate. Mesic meadows had the highest capture rates followed by willow/meadow habitats and sage habitat. Small mammals, especially voles (Microtus spp.), dominate the diet with ungulate remains becoming important in May through July, presumably elk (Cervus elaphus) calves and late winter, mostly scavenging. We have observed numerous successful and unsuccessful predation attempts on ungulates in our study areas. Coyotes appear to impact ungulate numbers in 3 ways: predation on calves and fawns shortly after birth (up to 8 weeks), predation on short-yearlings and adults during winter, and indirect impact from harassment of other predators at ungulate-kills. Coyotes may be the major ungulate predator on the northern range due to cooperative social and foraging behavior, their ability to take advantage of vulnerable ungulates, and their high population levels. Wolf (Canis lupus) extirpation has probably resulted in high coyote population densities.

Effects of isolated housing and chronic antidepressant treatment on cooperative social behaviour in rats

Effects of isolated housing and chronic antidepressant treatment on cooperative social behaviour in rats