Individual differences in foraging behavior and cortisol levels in recently emerged brook charr (Salvelinus fontinalis)

Recently emerged brook charr (Salvelinus fontinalis) foraging in still-water pools along the sides of streams tend to be either sedentary, feeding from the lower portion of the water column (a sit-and-wait tactic), or very active, feeding from the upper portion of the water column (an active search tactic). We tested whether the individual differences in foraging behavior were associated with baseline concentrations and responses of cortisol, a steroid hormone linked to personality differences in a variety of animals including fishes. We quantified the proportion of time spent on moving by focal charr in the field and then capturing them. Captured individuals were either (i) sacrificed immediately to quantify baseline cortisol concentrations, (ii) held in an unfamiliar field environment for 15 min and then sacrificed to quantify cortisol concentrations in response to handling and holding in a novel field environment, or (iii) held in an unfamiliar field environment with a white Plexiglas base (stressor) for 15 min to quantify cortisol concentrations in response to a novel object. Eleven statistical models relating cortisol concentrations to the proportion of time individuals spent on moving while searching for prey were compared using multi-model inferencing. Cortisol concentrations were higher for charr that spent a lower proportion of time on moving in the field than for charr that spent a higher proportion of time on moving. For a given proportion of time spent on moving, mean cortisol concentrations between baseline and experimental treatments, our measure of cortisol response, did not differ markedly. Our findings suggest that the foraging tactics displayed by wild brook charr in the field could reflect differences in how individuals perceive their environment.     

Foraging behaviour and brain morphology in recently emerged brook charr, Salvelinus fontinalis

Recently emerged brook charr (Salvelinus fontinalis) foraging in still-water pools along the sides of streams are either active, feeding on insects from the upper portion of the water column away from the stream bank, or sedentary, feeding on crustaceans emerging from the hyporheic zone near the stream bank. We tested whether the frequency of movement displayed by individual brook charr searching for prey in the field was related to the relative volume of the telencephalon, a brain region involved with movement and space use in fishes. Movement of individuals searching for prey was quantified in the field, individuals were captured and volumes of the telencephalon and of the olfactory bulbs, a brain region neighbouring the telencephalon but not implicated in space use, were measured. Individuals with larger telencephalon volumes moved more frequently on average while searching for prey in the field than did individuals with smaller telencephalon volumes. The frequency of movement was unrelated to differences in the volume of the olfactory bulbs, suggesting that the relationship between telencephalon volume and movement was not a consequence of differences in overall brain size. Demonstrating a correlation between foraging behaviour and brain morphology for brook charr exhibiting different foraging tactics suggests that diversification in brain structure and function could be important aspects of the foraging specialization believed to occur during early stages in the evolution and development of resource polymorphisms.     

Aggressiveness and foraging mode of young-of-the-year brook charr,Salvelinus fontinalis (Pisces,Salmonidae)

Summary Young-of-the-year brook charr in streams use either an active or a sit-and-wait foraging tactic and exhibit a range of resource defense from territoriality to tolerating conspecifics. We use simple graphical models, based on encounter rate with drift and the theory of economic defendability, to predict qualitative changes in the aggressiveness and mobility of brook charr in relation to current velocity. Aggressiveness (percent of conspecifics eliciting an overt response) initially increases with increasing current velocity, as does drift rate and foraging rate. However, aggressiveness decreases at high current velocities, probably because of increased costs of defense at these velocities. In standing water areas, brook charr use primarily an active foraging tactic, but mobility (percent time spent moving) decreases rapidly as current velocity increases. These results are generally consistent with the simple graphical models. A literature survey suggests that the models can be generalized for most species of stream salmonids.     

Innate antipredator responses of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) depend on predator species and their diet

The ability to discriminate between more dangerous and less dangerous predators can have serious fitness advantages for fish juveniles. This is especially true for hatchery-reared fish young used for stocking, because their post-release mortality is often much higher than that of wild-born conspecifics. We tested whether two coexisting fish predators and their different diets induce innate behavioral responses in predator-naive Arctic charr (Salvelinus alpinus) young originating from an endangered hatchery-bred population used for re-introductions. We predicted the antipredator responses of charr to be stronger towards chemical cues of brown trout (Salmo trutta) and pikeperch (Stizostedion lucioperca) than towards odorless control water. More pronounced antipredator behavior was predicted in treatments with predators fed on charr than when their diet consisted of another sympatric salmonid, European grayling (Thymallus thymallus), or when they were food-deprived. The Arctic charr young showed strong antipredator responses in all brown trout treatments, whereas odors of the less likely predator pikeperch were avoided with conspecific diet only. Freezing was the most sensitive antipredator behavior, as it was completely absent in control treatments. We found considerable individual variation in the amount and strength of antipredator responses. Although almost half of the charr failed to show antipredator behavior towards the piscivores, those with the innate ability showed highly sensitive recognition of predator odors. Our results indicate that the innate antipredator behavior of the juvenile fish is already finely tuned to respond specifically to chemical cues from different fish predators and even their diets.Communicated by J. Krause     

Shoal composition, body size and foraging in sticklebacks

Individuals which deviate from the majority in groups are likely to be most vulnerable to predation. This oddity effect, by definition, is frequency dependent, eventually fading at equal frequencies of the phenotypes in a group. It has been hypothesized that the increased predation risk of odd individuals may play an important role in the formation of phenotypically uniform shoals of fish. However, recent work has indicated that individuals may experience, or value, their predation hazard differently depending on their own size in relation to that of other group members: single large fish, but not small ones, appear concerned about their oddity in a shoal. Here I show that the apparent wariness of large fish is also expressed in a frequency-dependent manner, closely conforming to what is predicted if the oddity effect is responsible for their behavior. Using foraging activity of individuals as a means to evaluate their predation risk, I demonstrate with shoals comprising 12 threespine sticklebacks (Gasterosteus aculeatus) that large fish forage least actively when in a shoal consisting of 2 large and 10 small fish. An increase in the number of large fish to 4 among 8 small individuals clearly results in an increase in their foraging activity. However, having reached an equal frequency with small fish in a shoal, large fish do not seem to change their foraging activity much even when their number in a shoal increases further. In contrast, foraging activity of small sticklebacks remains fairly constant throughout the entire range of tested shoal compositions, providing further evidence that small and large fish respond to their oddity differently. Received: 12 February 1998 / Accepted after revision: 7 May 1998     

Determinants of multiple central-place territory use in wild young-of-the-year Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Patterns of space use provide key insights into how animals exploit local resources and are linked to both the fitness and distribution of individuals. We studied territory size, mobility, and foraging behavior of young-of-the-year Atlantic salmon Salmo salar in relation to several key environmental factors in Catamaran Brook, New Brunswick, Canada. The 50 study fish were all multiple central-place foragers (i.e., alternated among several sit-and-wait foraging stations) and showed great variability in territory size and the total distance traveled within the territories. Territory size increased with the mean distance traveled between consecutive foraging stations, the number of stations visited, and the mean foraging radius. Fish also varied greatly in how much of the total travel distance was associated with foraging at a station (14.8–91.8%) versus switching among stations (4.6–84.3%). As predicted, fish in slow-flowing waters, where drifting prey were scarce, used larger multiple central-place territories than individuals in faster, more productive waters. Interestingly, however, the most mobile fish did not inhabit slow-running waters as predicted but were found at intermediate (optimal) water current velocities. Hence, our study suggests that among some multiple central-place foragers, increased mobility may not only serve to increase prey encounter rate but may reflect an attempt to patrol territories in favorable habitats. Further studies are needed to determine the generality and the ultimate benefits of multiple central-place space use among stream-dwelling fish and other animals.     

The influence of prey distribution on the foraging strategy of the lizard Oligosoma grande (Reptilia: Scincidae)

The grand skink, Oligosoma grande, is a diurnal rock-dwelling lizard from the tussock grasslands of Central Otago, New Zealand, whose diet includes a variety of arthropods and fruit. We conducted a field experiment to examine the influence of prey distribution on foraging behavior and spacing patterns. On sites where prey distribution was unaltered (control sites), males and females differed in diet and foraging behavior. Most male feeding attempts were directed at large strong-flying insects, and males used a saltatory search pattern that involved relatively infrequent moves of long duration. Females spent more effort catching small weak-flying insects and visiting fruiting plants. Their search behavior involved frequent moves of short duration. The placement of meat-bait on experimental sites led to a redistribution of large flies without influencing other prey types. Experimental females switched foraging strategy by adopting a search pattern of relatively infrequent moves of long duration, increasing the frequency of attempts to capture large prey, and reducing the importance of fruit in their diet. The experimental manipulation appeared to influence space use. On control sites, both sexes had comparably sized home ranges. On experimental sites, male home ranges were significantly larger than female home ranges. Received: 3 November 1997 / Accepted after revision: 13 December 1998     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

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.     

Coordinated mechanics of feeding,swimming, and eye movements in <Emphasis Type="Italic">Tautoga onitis</Emphasis>, and implications for the evolution of trophic strategies in fishes

Fish feeding behavior results from successful coordination of the fins, jaws, and sensory systems, and the organization of this behavior may affect the fish’s foraging abilities and trophic ecology. Using quantitative kinematic methods, movements of the jaws, fins and eyes of Tautoga onitis (Teleostei: Labridae) were analyzed during feeding events. Tautog feeding events consisted of three phases: approach, strike, and recovery, each defined by a combination of kinematic events. The approach was characterized by changes in fin movements and in body position, with the eyes directed forward at the food item. The strike began with the onset of jaw opening and protrusion, then cranial elevation, with the eyes no longer looking at the food item. The end of the strike and the beginning of the recovery involved a braking maneuver with the pectoral fins; the fish turns and swims away from the original food location item after prey capture. The coordination performance variables of tautog were quantitatively compared to published data from closely related cheiline wrasses and parrotfishes, which represent different feeding ecologies within a monophyletic assemblage. Fishes feeding on molluscs and benthic invertebrates (Cheilinus fasciatus and Tautoga onitis) represented an intermediate coordination condition, with herbivores (the parrotfishes, Scarus quoyi and Sparisoma radians) at one extreme, and fishes feeding on elusive prey (Epibulus insidiator and Oxycheilinus digrammus) at the other extreme. The analysis suggests that the biomechanical demands of coordination for feeding on benthic invertebrates may represent a generalized, and perhaps ancestral behavior in the wrasses, whereas more specialized trophic niches have evolved divergent, more specialized demands. Examining the movement and coordination of the jaws, fins, and eyes during fish feeding provides a detailed mechanistic basis for behavior, and comparison of coordination patterns during feeding among different taxa can measure how these trophic strategies differ. Understanding the evolution of feeding ecologies in these demersal fishes may have implications for understanding their role within their shallow water reef community.     

Swimming behavior of juvenile anchovies (<Emphasis Type="Italic">Anchoa</Emphasis> spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O₂ l⁻¹, except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O₂ l⁻¹. We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s⁻¹ that were near the theoretical optimum speeds between 7.0 and 8.0 bl s⁻¹. An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s⁻¹, expected = 6.4 bl s⁻¹). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.     

Regulation of pollen foraging in honeybee colonies: effects of young brood, stored pollen, and empty space

Pollen storage in a colony of Apis mellifera is actively regulated by increasing and decreasing pollen foraging according to the “colony's needs.” It has been shown that nectar foragers indirectly gather information about the nectar supply of the colony from nestmates without estimating the amount of honey actually stored in the combs. Very little is known about how the actual colony need is perceived with respect to pollen foraging. Two factors influence the need for pollen: the quantity of pollen stored in cells and the amount of brood. To elucidate the mechanisms of perception, we changed the environment within normal-sized colonies by adding pollen or young brood and measured the pollen-foraging activity, while foragers had either direct access to them or not. Our results show that the amount of stored pollen, young brood, and empty space directly provide important stimuli that affect foraging behavior. Different mechanisms for forager perception of the change in the environment are discussed. Received: 13 June 1998 / Accepted after revision: 25 October 1998     

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     

Depth and muscle temperature of Pacific bluefin tuna examined with acoustic and pop-up satellite archival tags

Six Pacific bluefin tuna were tracked with ultrasonic telemetry and two with pop-up satellite archival tags (PSATs) in the eastern Pacific Ocean in 1997, 1998, and 1999. Both pressure and temperature ultrasonic transmitters were used to examine the behavior of the 2- to 4-year-old bluefin tuna. The bluefin spent over 80% of their time in the top 40 m of the water column and made occasional dives into deeper, cooler water. The mean slow-oxidative muscle temperatures of three fish instrumented with pressure and temperature transmitters were 22.0–26.1 °C in water temperatures that averaged 15.7–17.5 °C. The thermal excesses in slow-oxidative muscle averaged 6.2–8.6 °C. Variation in the temperature of the slow-oxidative muscle in the bluefin was not correlated with water temperature or swimming speeds. For comparison with the acoustic tracking data we examined the depth and ambient temperature of two Pacific bluefin tagged with pop-up satellite archival tags for 24 and 52 days. The PSAT data sets show depth and temperature distributions of the bluefin tuna similar to the acoustic data set. Swimming speeds calculated from horizontal distances with the acoustic data indicate the fish mean speeds were 1.1–1.4 fork lengths/s (FL s⁻¹). These Pacific bluefin spent the majority of their time in the top parts of the water column in the eastern Pacific Ocean in a pattern similar to that observed for yellowfin tuna. Received: 4 April 2000 / Accepted: 25 October 2000     

Prey ecology and behaviour affect foraging strategies in the Great Cormorant

The fine link between a particular dive pattern and a specific prey item represents a challenging task in the analysis of marine predator–prey relationships. There is growing evidence that prey type affects diving seabirds’ foraging strategies, dive shapes and underwater activity costs. This study investigates whether a generalist diver, the Great Cormorant Phalacrocorax carbo, modifies the time budget allocated to prey-capture behaviour and breathing strategies (reactive vs. anticipatory) with respect to the prey type (pelagic vs. benthic). Video recordings of 91 Great Cormorants show how the ecology and behaviour of their main prey, Mullets (Mugilidae) and Flounders Platichthys flesus, affect dive/surface durations and the diving pattern. The demersal habit and the low mobility of Flounders leads to an easy access to prey with an anticipatory strategy. Moreover, the patchy distribution of this fish species increases prey-capture rates. Conversely, Mullets exploit the whole water column and are highly mobile, and this is reflected in the need of performing two sequential dives to capture a prey, both longer and likely more expensive, with a consequent switch of strategy from reactive in the searching phase to anticipatory breathing during prey-capture events. This study provides evidence that a generalist diver may switch between different foraging strategies, and it shows how each of them may be optimal under particular ecological conditions. These constraints influence the dynamics that operate within the marine food chains and have relevant implications in managing lagoon areas, including fish ponds.     

Movement patterns of large bigeye tuna (Thunnus obesus) in the open ocean, determined using ultrasonic telemetry

 The horizontal and vertical movements of large bigeye tuna (Thunnus obesus Lowe, 1839; 25 to 50 kg) captured in the south Pacific Ocean (French Polynesia) were determined using pressure-sensitive ultrasonic transmitters. Bigeye tuna swam within the first 100 m below the surface during the night-time and at depths between 400 and 500 m during the daytime. The fish exhibited clear relationships with the sound scattering layer (SSL). They followed its vertical movements at dawn and dusk, and were probably foraging on the organisms of the SSL. Bigeye tuna did, however, make regular rapid upward vertical excursions into the warm surface layer, most probably in order to regulate body temperature and, perhaps, to compensate for an accumulated oxygen debt (i.e. to metabolize lactate). The characteristics of these dives differ from those reported from previous studies on smaller bigeye tuna (∼12 kg) near the main Hawaiian Islands. During the daytime, the large fish in French Polynesia made upward excursions approximately only every 2.5 h, whereas smaller fish in Hawaiian waters made upward excursions approximately every hour. Our data are the first observations on the role of body size in the vertical behavior of bigeye tuna. Received: 9 September 1998 / Accepted: 25 November 1999     

Protein–Surfactant interactions in the defensive skin secretion of the Red Sea trunkfish Ostracion cubicus

With the aid of column chromatography, solvent fractionation and toxicity assays we made the following observations. (1) The defensive skin secretion of the Red Sea trunkfish Ostracion cubicus consists, on a dry weight basis, of about 15% proteins, of 5 to 70 kDa molecular weight. (2) The proteins serve as an integral part of the chemical defense mechanism and fulfill two essential roles: (a) as ichthyotoxins which affect fish through external application into the surrounding water; and (b) as modifiers/regulators of the action of the well-known active ingredient of trunkfish secretion, the cationic, quaternary-ammonium, surfactant pahutoxin (PHN). The latter, when isolated from other components, plays only a limited role in ichthyotoxicity. (3) The regulatory role is performed by nontoxic, and as yet unidentified, proteins which potentiate PHN ichthyotoxicity. Potentiation is presumably preceded and mediated by the protein–PHN association, suggesting a guiding “pharmacokinetic” role of the regulatory proteins. (4) Finally, it may be concluded that the entire secretion process functions with unique pharmacological complexity due to the multiplicity of active substances and their interdependency. Received: 7 May 1998 / Accepted: 7 June 1999     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.     

Time budgets and foraging in a Malagasy primate: do sex differences reflect reproductive condition and female dominance?

Female mammals commonly employ behavioral tactics of modulating activity levels and foraging behavior to counter the energetic burden of reproduction; these behavioral changes are reflected as intersexual differences. Traditional views of Malagasy primates posit that high reproductive costs select for female dominance which guarantees to energetically stressed females priority of resource access. I tested predictions regarding reproductive influences on sex differences in time budgets and foraging behavior using two groups of Milne-Edwards' sifaka (Propithecus diadema edwardsi) in southeastern Madagascar. Compared to males, females increased neither feeding nor resting time during gestation or lactation. Sex differences were essentially absent in all foraging time variables examined (time, duration, rate). In contrast, dietary composition diverged between the sexes in some months. The possibility that females selected particular food items to boost nutrient and energetic intake to meet increased requirements during reproduction must be further clarified with nutritional analyses. Sex differences in plant part choices coincided with lactation in one of the two study groups. Thus, the timing of sex differences in feeding patterns of P. d. edwardsi only partially supports the prediction that sex differences are most pronounced during the period of greatest female energetic demand. A comparative review indicated no tight association between female dominance and sex differences in foraging among Malagasy primates. Traditional female dominance theory falls short of explaining the observed patterns. The results of my study coupled with recent evidence suggest that non-behavioral tactics involving energy conservation and storage require further attention as mechanisms by which female lemurs cope with reproductive costs. Received: 12 June 1998 / Accepted after revision: 10 October 1998     

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