Do Conversational Gutturals Help Florida Scrub‐Jays Coordinate Their Sentinel Behavior?

Florida scrub‐jays, Aphelocoma coerulescens, perform sentinel behavior in which individuals alternate bouts of watchfulness with little overlap. We examined how calls might facilitate sentinel coordination. Small soft calls labeled conversational gutturals were heard more often from sentinels than from foraging birds. Calls occurred infrequently throughout sentinel bouts but were more common later in bouts. The pattern of calling does not match predictions for a ‘watchman’s song’ at regular intervals nor for a signal at the end of a sentinel bout. Thus, our quantitative assessment of calling by sentinels did not find support for either standard hypothesis. Although Florida scrub‐jays clearly have information on each other’s sentinel behavior, our results suggest that calls provide perhaps a fraction of this information.     

Task‐Dependent Differences in Learning by Subordinate and Dominant Wild Arabian Babblers

Learning and innovation abilities have been studied extensively in flocking birds, but their importance and relevance in cooperatively breeding birds have been relatively unexplored. We studied the acquisition of novel foraging skills in 14 groups of wild, cooperatively breeding Arabian babblers (Turdoides squamiceps). While in a previous study we found that subordinate individuals were usually the first to learn to remove black rubber lids from a foraging grid, here we show that dominant individuals were the first to succeed in shifting from these black rubber lids to newly introduced white rubber lids. We also found that in all groups where one forager learned to shift to the white lids, the rest of the foragers also learned to do so, suggesting that this behaviour may be transmitted among group members. Although dominant individuals were almost always the first to remove white lids, once starting to remove white lids, dominants and subordinates learned equally well to prefer white over black lids based on differential reinforcement (food was provided only under white lids). Together with our previous study, our results suggest that differences in learning between dominants and subordinates may be task‐specific, which may represent different cognitive strategies: subordinates may explore a more diverse range of foraging opportunities, while dominants may be better at generalizing from familiar tasks to similar ones.     

Effects of habitat complexity,dominance and personality on habitat selection: Ideal despotic cichlids

Habitat structure can impede visibility and movement, resulting in lower resource monopolization and aggression. Consequently, dominant individuals may prefer open habitats to maximize resource gain, or complex habitats to minimize predation risk. We explored the role of dominance on foraging, aggression and habitat choice using convict cichlids (Amatitlania nigrofasciata) in a two‐patch ideal free distribution experiment. Groups of six fish of four distinct sizes first competed for shrimp in one‐patch trials in both an open and complex habitat; half the groups experienced each habitat type first. Following these one‐patch trials, each group then chose between habitat types in a two‐patch trial while competing for food. Finally, each fish underwent an individual behavioural assessment using a battery of “personality” tests to determine if behaviour when alone accurately reflected behaviour within a social context. In the one‐patch trials, dominant fish showed similar food consumption between habitats, but chased more in the complex habitat. In the two‐patch choice trials, dominants preferred and defended the complex habitat, forming an ideal despotic distribution with more than half the fish and competitive weight in the open habitat. Within the groups, individual fish differed in foraging and chasing, with repeatabilities of 0.45 and 0.23 across all treatments. Although a higher foraging rate during the individual assessment predicted foraging rate and use of the complex habitat during the group trials, aggression and boldness tests were not reflective of group behaviour. Across groups, heavier dominants and those with higher foraging rate in the open habitat used the open habitat more, suggesting that both risk and energetic state affect habitat preference in dominant convict cichlids.     

The effect of social dominance on fattening and food-caching behaviour in Carolina chickadees, Poecile carolinensis

Subordinates often have to wait for dominants to obtain food. As a result, their foraging success should be less predictable and they should therefore maintain a higher level of energy reserves compared with dominants. A corollary of this prediction is that subordinates should gain mass earlier in the day and maintain higher mass than dominants. We tested these predictions with captive Carolina chickadees. In two different experiments (one where birds were given ad libitum access to food and the other with food access limited to 60 min/day), we formed social flocks of two previously unfamiliar birds and compared their energy management (body fat and food caches) while they were in the flock with energy management when housed alone. Results from both experiments failed to support the predictions. Of all the parameters of body mass and food caching we measured only the following results were significant: (1) On the ad libitum food schedule, both subordinates and dominants accumulated more mass over the day when in a flock compared with when they were solitary, and there were no differences in mass gain between dominants and subordinates. (2) When analysed separately, dominants showed a higher evening mass in the flock compared with the solitary condition, a trend that runs opposite to the prediction. Our results suggest that when in favourable foraging conditions, social interactions might cause dominant and subordinate birds to accumulate more energy reserves as a result of competition. On the other hand, if food supply is limited, both dominants and subordinates may be forced to maintain similar fat reserves as an insurance against increased risk of starvation. Copyright 2000 The Association for the Study of Animal Behaviour.     

Benefits and Costs of Dominance in the Angelfish Centropyge bicolor

Social groups are often structured by dominance hierarchies in which subordinates consistently defer to dominants. High‐ranking individuals benefit by gaining inequitable access to resources, and often achieve higher reproductive success; but may also suffer costs associated with maintaining dominance. We used a large‐scale field study to investigate the benefits and costs of dominance in the angelfish Centropyge bicolor, a sequential hermaphrodite. Each haremic group contains a single linear body size‐based hierarchy with the male being most dominant, followed by several females in descending size order. Compared to their subordinate females, dominant males clearly benefited from disproportionately high spawning frequencies, but bore costs in lower foraging rates and greater aggressive defence of their large territories. Within the female hierarchy, more dominant individuals benefited from higher spawning frequencies and larger home ranges, but displayed neither higher foraging rates nor spawn order priority. However, dominance in females was also linked to aggressiveness, particularly towards immediate subordinates, suggesting that females were using energetically costly aggression to maintain their high rank. We further showed by experimentally removing dominant females that the linear hierarchy was also a social queue, with subordinates growing to inherit higher rank with its attendant benefits and costs when dominants disappeared. We suggest that in C. bicolor, the primary benefit of high rank is increased reproductive success in terms of current spawning frequency and the prospect of inheriting the male position in the near future, which may be traded off against the cost of aggressively defending rank and territory.     

Effect of the Presence of Subordinates on Dominant Female Behaviour and Fitness in Hierarchies of the Dwarf Angelfish Centropyge bicolor

In many hierarchical animal societies, dominant individuals control group membership owing to their power to evict subordinates. In such groups, the presence of subordinates, and therefore group stability, is continually dependent on subordinates being tolerated by dominants. The dominant decision to tolerate or evict is, in turn, dependent on the costs and benefits to dominants of subordinate presence. We investigated the effect of subordinate presence on dominants in the female dominance hierarchy of the dwarf angelfish Centropyge bicolor, using both observations of natural groups and experimental removals of subordinates. We found that the presence of subordinates had no effect on dominant access to resources, as measured by dominant foraging rates and home range areas, nor on dominant fitness, as measured by growth rates and spawning frequencies. Our results suggest that the presence of subordinates has a neutral effect on the current fitness of dominants, so that dominants have no great incentive to evict subordinates. We discuss the possibility that tolerance of subordinates might be further explained by considering future fitness, as dominant females in these haremic protogynous angelfish stand to inherit the male position, whereupon subordinate females change from potential competition to useful mates.     

Vocal Coordination of Group Movement by Green Woodhoopoes (Phoeniculus purpureus)

Communication signals are used by many species to maintain group cohesion when moving over larger areas. Groups of green woodhoopoes (Phoeniculus purpureus) generally move around their territory as a close‐knit unit. Dominant individuals were more likely than subordinates to initiate movement to a new foraging site, but there was no intersexual difference. Dominants were also more likely than subordinates to be followed immediately. Vocalizations were shown to play an important role in mobilization: in the thick forests inhabited by woodhoopoes, visual cues to coordinate movement are likely to be less successful. When responding to the rallying call of a neighboring group, dominants and subordinates were equally likely to lead, as were males and females. As other group members followed immediately on most of these occasions, vocalizations were less important in this context than when moving to a new foraging site.     

Incubation failure and nest abandonment by Leach's Storm‐Petrels detected using PIT tags and temperature loggers

ABSTRACT The nocturnal activity of burrow‐nesting seabirds, such as storm‐petrels and shearwaters, makes it difficult to study their incubation behavior. In particular, little is known about possible differences in the incubation behavior of adults at successful and unsuccessful nests. We combined the use of passive integrated transponder (PIT) technology and nest‐temperature data loggers to monitor the incubation behavior of 10 pairs of Leach's Storm‐Petrels (Oceanodroma leucorhoa). The mean incubation bout length was 3.31 ± 0.59 (SD) days for individual adults at successful nests (N= 4) and 1.84 ± 1.16 d for individuals at unsuccessful nests (N= 6). Mean bout length for pairs in successful burrows (3.51 ± 0.56 d) did not differ significantly (P= 0.07) from that for pairs in unsuccessful burrows (1.80 ± 1.20 d), perhaps due to one failed nest with a high mean bout length (4.15 d). The total number of incubation bouts per parent (4.3 ± 1.9 bouts) did not differ with hatching success. Adults whose nests failed repeatedly exhibited truncated incubation bouts (< 12 h) prior to complete nest failure and were more likely than successful parents to make brief visits to nearby, occupied nesting burrows. Our results suggest that the decision by Leach's Storm‐Petrels to abandon a nest is not an abrupt one. Rather, failed nesting attempts may be characterized by truncated incubation bouts where individuals pay the energetic cost of travel to and from the burrow, but do not remain long enough to successfully incubate the egg.     

Within‐group relatedness is correlated with colony‐level social structure and reproductive sharing in a social fish

In group‐living species, the degree of relatedness among group members often governs the extent of reproductive sharing, cooperation and conflict within a group. Kinship among group members can be shaped by the presence and location of neighbouring groups, as these provide dispersal or mating opportunities that can dilute kinship among current group members. Here, we assessed how within‐group relatedness varies with the density and position of neighbouring social groups in Neolamprologus pulcher, a colonial and group‐living cichlid fish. We used restriction site‐associated DNA sequencing (RADseq) methods to generate thousands of polymorphic SNPs. Relative to microsatellite data, RADseq data provided much tighter confidence intervals around our relatedness estimates. These data allowed us to document novel patterns of relatedness in relation to colony‐level social structure. First, the density of neighbouring groups was negatively correlated with relatedness between subordinates and dominant females within a group, but no such patterns were observed between subordinates and dominant males. Second, subordinates at the colony edge were less related to dominant males in their group than subordinates in the colony centre, suggesting a shorter breeding tenure for dominant males at the colony edge. Finally, subordinates who were closely related to their same‐sex dominant were more likely to reproduce, supporting some restraint models of reproductive skew. Collectively, these results demonstrate that within‐group relatedness is influenced by the broader social context, and variation between groups in the degree of relatedness between dominants and subordinates can be explained by both patterns of reproductive sharing and the nature of the social landscape.     

The Effect of Social Hierarchy on Captive Coyote (Canis latrans) Foraging Behavior

Foraging efficiency of individuals in pack forming species may be influenced by social dynamics within a pack. The effects of social hierarchy in particular may influence individual foraging behavior in canids, such as coyotes (Canis latrans). To examine the impact of social hierarchy on foraging behavior, we tested 16 captive coyotes in eight naturally established dominant–subordinate pairs, using the guesser–knower paradigm. We measured the efficiency of subordinate coyotes to relocate a food resource when alone and then allowed pairs to forage together, such that subordinates had prior knowledge of food location but dominants did not. To determine whether (1) subordinates used a direct or discursive strategy to obtain food in the presence of a dominant and (2) dominants used an exploitative or independent strategy to obtain food in the presence of a subordinate with previous knowledge, we measured their search efficiency (e.g., correct choice of area, feeder, and latency to correct feeder). Results showed subordinates learned to relocate food and increase efficiency when alone. In a social context, however, subordinate efficiency decreased. That is, subordinates approached the correct area, but searched more feeders before finding the correct one. Dominants initially used an independent search strategy but then quickly displaced the subordinate and monopolized the resource, reducing subordinate efficiency further. Despite continual displacement and reduction in efficiency, subordinates did not alter their foraging strategy over time. Our results suggest prior information can improve individual foraging advantage, but that social status strongly impacts individual foraging efficiency in social species such as coyotes.     

Aggression Levels Affect Social Interaction in the Non‐Breeding Territorial Aggression of the Weakly Electric Fish,Gymnotus omarorum

Animals typically decide whether to fight or retreat from conspecifics based on their individual estimates of the costs and benefits of fighting. Theoretical models predict how contenders solve a conflict, but the evaluation processes involved in these decisions depend on multiple factors that are difficult to explore experimentally. We addressed these questions using the non‐breeding territorial aggression of Gymnotus omarorum, in which subordinates make three distinctive decisions to signal their submission during a fight: (1) interruption of their electric discharges to hide from the dominant, (2) stop attacking and retreat, and (3) emission of ‘chirps’, transient submissive electric signals. We confirmed that subordinates take into account the aggressive performance of dominants to shape their own agonistic decisions and performance. The intensity of aggression is highly correlated with an agonistic dyad, and the decision of subordinates to retreat is influenced by the attack rates of dominants. When we lowered the aggression of expected dominants with a 5‐HT_1A receptor agonist, the correlation between the two contenders' aggression levels was lost and subordinates completely stopped emitting electric chirp signals. The aforementioned results contribute to the understanding of the decision‐making strategies driven by social challenge inherent to agonistic encounters.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Effects of reproductive condition,roost microclimate,and weather patterns on summer torpor use by a vespertilionid bat

A growing number of mammal species are recognized as heterothermic, capable of maintaining a high‐core body temperature or entering a state of metabolic suppression known as torpor. Small mammals can achieve large energetic savings when torpid, but they are also subject to ecological costs. Studying torpor use in an ecological and physiological context can help elucidate relative costs and benefits of torpor to different groups within a population. We measured skin temperatures of 46 adult Rafinesque's big‐eared bats (Corynorhinus rafinesquii) to evaluate thermoregulatory strategies of a heterothermic small mammal during the reproductive season. We compared daily average and minimum skin temperatures as well as the frequency, duration, and depth of torpor bouts of sex and reproductive classes of bats inhabiting day‐roosts with different thermal characteristics. We evaluated roosts with microclimates colder (caves) and warmer (buildings) than ambient air temperatures, as well as roosts with intermediate conditions (trees and rock crevices). Using Akaike's information criterion (AIC), we found that different statistical models best predicted various characteristics of torpor bouts. While the type of day‐roost best predicted the average number of torpor bouts that bats used each day, current weather variables best predicted daily average and minimum skin temperatures of bats, and reproductive condition best predicted average torpor bout depth and the average amount of time spent torpid each day by bats. Finding that different models best explain varying aspects of heterothermy illustrates the importance of torpor to both reproductive and nonreproductive small mammals and emphasizes the multifaceted nature of heterothermy and the need to collect data on numerous heterothermic response variables within an ecophysiological context.     

Sequential foraging of dusky dolphins with an inspection of their prey distribution

The aim of this work was to analyze the sequential foraging behavior of dusky dolphins (Lagenorhynchus obscurus). Foraging sequences were defined when more than two feeding bouts occur with a traveling bout between them. We hypothesized that traveling costs of searching for prey patches were related to the time spent feeding on a patch. In addition, the distribution and seasonal variation of anchovy schools were studied in the area to better understand dolphins' behavior. We observed dolphins from a research vessel from 2001 to 2007, and recorded their location and behavior. Anchovy data were collected during two hydro‐acoustic surveys. Dusky dolphin behaviors varied seasonally; they spent a greater proportion of time traveling and feeding in the warm season (Kruskal‐Wallis: H = 172.07, P < 0.01). During the cold season dolphin groups were more likely to exhibit diving behavior and less surface feeding. We found a positive correlation between searching and foraging time (r = 0.88, P = 0.019), suggesting that the costs associated with searching were compensated by an increase in the energy intake during the foraging bout. There was an association between dusky dolphin and anchovy distribution, in that they co‐varied spatially and seasonally.     

Energetic solutions of Rock Sandpipers to harsh winter conditions rely on prey quality

Rock Sandpipers Calidris ptilocnemis have the most northerly non‐breeding distribution of any shorebird in the Pacific Basin (upper Cook Inlet, Alaska; 61°N, 151°W). In terms of freezing temperatures, persistent winds and pervasive ice, this site is the harshest used by shorebirds during winter. We integrated physiological, metabolic, behavioural and environmental aspects of the non‐breeding ecology of Rock Sandpipers at the northern extent of their range to determine the relative importance of these factors in facilitating their unique non‐breeding ecology. Not surprisingly, estimated daily energetic demands were greatest during January, the coldest period of winter. These estimates were greatest for foraging birds, and exceeded basal metabolic rates by a factor of 6.5, a scope of increase that approaches the maximum sustained rate of energetic output by shorebirds during periods of migration, but far exceeds these periods in duration. We assessed the quality of their primary prey, the bivalve Macoma balthica, to determine the daily foraging duration required by Rock Sandpipers to satisfy such energetic demands. Based on size‐specific estimates of M. balthica quality, Rock Sandpipers require over 13 h/day of foraging time in upper Cook Inlet in January, even when feeding on the highest quality prey. This range approaches the average daily duration of mudflat availability in this region (c. 18 h), a maximum value that annually decreases due to the accumulation of shore‐fast ice. Rock Sandpipers are likely to maximize access to foraging sites by following the exposure of ice‐free mudflats across the upper Cook Inlet region and by selecting smaller, higher quality M. balthica to minimize foraging times. Ultimately, this unusual non‐breeding ecology relies on the high quality of their prey resources. Compared with other sites across their range, M. balthica from upper Cook Inlet have relatively light shells, potentially the result of the region's depauperate invertebrate predator community. Given the delicate balance between environmental and prey conditions that currently make Cook Inlet a viable wintering area for Rock Sandpipers, small variations in these variables may affect the suitability of the site in the future.     

Social rank‐dependent expression of arginine vasotocin in distinct preoptic regions in male Oryzias latipes

This study examined arginine vasotocin (AVT) expression in the brains of dominant and subordinate male medaka Oryzias latipes after short‐ and long‐term competition. High AVT expression in distinct preoptic regions was found in dominants and subordinates within minutes of encountering each other. During long‐term competition, AVT expression remained high in dominants but not in subordinates.     

Advancing breeding phenology does not affect incubation schedules in chestnut‐crowned babblers: Opposing effects of temperature and wind

Projecting population responses to climate change requires an understanding of climatic impacts on key components of reproduction. Here, we investigate the associations among breeding phenology, climate and incubation schedules in the chestnut‐crowned babbler (Pomatostomus ruficeps), a 50 g passerine with female‐only, intermittent incubation that typically breeds from late winter (July) to early summer (November). During daylight hours, breeding females spent an average of 33 min on the nest incubating (hereafter on‐bouts) followed by 24‐min foraging (hereafter off‐bouts), leading to an average daytime nest attentiveness of 60%. Nest attentiveness was 25% shorter than expected from allometric calculations, largely because off‐bout durations were double the expected value for a species with 16 g clutches (4 eggs × 4 g/egg). On‐bout durations and daily attentiveness were both negatively related to ambient temperature, presumably because increasing temperatures allowed more time to be allocated to foraging with reduced detriment to egg cooling. By contrast, on‐bout durations were positively associated with wind speed, in this case because increasing wind speed exacerbated egg cooling during off‐bouts. Despite an average temperature change of 12°C across the breeding season, breeding phenology had no effect on incubation schedules. This surprising result arose because of a positive relationship between temperature and wind speed across the breeding season: Any benefit of increasing temperatures was canceled by apparently detrimental consequences of increasing wind speed on egg cooling. Our results indicate that a greater appreciation for the associations among climatic variables and their independent effects on reproductive investment are necessary to understand the effects of changing climates on breeding phenology.     

From facilitation to competition: temperature‐driven shift in dominant plant interactions affects population dynamics in seminatural grasslands

Biotic interactions are often ignored in assessments of climate change impacts. However, climate‐related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co‐occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad‐scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population‐level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.     

Quality versus quantity: Foraging decisions in the honeybee (Apis mellifera scutellata) feeding on wildflower nectar and fruit juice

Foraging animals must often decide among resources which vary in quality and quantity. Nectar is a resource that exists along a continuum of quality in terms of sugar concentration and is the primary energy source for bees. Alternative sugar sources exist, including fruit juice, which generally has lower energetic value than nectar. We observed many honeybees (Apis mellifera scutellata) foraging on juice from fallen guava (Psidium guajava) fruit near others foraging on nectar. To investigate whether fruit and nectar offered contrasting benefits of quality and quantity, we compared honeybee foraging performance on P. guajava fruit versus two wildflowers growing within 50 m, Richardia brasiliensis and Tridax procumbens. Bees gained weight significantly faster on fruit, 2.72 mg/min, than on either flower (0.17 and 0.12 mg/min, respectively). However, the crop sugar concentration of fruit foragers was significantly lower than for either flower (12.4% vs. 37.0% and 22.7%, respectively). Fruit foragers also spent the most time handling and the least time flying, suggesting that fruit juice was energetically inexpensive to collect. We interpret honeybee foraging decisions in the context of existing foraging models and consider how nest‐patch distance may be a key factor for central place foragers choosing between resources of contrasting quality and quantity. We also discuss how dilute solutions, such as fruit juice, can help maintain colony sugar–water balance. These results show the benefits of feeding on resources with contrasting quality and quantity and that even low‐quality resources have value.