Territorial cooperation and social monogamy: factors affecting intersexual behaviours in pair-living snapping shrimp

Among the factors that may contribute to the evolution of social monogamy are selection for extended mate guarding of females and selection for territorial ‘cooperation’. Many socially monogamous taxa are also territorial, with ‘partners’ sharing a single territory, suggesting that one or both partners may benefit by sharing territorial maintenance. Snapping shrimp (genus Alpheus) are socially monogamous and territorial, living in excavated burrows or with host organisms, with females performing all parental care. The territorial cooperation hypothesis predicts that male and female partners share (1) territorial defence, resulting in a reduction in the risk of eviction from the burrow, (2) burrow construction duties, such that individuals in pairs spend less time in burrow construction relative to solitary individuals, and/or (3) foraging duties, by returning food to the burrow, where it is consumed by both partners. UsingA. angulatus as a model species, a territorial defence experiment revealed that females in pairs were significantly less likely than solitary females to be evicted by female intruders, but males in pairs were not significantly less likely than solitary males to be evicted by male intruders. A subsequent experiment revealed that paired males were significantly less likely to be evicted by an intruding male if paired with sexually receptive females than if paired with nonreceptive females. Another experiment revealed that (1) paired females spent significantly more time in burrow construction than paired males, and (2) both males and females consistently returned food items to the burrow, perhaps incidentally provisioning their mates. These data suggest that social monogamy may have been selected for in part because of the advantages of territorial cooperation, as both males and females are likely to benefit by dividing the labour of territorial defence and maintenance. These tests of the territorial cooperation hypothesis are synthesized with data from tests of the extended mate-guarding hypothesis to place snapping shrimp pairing behaviour into a larger construct incorporating both the influence of ecological pressures (territoriality) and mating interactions between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Scent-marking in free-ranging coyotes, Canis latrans

From November 1977 to March 1978 we recorded the scent-marking behaviour of free-ranging coyotes in southern Québec. Fourteen animals were marked by foot-pad ablation, which made it possible to recognize individuals' tracks in the snow and therefore to relate scent-marking with social organization. We observed 949 scent-marks. The identity of the animals was known for 50% of the tracking distances. Animals living in territorial groups (pairs and pairs with juveniles) marked more than twice as much as solitary non-territorial individuals. In addition the former used stronger forms of marking significantly more often than the latter.     

Poor health is associated with use of anthropogenic resources in an urban carnivore

Rates of encounters between humans and wildlife are increasing in cities around the world, especially when wildlife overlap with people in time, space and resources. Coyotes (Canis latrans) can make use of anthropogenic resources and reported rates of conflict have increased in cities across North America. This increase may be linked to individual differences in the use of human food and developed areas. We compared the relationships between coyote age, sex or health and the use of anthropogenic resources, which we defined as using developed areas over large home ranges, being active during the day, and consuming anthropogenic food. To do so, we applied GPS collars to 19 coyotes and sampled hair for stable isotope analysis. Eleven coyotes appeared to be healthy and eight were visibly infested with sarcoptic mange (Sarcoptes scabiei), a mite that causes hair loss. Diseased coyotes used more developed areas, had larger monthly home ranges, were more active during the day, and assimilated less protein than coyotes that appeared to be healthy. We speculate that anthropogenic food provides a low-quality but easily accessible food source for diseased coyotes, which in turn may increase reliance on it and other anthropogenic resources to promote encounters with people.     

Behavioral Ecology of Coyotes: Social Organization,Rearing Patterns,Space Use,and Resource Defense

Two groups of coyotes in which genealogical relationships were known were studied in the Grand Teton National Park, outside of Jackson, Wyoming, U.S.A., from 1977–1982. One group, a pack consisting of parents and some non-dispersing and non-breeding offspring, defended a territory and the food (mainly elk carrion) contained within it, especially during winter, and also had helpers at den sites (5 of 6 were males). The other group, a mated resident pair, all of whose young dispersed during the first year of life, did not defend a territory and never had helpers at dens. Delayed dispersal and retention of some offspring as helpers was related to the presence of an abundant, clumped, and defendable winter food resource. Dispersing yearlings suffered higher mortality than did non-dispersing individuals. Litter size was the same for the pack and resident pair; litter size was not significantly correlated with number of adults in the group or with the number or percentage of pups that survived to 5–6 months of age. The presence of pack helpers was not significantly correlated with pup survival, although there was a positive correlation (r^s = +0.37) between the number of adults attending a den(s) and pup survival. Helpers rarely fed pups and their presence had no appreciable effect on juvenile weight. Helpers partook in den-sitting (pup-guarding), but they did not reduce the amount of time that parents spent at den sites. Helpers also actively initiated and took part in territorial and food defense. The proportion of times that pack members initiated defense was inversely related to intruder density (r = –0.94).     

Male copulation behaviour and the risk of sperm competition

When females mate with more than one male during their reproductive cycle, males may increase their share of paternity by copulating repeatedly with the same female. Accordingly, males should mate repeatedly with the same female more frequently when the risk of sperm competition is greater. We examined this idea experimentally in the orb-web spiderNephila edulis , which is characterized by both extreme sexual size dimorphism and extreme male size variation. Comparison of the mating behaviour of solitary and pairs of males on the webs of virgin and mated females revealed that males adjust the frequency and duration of copulation according to the mating history of the female and the presence of rival males. Males copulated more frequently and for longer with virgin than mated females. The copulation behaviour of males in the presence of rivals depended upon their relative size. Typically, larger males prevented smaller rivals from gaining access to the female and therefore were able to copulate more frequently. Smaller males copulated less frequently, but for longer periods, which may have increased their share of paternity. The size of male N. edulis can vary by an order of magnitude, and our results suggest that this variation may be maintained by the alternative size-dependent strategies of preventing or winning sperm competition. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Fine‐scale movements and behaviors of coyotes (Canis latrans) during their reproductive period

In canids, resident breeders hold territories but require different resources than transient individuals (i.e., dispersers), which may result in differential use of space, land cover, and food by residents and transients. In the southeastern United States, coyote (Canis latrans) reproduction occurs during spring and is energetically demanding for residents, but transients do not reproduce and therefore can exhibit feeding behaviors with lower energetic rewards. Hence, how coyotes behave in their environment likely differs between resident and transient coyotes. We captured and monitored 36 coyotes in Georgia during 2018–2019 and used data from 11 resident breeders, 12 predispersing residents (i.e., offspring of resident breeders), and 11 transients to determine space use, movements, and relationships between these behaviors and landcover characteristics. Average home range size for resident breeders and predispersing offspring was 20.7 ± 2.5 km² and 50.7 ± 10.0 km², respectively. Average size of transient ranges was 241.4 ± 114.5 km². Daily distance moved was 6.3 ± 3.0 km for resident males, 5.5 ± 2.7 km for resident females, and 6.9 ± 4.2 km for transients. We estimated first‐passage time values to assess the scale at which coyotes respond to their environment, and used behavioral change‐point analysis to determine that coyotes exhibited three behavioral states. We found notable differences between resident and transient coyotes in regard to how landcover characteristics influenced their behavioral states. Resident coyotes tended to select for areas with denser vegetation while resting and foraging, but for areas with less dense vegetation and canopy cover when walking. Transient coyotes selected areas closer to roads and with lower canopy cover while resting, but for areas farther from roads when foraging and walking. Our findings suggest that behaviors of both resident and transient coyotes are influenced by varying landcover characteristics, which could have implications for prey.     

Social interactions regulate resource utilization in a Tephritidae fruit fly

Previous studies have suggested that social interactions (e.g., the actions and reactions elicited by the interaction of co-specific individuals) induce individual fruit flies (Tephritidae) to ingest more food, especially protein-rich food. Changes in feeding behavior related to social interactions have been associated with reproduction (e.g., when different sexes are present), reproductive facilitation (e.g., when two females interact) and stress and aggression (e.g., flies of the same sex, or crowdedness). The present study investigated the effect of social interaction on the feeding, longevity and resource management of the Ethiopian fruit fly, Dacus ciliatus. Single flies and pairs of flies (of the same or different sexes) were confined to a small arena (the PUB system), in which we measured the amount of liquid food ingested daily by each fly. In addition, we sampled flies of different ages, extracted and quantified their lipid and protein contents, and related individual metabolic contents to the ingestion of a fructose and protein hydrolysate solution. Results showed that individual ingestion was significantly higher in flies maintained in pairs than in flies kept as solitary individuals. The highest intake rates were observed for the female–female pairs. In general, females ingested significantly greater volumes than males. Lipid contents tended to decrease progressively with age in flies kept as solitary individuals, especially in female flies, while lipid levels decreased and then increased in flies maintained in pairs. Protein trends were similar, although less pronounced than the patterns observed for the lipids. The flies kept as solitary individuals lived significantly longer than those kept in pairs. A resource-management analysis points to a decreased metabolic rate in flies kept as solitary individuals, as compared to paired flies. Results are discussed in view of theories of resource management and survival strategies.     

Dominance and neophobia in coyote (Canis latrans) breeding pairs

Previous research suggests that within social animals, subordinate individuals are less neophobic than dominant individuals. We investigated the effect of social status on neophobic responses using 10 captive coyote breeding pairs. Social status was determined from observations of feeding behavior and agonistic interactions during a series of reference trials. Once dominance was established, we surrounded an experimental area with novel stimuli (ropes adorned with interspersed flags) to create a novel context around a familiar food source. Contrary to hypotheses, dominant coyotes were first to feed, showed more interest toward novel stimuli, and eventually crossed the barrier, which subordinates never did. Our results indicate that dominant coyotes are less neophobic of novel settings that contain familiar food than subordinates are. Since a reduction in neophobia can be interpreted as an increase in risk taking, our results support previous observations that dominant (alpha) coyotes take more risks than subordinates. Our results also suggest reasons for differential observations of coyote behavior in the field: artificial selection against bold behavior in populations undergoing predator control.     

Laboratory breeding studies of freshwater crayfish, Austropotamobius pallipes (Lereboullet)

SUMMARY. 1. Mature crayfish, collected from an Irish lake before breeding had started, were held in breeding combinations and their mating and brooding activities observed.
2. All mating attempts were initiated by the male. A single mating led to spawning within 6 days but a subsequent mating cancelled the effects of the first. Males mated more often when there were more females present. Males lacking a major cheliped mated less often than did normal males.
3. Larger males mated more often than did smaller males, and although males showed no female size preference, matings were less frequent and generally unsuccessful when males were much larger than females; the female was usually killed. Large females mated successfully with smaller males.
4. Females held at high densities with a larger male mated earlier than at low densities. However, aggression also increased with density; at high densities males fought and killed females.
5. Males held in pairs without females fought; in occasional mating attempts spermatophores were not positioned correctly. Paired females rarely fought; all spawned normally although unmated. Although their eggs soon died and were removed during grooming, brooding behaviour continued for at least 2 months.
6. Brooding females held in pairs shed pleopodal eggs during aggressive encounters. Females held singly showed a lower initial rate of egg loss.     

Greater consumption of protein‐poor anthropogenic food by urban relative to rural coyotes increases diet breadth and potential for human–wildlife conflict

Reports of encounters between people and generalist urban‐adapted carnivores are increasing around the world. In North America, coyotes Canis latrans are among the carnivores that appear to be especially capable of incorporating novel anthropogenic food types, including those found in cities. Consuming anthropogenic food may benefit coyotes by increasing their dietary diversity, but it may also lead to increased interactions and conflicts with humans. To test these hypotheses, we compared the diets of urban and rural coyotes from two urban and three rural sites spanning 32 200 km² in Alberta, Canada. We analyzed scat samples to calculate diet diversity at the level of both individuals (species per scat) and populations (Shannon index) and to determine the frequency of anthropogenic food consumption. We complemented this comparison with stable isotope analyses of hair samples taken from individual urban and rural coyotes that were or were not reported by the public for repeatedly visiting backyards and schoolyards during the day. Relative to rural coyotes, urban coyotes had more diverse diets at the level of both individuals and populations, consumed anthropogenic food more often, and animals less often, than rural coyotes. Although urban coyotes assimilated more anthropogenic food than the rural coyotes overall, the urban coyotes reported for conflict assimilated less protein and were more likely to be diseased. Our results suggest that processed anthropogenic food may contribute to the success of urban coyotes, but does not entirely correlate with conflict. Instead, some seemingly innocuous, but low‐protein food sources such as bird feeders, compost, and cultivated fruit trees may contribute disproportionately to encounters with people for coyotes and other urban‐adapted opportunistic carnivores.     

Resident and transient coyotes exhibit differential patterns of movement behavior across heterogeneous landscapes in the southeastern United States

Coyotes (Canis latrans) are a highly adaptable canid species whose behavioral plasticity has allowed them to persist in a wide array of habitats throughout North America. As generalists, coyotes can alter movement patterns and change territorial strategies between residency (high site fidelity) and transiency (low site fidelity) to maximize fitness. Uncertainty remains about resident and transient coyote movement patterns and habitat use because research has reached conflicting conclusions regarding patterns of habitat use by both groups. We quantified effects of habitat on resident and transient coyote movement behavior using first passage time (FPT) analysis, which assesses recursive movement along an individual''s movement path to delineate where they exhibit area‐restricted search (ARS) behaviors relative to habitat attributes. We quantified monthly movement rates for 171 coyotes (76 residents and 53 transients) and then used estimated FPT values in generalized linear mixed models to quantify monthly habitat use for resident and transient coyotes. Transients had greater movement rates than residents across all months except January. Resident FPT values were positively correlated with agricultural land cover during fall and winter, but negatively correlated with agriculture during spring. Resident FPT values were also negatively correlated with developed habitats during May–August, deciduous land cover during June–August, and wetlands during September–January except November. FPT values of transient coyotes were positively correlated with developed areas throughout much of the year and near wetlands during July–September. Transient FPT values were negatively correlated with agriculture during all months except June and July. High FPT values (ARS behavior) of residents and transients were generally correlated with greater densities of edge habitat. Although we observed high individual variation in space use, our study found substantive differences in habitat use between residents and transients, providing further evidence that complexity and plasticity of coyote habitat use is influenced by territorial strategy.     

The Influence of Snowmobile Trails on Coyote Movements during Winter in High-Elevation Landscapes

Competition between sympatric carnivores has long been of interest to ecologists. Increased understanding of these interactions can be useful for conservation planning. Increased snowmobile traffic on public lands and in habitats used by Canada lynx (Lynx canadensis) remains controversial due to the concern of coyote (Canis latrans) use of snowmobile trails and potential competition with lynx. Determining the variables influencing coyote use of snowmobile trails has been a priority for managers attempting to conserve lynx and their critical habitat. During 2 winters in northwest Wyoming, we backtracked coyotes for 265 km to determine how varying snow characteristics influenced coyote movements; 278 km of random backtracking was conducted simultaneously for comparison. Despite deep snow (>1 m deep), radio-collared coyotes persisted at high elevations (>2,500 m) year-round. All coyotes used snowmobile trails for some portion of their travel. Coyotes used snowmobile trails for 35% of their travel distance (random: 13%) for a mean distance of 149 m (random: 59 m). Coyote use of snowmobile trails increased as snow depth and penetrability off trails increased. Essentially, snow characteristics were most influential on how much time coyotes spent on snowmobile trails. In the early months of winter, snow depth was low, yet the snow column remained dry and the coyotes traveled off trails. As winter progressed and snow depth increased and snow penetrability increased, coyotes spent more travel distance on snowmobile trails. As spring approached, the snow depth remained high but penetrability decreased, hence coyotes traveled less on snowmobile trails because the snow column off trail was more supportive. Additionally, coyotes traveled closer to snowmobile trails than randomly expected and selected shallower snow when traveling off trails. Coyotes also preferred using snowmobile trails to access ungulate kills. Snow compaction from winter recreation influenced coyote movements within an area containing lynx and designated lynx habitat.     

The effects of mate separation on pair re-formation in the Texas cichlid fish (Cichlasoma cyanoguttatum)

This study tested whether mated pairs of Texas cichlid ($\text{Cichlasoma cyanoquttatum}$) would reform and continue rearing their offspring after being separated for either one, four, or ten days. All pairs successfully re-formed after one day, while only 50% re-formed after 4 days. No pair re-formed after 10 days of separation. In successfully re-formed pairs, the female was always more aggressive than her mate, irrespective of whether the female was the resident or the returning individual. Aggressiveness seemed related to the female's attempt to control the male's parental behavior. For pairs which did not re-form, the resident parent, either the male or female, violently and continously attacked the returned mate.     

Male Defense of the Breeding Cavity and Factors Affecting the Persistence of Breeding Pairs in the Stomatopod,Gonodactylus bredini (Manning) (Crustacea: Hoplocarida)

In Caribbean Panama, nonreproductive male and female stomatopods are solitary and defend their own coral-rubble cavities. When breeding pairs form, however, males assume all responsibility for cavity defense. To compare success in cavity defense and defensive tactics among paired and unpaired males, and to examine the tendency for paired stomatopods to exchange their present mates for larger (higher quality) individuals, we introduced same-sized and 15% larger male, and same-sized and 15% larger reproductive female intruders to paired and unpaired male residents in a balanced design. Paired males were more successful at cavity defense than unpaired males, evidently because paired males strike intruders more than unpaired males, and because intruders fight less intensely against paired males than against unpaired males. Paired males occasionally attempted extrapair copulations, but showed little tendency to abandon their mates in favor of larger females. Paired females, however, mated readily with intruder males that evicted resident males. Populationwide female breeding synchrony and prolonged female receptivity before oviposition reduce variance in male mating success and may force males to guard the breeding cavity to assure their paternity. Uncertainty about the reproductive condition of intruder females may prevent males from exchanging mates.     

Winter space use of coyotes in high-elevation environments: behavioral adaptations to deep-snow landscapes

In the last century, coyotes (Canis latrans) have expanded their range geographically, but have also expanded their use of habitats within currently occupied regions. Because coyotes are not morphologically adapted for travel in deep snow, we studied coyote space use patterns in a deep-snow landscape to examine behavioral adaptations enabling them to use high elevations during winter. We examined the influence of snow depth, snow penetrability, canopy cover, and habitat type, as well as the rates of prey and predator track encounters, on coyote travel distance in high-elevation terrain in northwestern Wyoming, USA. We backtracked 13 radio-collared coyotes for 265.41 km during the winters of 2006–2007 and 2007–2008, and compared habitat use and movement patterns of the actual coyotes with 259.11 km of random travel paths. Coyotes used specific habitats differently than were available on the landscape. Open woodlands were used for the majority of coyote travel distance, followed by mixed conifer, and closed-stand spruce–fir. Prey track encounters peaked in closed-stand, mature Douglas fir, followed by 50- to 150-year-old lodgepole pine stands, and 0- to 40-year-old regeneration lodgepole pine stands. Snowmobile trails had the most variation between use and availability on the landscape (12.0 % use vs. 0.6 % available). Coyotes increased use of habitats with dense canopy cover as snow penetration increased and rates of rodent and red squirrel track encounters increased. Additionally, coyotes spent more time in habitats containing more tracks of ungulates. Conversely, use of habitats with less canopy cover decreased as snow depth increased, and coyotes traveled more directly in habitats with less canopy cover and lower snow penetration, suggesting coyotes used these habitats to travel. Coyotes persisted throughout the winter and effectively used resources despite deep snow conditions in a high-elevation environment.     

Male Mating Success in the Presence of a Conspecific Opponent in a Madagascar Hissing Cockroach,Gromphadorhina portentosa (Dictyoptera: Blaberidae)

The influence of a conspecific competitor on male mating behavior was examined in a Madagascar hissing cockroach, Gromphadorhina portentosa. Previous studies have suggested that both male-male competition and female discrimination during courtship interactions may influence male mating success. Familiar pairs of males with a known social association were placed in an arena with a single virgin female and observed. As expected, subordinate males mated significantly less often than their dominant opponents. In pairs in which one male mated, dominant individuals limited the access of subordinates to females. Dominant males displayed an increased frequency and duration of interaction with the female. However, in pairs where both males remained unmated, the mating behavior of dominant and subordinate males did not differ significantly. As interactions progressed, as in the case of males that remained unmated, subordinate males gained increased access to the female. Mated males tended to be larger than their opponent although within a rank, males that mated were no larger than those that remained unmated. These results are discussed in light of the possible roles of male-male competition and female discrimination during courtship interactions.     

Travel distance and mass gain in wintering blackbirds

Birds that range over a large area will have a greater mass-dependent risk of predation than more sedentary birds. Birds that travel more may then reduce winter mass gain to compensate for the increased predation risk that greater travelling entails. I tested whether European blackbirds, Turdus merula, that travelled more in winter had a lower mass than more sedentary birds, independently of any confounding effects of food supply on both ranging behaviour and mass gain. I measured change in winter mass and amount of food eaten in conjunction with the distance that blackbirds travelled to a randomly sited mobile feeder. Blackbirds that travelled shorter distances (per trip and in total) and less often to the feeder had the highest mass midwinter relative to their spring mass. Blackbirds with a higher mean mass midwinter also travelled, on average, shorter distances to the feeder. The distance an individual blackbird travelled to the feeder at any one time was probably independent of the state of its daily energy reserves (how much of its daily total mass gain it had achieved at that point). The relationship between distance travelled and mass was probably independent of food supply because distances actually increased at the end of the winter and the amount of food eaten per individual changed little. More mobile blackbirds were therefore likely to have compensated for any increases in predation risk associated with their greater ranges by decreasing winter mass gain, but will have had an increased risk of starvation because of their lower mass. Copyright 1999 The Association for the Study of Animal Behaviour.     

江西鄱阳湖流域中华秋沙鸭越冬期间的集群特征

2010年11月—2011年3月,采用样线法对江西鄱阳湖流域8个河段的中华秋沙鸭Mergus squamatus的集群特征进行了研究。中华秋沙鸭的集群类型包括雄性群、雌性群、混合群、雌性孤鸭和雄性孤鸭5种类型。越冬期间共记录到89群次,432只次中华秋沙鸭。其中,混合群是最多的一种集群方式,孤鸭也是越冬期间出现频次较高的一种特殊的集群方式。集群类型存在时间变化(P<0.01)。混合群中,大多数群体中雌性个体数多于雄性个体或者是一雌一雄。89群次中,孤鸭和2—8只群占总群数的87.64%,提示中华秋沙鸭主要以集小群分散活动。中华秋沙鸭越冬期间的群体大小是(4.85±4.97)只/群。不同集群类型的群体大小差异较大。集群大小可能与采砂、捕鱼、食物的丰富度等环境因子有关。调查结果显示,中华秋沙鸭越冬期间的性比是1∶0.74(n=432)。中华秋沙鸭群体大小与性比有一定的关系。     

Kin recognition and adjustment of reproductive effort in zebra finches

The differential allocation theory predicts that females should invest more in offspring produced with attractive partners, and a number of studies support this prediction in birds. Females have been shown to increase reproductive investment when mated to males showing elaborated sexual traits. However, mate attractiveness might also depend on the interaction between male and female genotypes. Accordingly, females should invest more in offspring sired by individuals that are genetically dissimilar or carry superior alleles. Here, we show in zebra finches (Taeniopygia guttata) that pairs of unfamiliar genetic brothers and sisters are less likely to reproduce in comparison with randomly mated pairs. Among the brother–sister pairs, those that attempted to breed laid smaller clutches and of lower total clutch mass. Our results provide the first experimental evidence that females adjust their reproductive effort in response to the genetic similarity of their partners. Importantly, these results imply a female ability to assess relatedness of a social mate without prior association.     

Effects of Pair-Bond and Social Context on Male–Female Interactions in Captive Titi Monkeys (Callicebus moloch, Primates: Cebidae)

In monogamous species, an abiding relationship between a specific adult male and a specific adult female is a defining feature of the social system. The interactions between these individuals are influenced by many factors, including not only the history of their relationship (for example, development of a mutual bond), but also the immediate effects of the prevailing social context (for example, presence and sex of extra‐pair conspecifics). In this study we examined the effects of an existing bond and of social context on interactions between adult heterosexual pairs of the monogamous titi monkeys (Callicebus moloch). Twelve adult males and 12 adult females were tested with their cagemates and with an unfamiliar partner of the opposite sex in five social contexts: (1) mated male–female pair; (2) unfamiliar pair; (3) single female; (4) single male; and (5) empty stimulus cage. Results show that mated pairs were more affiliative than unfamiliar pairs and differentiated social contexts more sharply. Males were more responsive to context than females. Distance between mates was less and physical contact was more frequent in the presence of male–female pairs or a single male, than in the presence of a single female or an empty cage. These findings suggest that the presence and sex of strangers have a stronger influence on male–female interactions when the pair has an existing relationship.