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     

Is boldness affected by group composition in young-of-the-year perch (Perca fluviatilis)?

In young-of-the-year perch (Perca fluviatilis), individuals within groups differed in the degree of boldness, estimated by habitat utilisation and feeding activity in visual contact with a potential predator. We looked at changes in individual behaviour in connection with change of group composition. During the first period, perch were randomly assigned to groups, and time spent in open habitat versus in vegetation and number of prey attacks were registered. The perch were then categorised into personality types (shy, bold, intermediate) according to their behaviour. During the second period, fish were observed when sorted into new groups, each containing only one personality type. Shy individuals showed the largest changes in behaviour, and increased both the time spent in the open and the number of prey attacks when placed into the new groups. Feeding activity in shy fish during the second period was affected by group composition during the first period. After regrouping, bold individuals decreased their time in the open, whereas intermediate individuals did not change behaviour. Time in the open habitat was, to some extent, influenced by the behaviour of the other members of the group, but number of prey attacks was not. The behaviour of fish of the different personality types we have defined in this study seemed to be based on innate traits, but also modified by the influence of other group members and by habituation to the environment.Communicated by J.Krause     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Predator inspection behaviour in minnow shoals: differences between populations and individuals

Summary When minnows (Phoxinus phoxinus) detect a stalking pike (Esox lucius) one of their first responses is to perform inspection behaviour during which individuals or small groups approach the predator. This paper compares the inspection behaviour of two contrasting groups of minnows: Dorset minnows which have been heavily predated by pike for many thousands of years and Gwynedd minnows which have spent an equivalent period of time in a pike free environment. Minnows sympatric with pike inspected a realistic model pike more frequently and in larger shoals. Although they commenced inspection earlier they were more timid and kept a greater distance between themselves and the predator. After an inspection they were less likely to recommence foraging than minnows from the Gwynedd population. Individual differences in inspection found within the two populations suggest that selfish behaviour was present.     

Feeding rates and diel vertical migration of copepods near South Georgia: comparison of shelf and oceanic sites

Seventeen Longhurst Hardy Plankton Recorder profiles were taken over a diel cycle in January 1990 to study the feeding of four major copepods over the South Georgia shelf. Ontogenetic changes in vertical migration were followed and feeding cycles determined by gut fluorometry for Calanoides acutus Stage CV, Calanus sinillimus CV and CVI, C. propinquus CV and Rhincalanus gigas CV and CVI. In common with a neighbouring oceanic site visited two weeks later and reported elsewhere, all four species had a diel cycle of feeding and migration. The vertical distributions of C. simillimus (all stages), R. gigas (nauplii) and Euphausia frigida (postlarvae) were similar at both sites, the night being spent within the chlorophyll maximum at 15 to 30 m. However, the biomass dominants, C. acutus and R. gigas, dwelt below the chlorophyll maximum, about 30 m deeper than their oceanic counterparts. Unlike the oceanic site, feeding at the shelf site was not restricted to darkness, but increased 6 to 10 h before nightfall and finished at dawn; the intervening period coincided with sinking and digestion. Daylight feeding may have been induced by the shorter night, lower light levels or greater food requirements at the shelf site, despite planktonic predators being over three times more abundant. Daily ration estimates for R. gigas at both sites were only 2% body carbon per day. These low values contrast with its smaller competirors, whose rations were in the range 5.6 to 27%.     

Feeding ecology of a size-structured predator population,the South American sun-star Heliaster helianthus

Patterns of feeding in a population of Heliaster helianthus (Lamarck), a common and dominant species of starfish indigenous to the Pacific South American coast, were investigated in an intertidal habitat in central Peru from October 1986 to April 1987. The H. helianthus population comprised individuals of 3.5 to 30.2 cm body size (diameter) with two modal size classes. The number of rays ranged between 18 and 40, and individuals with 31 to 33 rays accounted for ca. 42% of the total population. There was a higher rate of increase in ray number with body size amongst small individuals(<13.0 cm diam). H. helianthus is capable of feeding on more than one prey item at a time (average of 5.6 to 13.2 prey items handled, with several predators observed to hold >100), and both the number of prey individuals captured and the total prey biomass were significantly correlated with predator size. Amongst a total of 1132 feeding observations, the largest number of predators (an average of 85.4% of those feeding) were preying on the mussel Semimytilus algosus whilst another mussel, Perumytilus purpuratus, ranked second with 21,9% of predators feeding. The proportion of S. algosus in the diet increased from 65.4% in the smallest predator size-group (10.9 cm diam) to 91.2% in the largest (19.0 cm). In contrast, P. purpuratus and barnacles were more highly represented in the diet of small H. helianthus. The smallest size-group (10.9 cm) had low dietary overlap with larger sizes and less specialized prey utilization. Two geographically separated populations of H. helianthus in Peru and Chile showed contrasting patterns of prey utilization. S. algosus and P. purpuratus comprised 85.5 and 6.5% by number in the diet of the Peruvian population, respectively, whilst corresponding figures for the Chilean population were 8.3 and 60.5%, with barnacles attaining a higher share (22.6%). However, the total number of prey individuals per feeding predator was almost the same in Peru and Chile, with 10.0 and 10.7 individuals, respectively. H. helianthus individuals of different sizes occupy slightly different microhabitats within the intertidal area, which, coupled with differential spatial distribution of prey species, results in the predator population being able to utilize a wide range of resources.     

Large male advantage for access to females: evidence of male-male competition and female discrimination in a territorial salamander

Summary In the natural habitat visited in this study, adult male red-backed salamanders (Plethodon cinereus) found near females were significantly larger than adult males found alone; there was no evidence for size assortative associations. Previous research has indicated that females associate preferentially with males that occupy high quality territories and that larger individuals are more successful at obtaining higher quality areas. In laboratory experiments, when resource quality was held constant and males were restrained so that male-male interactions were prohibited, females discriminated behaviorally between large and small males by spending more time visually and chemically assessing larger males and more time apparently attempting to leave the chamber when near smaller males. Females were found near the larger males at the end of the trials significantly more often than predicted by chance. In a separate experiment in which only females were restrained, larger males spent significantly more time in aggressive postures when paired with smaller males than when alone with the female. Smaller males spent significantly more time in submissive postures when paired with larger males and more time near the female when alone. Therefore, large body size of males may positively affect both intra- and intersexual interactions and, ultimately, mating success of male P. cinereus.     

Dominance status influences female reproductive strategy in a territorial African rodent <Emphasis Type="Italic">Rhabdomys pumilio</Emphasis>

Dominance status influences the fitness of many mammals. Using African striped mice Rhabdomys pumilio, we tested whether (1) dominant females have greater reproductive success than subordinate females, (2) dominant females influence the reproductive output of subordinate females when they are housed in close proximity, (3) reproductive output of a female changes in response to the dominance status of her neighbours, and (4) whether prolonged association between individuals influences the variance in reproductive success between dominants and subordinates (i.e. the ‘dear enemy’ phenomenon). The size and mass of litters of dominants increased significantly when housed adjacent to subordinates than when housed apart. The litter size and mass of subordinates remained unchanged, although subordinates spent significantly more time with their pups when housed close to dominants than when housed apart; time spent with pups by dominants remained unchanged. Moreover, females modified their reproductive output and behaviour in relation to the dominance status of their neighbours. Following prolonged association, dominants still had greater reproductive success, but now, the time spent with pups decreased in subordinates. We suggest that dominants adopt a strategy to increase the reproductive value of their litter, whereas subordinates adopt a pup defence strategy. These strategies are flexible and are influenced by the dominance status and period of association between neighbours, so that females could maximize their fitness in response to varying social conditions.     

Effects of feeding experience in the copepod Eucalanus pileatus: a cinematographic study

High-speed microcinematography was used to examine the effects of prior experience with particular cell types on the feeding efficiency of a calanoid copepod. Female Eucalanus pileatus were fed monocultures of either the 5-m diatom Thalassiosira pseudonana or the 11-m diatom T. weissflogii during a 2-to 3-d preconditioning period. The smaller diatoms are accumulated passively by the second maxillae while the larger diatoms are detected and actively captured as individual cells. Four females from each preconditioning culture were transferred to a monoculture of the large cells and their behavior filmed at five intervals over a 24-h period to determine whether a loss of efficiency occurs when the copepods must shift capture modes. Ingestion rates for females experienced with the larger cells were approximately 2.5 times higher than those of inexperienced females. Six sequential behavioral steps in the feeding process could alter ingestion rates: (1) amount of time spent flapping the feeding appendages. (2) rate of flapping of the feeding appendages, (3) ability to detect individual cells, (4) success rate of capture attempts, (5) capture and handling time per cell and (6) rejection rate of captured cells. An increased ability to detect cells and a decreased rejection rate contributed significantly to the higher ingestion rate of experienced feeders, indicating that copepods have the ability to learn during the feeding process. Grazing rates may be seriously underestimated in experiments which do not include a preconditioning period, especially those which calculate ingestion over short time intervals. Such effects may also influence the feeding of copepods in the field when encountering changes in particle spectra through vertical migration or horizontal displacement.     

Nutritional constraints on mountain baboons (Papio ursinus): Implications for baboon socioecology

Summary Populations of baboon (Papio sp.) at geographic and climatic extremes for the genus show a tendency to one-male organization, whereas most baboons live in multimale social groups; this effect has been attributed largely to limitation of food supply, but baboons' complex diet has hindered proper nutritional analyses. To test these optimal-diet explanations of social variation, we quantified intake and used phytochemical analysis of foods to compare the nutrition, during seasonal changes, of two groups of mountain baboons (P. ursinus) living at different altitudes of a continuous grassland habitat. The majority of plant foods were eaten uniquely by one or other group, though their altitudinal separation was only 400 m, and the time budget of feeding choices varied with age-sex class as well as season. Converting to a common currency of nutrients reveals that baboons gained the same yield from a unit time spent foraging (whether this is measured in edible dry weight, or simply protein) in both groups, despite their differing mean altitude, whereas seasonal variation was large and statistically significant. Increased feeding time at the winter bottleneck made no effective compensation for the poorer food yields: in late winter there was a minimum for daily nutrient gain at both altitudes. Apparently this population is already at an extreme for the time animals devote to foraging in winter, when they rely on inconspicuous and slow-to-harvest swollen shoot bases and underground plant storage organs. Since an individual's nutrient yield does not vary with altitude, we conclude that socioecological parameters are effectively optimized for feeding. Since contest competition is absent, this adjustment of foraging efficiency is largely through the effect of differential density on scramble competition. Differences in social structure are considered to be a secondray consequence of optimal foraging, mediated through altitudinal variation in either population density or in day range limits.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

Flow rate and particle concentration within the house of the pelagic tunicate Oikopleura vanhoeffeni

The food-concentrating filter within the house of Oikopleura vanhoeffeni Lohmann consists of three layers, upper and lower fine-mesh layers and an intermediate coarse-mesh layer. The filter is a natural analogue of man-made tangential, or cross-flow, filters that are used to concentrate fine particulate and colloidal material by the exclusion of water. Flow analysis using video microscopy of O. vanhoeffeni, collected in 1989 and 1990 at Logy Bay, insular Newfoundland, indicates concentration of particles of 74 to 1089 times that of ambient seawater ( ,n=23), for individuals 1.8 to 5.4 mm long. The smallest individuals had the highest concentration factors. Flow through the house is predominately affected by viscous forces and values of Reynolds numbers are 1. Flow rates through the house of O. vanhoeffeni indicate a maximum filtration capacity of ca. 1.4 to 26 liters d^-1 and clearance rates (assuming ca. 50% of the time is spent not actively pumping) of 0.7 to 13 liters d^-1, depending on body size. The estimated thickness of the boundary layer around individual fibers of the food-concentrating filter is 0.08 m, which is less than the pore width. The estimated concentration factors suggest that colloidal and fine particulate material may aggregate into larger particles within the food-concentrating filter and food tube during feeding.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.     

Density and distribution patterns of the temperate marine fish Cheilodactylus spectabilis (Cheilodactylidae) in a reef environment

Cheilodactylus spectabilis (Hutton) is common over shallow reefs in north-eastern New Zealand. Replicated transect between-area differences in density and in size frequency. C. spectabilis is not nocturnally active. The between-area differences in density and size frequency remained constant over daylight hours, demonstrating that C. spectabilis does not undergo any systematic feeding migrations. Topographic complexity was shown to have a significant positive influence on fish density. Mean size was related to water depth at 5 localities investigated, with small (<200 mm standard length, SL) individuals being restricted to shallow water. All sizes of C. spectabilis examined had fed on small invertebrates, with gammarid amphipods predominating. No evidence of size-related differentiation in feeding patterns was observed, although small individuals spent significantly more time feeding than large ones. Small (<200 mm SL) individuals occupied feeding areas and shelter sites from which they excluded other small C. spectabilis. Large individuals showed no site-associated aggression, covered a greater area during daily movements and overlapped with other large and small C. spectabilis while feeding. No evidence of dawn or dusk peaks of feeding activity or movement was collected.     

Individual and species recognition in centrarchid fishes: evidence and hypotheses

Summary We conducted laboratory experiments to determine if juveniles of three species of centrarchid fishes displayed abilities for species or individual recognition. In one experiment we reared rock bass (Ambloplites rupestris) in social isolation and in social groups. Both the isolated and group-reared fish spent significantly more time close to conspecifics than to heterospecifics. These results suggest that species recognition in rock bass is controlled by a closed genetic program and that social experience is not necessary. A second set of experiments was conducted to determine if year-old blue-gill (Lepomis macrochirus), pumpkinseed (L. gibbosus), and rock bass juveniles could discriminate between familiar and unfamiliar conspecifics. Bluegill spent significantly more time with familiar conspecifics than with unfamiliar conspecifics, while the other two species displayed no such trend. Bluegill spent significantly more time with familiar conspecifics than did either of the other two species. We postulate that these interspecific differences in recognition abilities may be related to differences in habitat and group parameters among the species.     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Comparisons of the feeding activity and diets of male and female cunners Tautogolabrus adspersus (Pisces: Labridae)

The diel distribution of feeding activity during the spawning season was compared for territorial male and female cunners Tautogolabrus adspersus in Conception Bay, Newfoundland, Canada between June and August, 1978. Territorial males feed significantly less often than females and concentrate their feeding activities in the morning, whereas females feed as frequently in the afternoon as in the morning. Based on the contents of alimentary tracts, territorial males, non-territorial males, and females have different diets. Since these three groups share the same habitat, the observed dietary differences probably reflect differences in their foraging behaviour. The relevance of these findings to the concept of time minimizer and energy maximizer is discussed.     

Home range and habitat utilization of adult California sheephead,<Emphasis Type="Italic"> Semicossyphus pulcher</Emphasis> (Labridae), in a temperate no-take marine reserve

The California sheephead, Semicossyphus pulcher Ayres (Labridae), is a carnivorous, temperate, rocky-reef/kelp-bed species that is highly sought in recreational and commercial fisheries. Fine-scale acoustic telemetry tracking was used to ascertain the home range and habitat utilization of S. pulcher. Sixteen adult S. pulcher (26–38 cm SL) were surgically fitted with small acoustic transmitters and manually tracked for up to 144 h during multiple, 24-h periods between March 2001 and August 2002 within the Catalina Marine Science Center Marine Life Reserve (33°26N; 118°29W). A geographic information system was used to calculate home range sizes (95% kernel utilization distributions) and habitat use. Tracking of the first five fish over 24 h confirmed that S. pulcher were strictly diurnal, so the remaining 11 fish were tracked from 1 h before sunrise to 1 h after sunset. Home ranges varied greatly, from 938 to 82,070 m², with a mean (±SD) of 15,134±26,007 m². Variability in home range sizes among fish was attributed to differences in habitat shape (embayment vs. contiguous coastline) and to natural habitat boundaries (deep, sandy expanses) in adjacent areas within the reserve. There was a significant relationship between fish length and proportion of time spent in different habitats (sand vs. reef). S. pulcher were found within rocky-reef areas 54% of the time, and, within these areas, a greater percentage of daytime was spent in high-relief areas. Based on the relatively small size and persistence of home ranges of adult S. pulcher, no-take reserves, if they contain appropriate habitat, would provide adequate protection for their stocks.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1573-1.Communicated by J.P. Grassle, New Brunswick     

Behavior of juvenile American lobsters, Homarus americanus, under predation risk

The influence of predation risk and food deprivation on the behavior and activity of juvenile American lobsters, Homarus americanus Milne Edwards, was examined in single and paired individuals in laboratory experiments performed during 1988 and in the winter of 1991/92. In the presence of a predator (the tautog Tautoga onitis Linnaeus) restrained behind a barrier, single lobsters significantly reduced the time spent feeding at night, consumed fewer mussels, and quickly brought them back to shelter. Single lobsters did not forage during the day in any treatment. If deprived of food for 60 h, they consumed more mussels and spent more time walking than recently fed (12-h food-deprived) lobsters. Paired lobsters did forage during the day in the presence of a predator. The smaller lobsters (subdominant) in the pairs foraged for a longer time in the presence than in the absence of a predator and significantly longer than single individuals. Shelter occupancy was significantly shorter in single, recently fed lobsters in the presence of a predator compared to time spent sheltering in its absence. Among food-deprived lobsters, paired individuals spent a significantly shorter time within the shelter than single lobsters in the absence of a predator. Larger (dominant) lobsters, however, spent more time than subdominant lobsters within the shelter during all periods of the day. Without a predator, paired lobsters spent significantly more time than single ones in shelter-related activities. Under predation risk, subdominant lobsters concentrated shelter-building time during the day and built a higher percent of alternative shelters than either single or dominant lobsters. In the absence of a predator, paired lobsters walked in the open area for a significantly longer time than single ones in the absence of a predator. This apparently was associated with fighting between dominant and subdominant lobsters and the attempts of the larger lobster to drive the smaller one from its shelter. During the day, lobsters fought for a significantly longer time in the presence than in the absence of a predator. When the tautog was not constrained, mortality rate was similar in both single and paired lobsters. Mortality rate among subdominant lobsters, however, was seven times higher than among dominant lobsters. We suggest that the risk of predation interferes with the ability of single juvenile lobsters to acquire and consume food. They appear to trade off energetic consideration against risk of predation when foraging away from the shelter. The introduction of a conspecific competitor to the system may further increase risk (of the subdominant) to the predator. Intraspecific interactions tend to increase the risk of predation to smaller lobsters but increase the survival rate among larger lobsters. Received: 6 February 1995 / Accepted: 2 September 1997