Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.Communicated by M. Zuk     

Echolocation and foraging behavior of the lesser bulldog bat, Noctilio albiventris : preadaptations for piscivory?

We studied variability in foraging behavior of Noctilio albiventris (Chiroptera: Noctilionidae) in Costa Rica and Panamá and related it to properties of its echolocation behavior. N. albiventris searches for prey in high (>20 cm) or low (<20 cm) search flight, mostly over water. It captures insects in mid-air (aerial captures) and from the water surface (pointed dip). We once observed an individual dragging its feet through the water (directed random rake). In search flight, N. albiventris emits groups of echolocation signals (duration 10–11 ms) containing mixed signals with constant-frequency (CF) and frequency-modulated (FM) components, or pure CF signals. Sometimes, mostly over land, it produces long FM signals (duration 15–21 ms). When N. albiventris approaches prey in a pointed dip or in aerial captures, pulse duration and pulse interval are reduced, the CF component is eliminated, and a terminal phase with short FM signals (duration 2 ms) at high repetition rates (150–170 Hz) is emitted. Except for the last pulses in the terminal phase N. albiventris avoids overlap between emitted signals and echoes returning from prey. During rakes, echolocation behavior is similar to that in high search flight. We compare N. albiventris with its larger congener, N. leporinus, and discuss behavioral and morphological specializations that can be interpreted as preadaptations favoring the evolution of piscivory as seen in N. leporinus. Prominent among these specializations are the CF components of the echolocation signals which allow detection and evaluation of fluttering prey amidst clutter-echoes, high variability in foraging strategy and the associated echolocation behavior, as well as morphological specializations such as enlarged feet for capturing prey from the water surface. Received: 21 April 1997 / Accepted after revision: 12 January 1998     

Foraging tactics of an ambush predator: the effects of substrate attributes on prey availability and predator feeding success

The foraging sites selected by an ambush forager can strongly affect its feeding opportunities. Foraging cane toads (Rhinella marina) typically select open areas, often under artificial lights that attract insects. We conducted experimental trials in the field, using rubber mats placed under lights, to explore the influence of substrate color and rugosity on prey availability (numbers, sizes, and types of insects) and toad foraging success. A mat's color (black vs. white) and rugosity (smooth vs. rough) did not influence the numbers, sizes, or kinds of insects that were attracted to it, but toads actively preferred to feed on rugose white mats (50% of prey-capture events, vs. a null of 25%). White backgrounds provided better visual contrast of the (mostly dark) insects, and manipulations of prey color in the laboratory showed that contrast was critical in toad foraging success. Insects landing on rugose backgrounds were slower to leave, again increasing capture opportunities for toads. Thus, cane toads actively select backgrounds that maximize prey-capture opportunities, a bias driven by the ways that substrate attributes influence ease of prey detection and capture rather than by absolute prey densities.     

Trophic segregation between sexes in the Black Skimmer revealed through the analysis of stable isotopes

The degree of individual or gender variation when exploiting food resources is an important aspect in the study of foraging ecology within a population. Previous information on non-breeding skimmers obtained through conventional methodologies suggested sex-related differences in prey species. In this study, stable isotope techniques were used to investigate the intraspecific segregation in diet and foraging habits of the Black Skimmer (Rynchops niger intercedens) at Mar Chiquita Coastal Lagoon (37°40′S, 57°22′W), Argentina. These results were compared with contemporary data on the trophic composition obtained by conventional methodologies. Blood samples were taken from birds captured with mist-nets during their non-breeding season. The isotopic signatures of skimmers showed a diet mainly composed of marine prey with some degree of estuarine fish intake. When comparing diet between sexes, males showed enrichment in ¹⁵N compared to females, while no differences were observed in ¹³C. The use of mixing models revealed differences in the relative composition of prey in the diet of male and female skimmers. This study highlights stable isotope analysis as a valuable tool to test inter-individual differences and sexual segregation in trophic ecology of Black Skimmers as compared to conventional methodologies. The results show a trophic segregation in the Black Skimmer during the non-breeding season that can be explained by differences in prey species and larger prey sizes of male skimmers. Our findings have significant implications for conservation since any environmental change occurring at wintering areas might have profound effects on several avian life-history traits, and could be different for males and females due to trophic segregation.     

Network analysis reveals contrasting effects of intraspecific competition on individual vs. population diets

Optimal foraging theory predicts that individuals should become more opportunistic when intraspecific competition is high and preferred resources are scarce. This density-dependent diet shift should result in increased diet breadth for individuals as they add previously unused prey to their repertoire. As a result, the niche breadth of the population as a whole should increase. In a recent study, R. Svanb?ck and D. I. Bolnick confirmed that intraspecific competition led to increased population diet breadth in threespine stickleback (Gasterosteus aculeatus). However, individual diet breadth did not expand as resource levels declined. Here, we present a new method based on complex network theory that moves beyond a simple measure of diet breadth, and we use the method to reexamine the stickleback experiment. This method reveals that the population as a whole added new types of prey as stickleback density was increased. However, whereas foraging theory predicts that niche expansion is achieved by individuals accepting new prey in addition to previously preferred prey, we found that a subset of individuals ceased to use their previously preferred prey, even though other members of their population continued to specialize on the original prey types. As a result, populations were subdivided into groups of ecologically similar individuals, with diet variation among groups reflecting phenotype-dependent changes in foraging behavior as prey density declined. These results are consistent with foraging theory if we assume that quantitative trait variation among consumers affects prey preferences, and if cognitive constraints prevent individuals from continuing to use their formerly preferred prey while adding new prey.     

Demographic and temporal structure of an allele frequency cline in the mussel Mytilus edulis

Feeding rates, patterns of prey selection, and starvation tolerance were investigated for adult males and females of the cyclopoid copepod Corycaeus anglicus collected from the waters of Friday Harbor, Washington, USA. Selection by C. anglicus was determined largely by prey body-size, but was also affected by species and developmental stage. Small developmental stages of all prey species were fed upon at relatively low rates. The small calanoid species Acartia clausii was increasingly vulnerable to predation by C. anglicus as it progressed through successive developmental stages. Larger prey species, Pseudocalanus sp. and Calanus pacificus, were more vulnerable in intermediate stages, the C3 and N6 stages, respectively. Larger and smaller prey were characteristically attacked at different sites on their bodies; however, attack sites fell within a similar range of body widths, 130 to 170 m. Males of Corycaeus anglicus killed a maximum of 1.4 prey d^-1 when feeding on the optimally-sized adult females of Acartia clausii, which are approximately equivalent to its own body length. Males fed at approximately double the rates of females. Despite its small size and apparent lack of metabolic stores, this cyclopoid is highly tolerant of starvation conditions. Median survival time without food is at least 2 wk for both males and females. In its predatory behavior, C. anglicus employs an ambush-type strategy and seems to be adapted for infrequent encounters with relatively large prey.Contribution No. 1412 from the School of Oceanography, University of Washington, Seattle     

Pursuit-deterrent communication between prey animals and timber rattlesnakes (Crotalus horridus): the response of snakes to harassment displays

A thorough understanding of communication requires an evaluation of both the signaler and receiver. Most analyses of prey–predator communication are incomplete because they examine only the behavior of the prey. Predators in these systems may be understudied because they are perceived as less tractable research subjects, due to their more cryptic hunting behaviors and secretive lifestyles. For example, research on interactions between rodents and rattlesnakes has focused on the behavior of rodent signalers, while responses of snakes have been virtually unexamined. Rattlesnakes are ambush predators, and capture rodents by waiting at foraging sites for long periods of time. In this study, I take advantage of the sedentary nature of this foraging strategy and use fixed videography to record natural encounters between timber rattlesnakes (Crotalus horridus) and their prey. Three different prey species were found to exhibit conspicuous visual displays to snakes, both when snakes were actively foraging, and when they were basking. After receiving displays, foraging snakes left their ambush sites and moved long distances before locating subsequent ambush sites, indicating that they responded to displays by abandoning attempts to ambush prey in the vicinity of signalers. This study represents the first quantitative analysis of the response of free-ranging snakes to signals from their prey, and elucidates a technique by which such quantitative data can be more easily obtained.     

Contrasting home-range size and spatial partitioning in cryptic and sympatric pipistrelle bats

The extent of spatial partitioning in insectivorous bats, whose prey is patchily distributed and transient in nature, remains a contentious issue. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and Pipistrellus pygmaeus, which are morphologically similar and sympatric, provides an opportunity to examine this question. The present study used radio telemetry to address the spatial distribution and foraging characteristics of P. pipistrellus and P. pygmaeus in northeast Scotland, to test the hypothesis that coexistence between these species is facilitated through spatial segregation. We reveal large and significant differences in the spatial distribution and foraging characteristics of these two cryptic species. Individual P. pipistrellus home ranges were on average three times as large as that of P. pygmaeus, and they foraged for approximately an hour longer each night. Inter-specific spatial overlap was minimal (<5%) and core foraging areas of either species were essentially mutually exclusive despite the proximity of the two roosts. Inter-specific differences in range size were associated with the spatial dispersion of productive foraging sites within individual foraging ranges. P. pipistrellus foraging sites were highly dispersed, necessitating larger ranges. It is predicted that the spatial segregation revealed by the present study is a result of selection favouring the avoidance of competition in these species through differential habitat use.     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.     

Competition among penguins and cetaceans reveals trophic cascades in the western Ross Sea, Antarctica

An apparent trophic cascade that appears during summer in the western Ross Sea, Antarctica, explains why the Antarctic silverfish (Pleuragramma antarcticum) there becomes cannibalistic; its principal prey, crystal krill (Euphausia crystallorophias) becomes scarce; and the diatom community is minimally grazed compared to adjacent areas. The krill is the major grazer of diatoms. On the basis of fieldwork at Ross Island, we suggest that the cascade results from foraging by unusually numerous Adélie Penguins (Pygoscelis adeliae), minke whales (Balaenoptera bonaerensis), and fish-eating killer whales (Orcinus orca). These species and other top predators apparently deplete the krill and silverfish. In drawing our conclusions, we were aided by two "natural experiments." In one "experiment," large, grounded icebergs altered the seasonal pattern of change in regional sea-ice cover, but not the seasonal change in penguin diet and foraging behavior that was also detected during the pre-iceberg era. In the other "experiment," a short-term polynya (opening in the ice) brought penguins and whales together in a confined area, this time altering both penguin diet and foraging behavior. We conclude that the foraging of penguins and whales, and not a formerly hypothesized seasonal decrease in sea-ice cover, explains (1) the annual switch in the penguins' prey from krill to silverfish, (2) the subsequent lengthening of penguin foraging trips, and (3) a marked decline of cetaceans in the area later in the season. Reduction in the middle-trophic-level prey is expressed in the relaxed grazing pressure on phytoplankton.     

Linking marine predator diving behavior to local prey fields in contrasting habitats in a subarctic glacial fjord

Foraging theory predicts that animals will adjust their foraging behavior in order to maximize net energy intake and that trade-offs may exist that can influence their behavior. Although substantial advances have been made with respect to the foraging ecology of large marine predators, there is still a limited understanding of how predators respond to temporal and spatial variability in prey resources, primarily due to a lack of empirical studies that quantify foraging and diving behavior concurrently with characteristics of prey fields. Such information is important because changes in prey availability can influence the foraging success and ultimately fitness of marine predators. We assessed the diving behavior of juvenile female harbor seals (Phoca vitulina richardii) and prey fields near glacial ice and terrestrial haulout sites in Glacier Bay (58°40′N, ?136°05′W), Alaska. Harbor seals captured at glacial ice sites dived deeper, had longer dive durations, lower percent bottom time, and generally traveled further to forage. The increased diving effort for seals from the glacial ice site corresponded to lower prey densities and prey at deeper depths at the glacial ice site. In contrast, seals captured at terrestrial sites dived shallower, had shorter dive durations, higher percent bottom time, and traveled shorter distances to access foraging areas with much higher prey densities at shallower depths. The increased diving effort for seals from glacial ice sites suggests that the lower relative availability of prey may be offset by other factors, such as the stability of the glacial ice as a resting platform and as a refuge from predation. We provide evidence of differences in prey accessibility for seals associated with glacial ice and terrestrial habitats and suggest that seals may balance trade-offs between the costs and benefits of using these habitats.     

Movements of vervets (Cercopithecus aethiops) and patas monkeys (Erythrocebus patas) as estimators of food resource size, density, and distribution

The effect of food resources on behavior has been difficult to measure. Here we use animals themselves to describe “effective” food abundance and distribution by comparing, relative to where individuals stopped to eat, movements of (1) adult females living in a small group of vervet monkeys (Cercopithecus aethiops) with those living in a large group and (2) vervets and patas monkeys (Erythrocebus patas). Although females in the large vervet group travelled farther and stopped to eat more often than females in the small vervet group, these differences resulted from foraging in Acacia drepanolobium habitat. In A. xanthophloea habitat, females in the large group travelled less far, travelled shorter distances between foods, and stopped as often as females in the small group. Greater foraging costs of females in larger vervet groups may be offset by access to home ranges of better quality. Compared to patas, vervets travelled shorter distances, moved shorter distances between food sites, stopped less often, and had longer feeding bouts, suggesting that foods of vervets are denser and larger, overall, than foods of patas. When vervets foraged in A. drepanolobium habitat, also the habitat of patas, their foraging behavior became more like that of patas. Vervets travelled farther, stopped more often, and spent less time at food sites in A. drepanolobium habitat than in A. xanthophloea habitat, suggesting that foods are smaller and less usurpable in A. drepanolobium habitat. Distance between foods, a component of food distribution, did not increase, however. The critical variable underlying usurpability of foods may be food site depletion time, a temporal measure. Received: 14 March 1997 / Accepted after revision: 19 October 1997     

Underwater and above-water search patterns of an Arctic seabird: reduced searching at small spatiotemporal scales

How predators vary search patterns in response to prey predictability is poorly known. For example, marine invertebrates may be predictable but of low energy value, while fish may be of higher energy value but unpredictable at large (pelagic schools) or small (solitary benthics) spatial scales. We investigated the search patterns of the thick-billed murre (Uria lomvia), an Arctic seabird feeding on invertebrates, pelagic fish, or benthic fish. Foraging ranges at the Coats Island colony are generally smaller (<240 min per trip) than at larger colonies, and many birds specialize in foraging tactics and diet. Underwater search times for benthic fish were higher than for pelagic fish or invertebrates while above-water search times for pelagic fish were higher than for benthic fish or invertebrates. There were few stops during trips. Total trip time, flying time, number of flights, and number of dives were intercorrelated and increased with prey energy content, suggesting that longer trips involved fewer prey encounters due to selection of higher-quality, but rarer, prey items. Flight times were not Lévy-distributed and seabirds may have used area-restricted searches. The high degree of specialization, apparent absence of information center effects, and reduced above-water searching times may be linked to the relatively small colony size and the resulting short commuting distances to feeding areas, leading to greater prey predictability. We concluded that prey predictability over various scales affected predator search patterns.     

Mate quality,not aggressive spillover,explains sexual cannibalism in a size-dimorphic spider

Sexual cannibalism particularly before mating is costly for the male victim but also for the female aggressor if she risks remaining unmated. The aggressive spillover hypothesis explains the persistence of this behavior as a maladaptive side effect of positive selection on aggressiveness in a foraging context. The hypothesis predicts that the occurrence of sexual cannibalism is explained by female aggressiveness but is not related to male phenotype or behavioral type. An alternative hypothesis invokes sexual selection and makes the opposite prediction namely that sexual cannibalism is an expression of female choice and should hence mainly target males of low quality. We tested the above hypotheses on a sexually dimorphic nephilid spider Nephilengys livida, known for male monopolization of females via genital damage, female genital plugging, and mate guarding, by staging mating trials during which we recorded mating behaviors and occurrences of pre- and postcopulatory cannibalism. We did not restrict assessment of aggressiveness to the mating and foraging context but also included aggression against same sex conspecifics. To assess female personalities, i.e., consistent individual differences in behavior including aggressiveness, we repeatedly tested them for intra-sex aggression, voracity towards prey, locomotory activity, and boldness. Females exhibited consistent differences in intra-sex aggressiveness, latency to attack prey, and boldness. Aggressive females had shorter latencies to attack prey and were more active than non-aggressive ones. In contrast to the predictions of the aggressive spillover hypothesis, females that were aggressive towards prey and towards other females were not more likely to attack a male than non-aggressive females. In support of the mate choice hypothesis, less aggressive males were more likely attacked and cannibalized than more aggressive ones. This hints at sexual selection for aggressiveness in males and raises the question of mechanisms that maintain variation in male aggressiveness.     

Male copulatory guarding enhances female foraging in a water strider

Summary In the laboratory, females of Gerris remigis foraging singly after being separated from a copulating male averaged 16 times longer to repel male copulatory attempts compared to repelling time when copulating (60 trials). In<10 min after separation, all females either became immobile at the edge (in 34% of all trials), or recopulated and either continued to forage (61%) or swam to the edge and became immobile (5%). Copulating females captured 85% of imitation prey, but single females, harassed by males, captured only 32% of prey.During a census of 2 small streams, in the pools with swimming single males, >99% of all swimming females were copulating, <1% being single. Ninety seven percent of all single females were immobile at the edge, whereas only 11% of all pairs were at the edge. In pools without swimming single males, 32% of all single females were swimming, compared to <1% in pools with swimming males. Introductions of a male into 4 pools with a female swimming singly in each resulted in the females either becoming immobile at the edge or copulating. The 2 streams had a : sex ratio of 1.6:1, and 79% of all females were copulating during the census. Foraging, copulation and copulatory attempts continued at a reduced level during the night.Thus in streams where male G. remigis are attempting to copulate, females can forage effectively only by carrying a copulating male who apparently repels copulatory attempts by other males.     

Adaptive peaks and alternative foraging tactics in brook charr: evidence of short-term divergent selection for sitting-and-waiting and actively searching

Some recently emerged brook charr (Salvelinus fontinalis) inhabiting still-water pools along the sides of streams are sedentary and eat crustaceans from the lower portion of the water column. Others are more active and eat insects from the upper portion of the water column. We provide evidence that this divergent foraging behavior reflects short-term divergent selection brought about by intraspecific competition in the presence of alternative food sources. Rates of encounters and interactions between individuals were density dependent, and encounter and interaction events were closely timed with prey capture attempts. In addition, aggressive fish made more foraging attempts per minute than nonaggressive fish. Aggressive fish were also either inactive or very active, while nonaggressive fish exhibited intermediate levels of activity. Growth rate potential, an important component of fitness during the early life stages of brook charr, was assessed using tissue concentrations of RNA and found to be highest for sedentary fish and for active fish making frequent foraging attempts, and lower for fish exhibiting intermediate levels of activity. Our findings support contentions that individual behavior plays an important role during initial steps in the evolution of resource polymorphisms. Received: 27 July 1998 / Accepted after revision: 16 November 1998     

Swim speeds of free-ranging great cormorants

Information about foraging speeds is particularly valuable when the impact of a predator species upon a community of prey has to be defined, as in the case of great cormorants. We measured the swim speed of 12 (six males and six females) free-ranging great cormorants Phalacrocorax carbo, foraging off the Greenland coast during the summer of 2003, using miniaturized data-loggers. Although mean body mass of males was 27% greater than that of females, and mean swim speed of males were 29–57% higher than that of females during foraging phases (but not descent phases) of dives, these differences in speeds were not significant due to high variances. Birds descended to the mean maximum depth of 4.7 m at an average speed of 1.6±0.5 m s⁻¹, a speed similar to that measured in captive cormorants in previous studies. Although bursts of up to 4 m s⁻¹ were recorded, speed usually decreased during the deepest (foraging) phase of dives, being on average 0.8±0.6 m s⁻¹. Speeds measured here should be taken with caution, because the large propeller loggers used to measure speed directly decreased descent speeds by up to 0.5 m s⁻¹ when compared to smaller depth-only loggers. Cormorants in Greenland seem to combine two searching strategies, one requiring low speed to scan the water column or benthos, and one requiring high speed to pursue prey. These two strategies depend on the two main habitats of their prey: pelagic or demersal.     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

Home range and social behavior of three species of European Pitymys (Mammalia,Rodentia)

Summary Habits, home range, and social behavior of three species of voles of the genus Pitymys, P. multiplex, P. subterraneus, and P. savii, were studied in Tessin (Switzerland) by radioactive tagging. P. multiplex and P. savii are fossorial and often use the burrow systems of moles (Talpa); P. subterraneus moves around on the surface under dense vegetation. Males and females of P. multiplex were territorial and their home ranges averaged 342 and 229 m² respectively. The mating system appeared to be monogamous. P. savii male and female home ranges averaged 445 and 298 m² respectively. Groups of up to one to three adult females, one or more adult males, and several young, occupied exclusive communal territories. The mating system appeared to be flexible, including monogamy, polygyny, and polyandry. P. subterraneus females occupied exclusive territories (mean 256 m²) overlapped by the home ranges of one or more males (1025 m²). The mating system appeared to be promiscuous. Individuals of the three species occupied one to seven nests. In P. savii all family members utilized three or more communal nests at the same time or alternatively. Co-nesting between males and females of P. multiplex or P. subterraneus were observed less frequently.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.