Dwarf mongoose and hornbill mutualism in the Taru desert,Kenya

Summary Dwarf mongooses in the Taru desert region of Kenya form foraging communities with a variety of endemic bird species, especially hornbills. The prey spectra of the mongooses and hornbills overlap almost completely. For the other bird species forming the foraging community only partial overlap exists. The association between the birds and mongooses is actively sought by both parties. The birds wait in tress around the termite mound where the monogooses are sleeping for them to emerge and the mongooses delay their foraging departure if no birds are present. There is a positive relationship between the number of mongooses in the group and the number of birds accompanying them. A true mutualism only exists between the mongooses and the two hornbill species Tockus deckeni and T. flavirostris since their presence or arrival affects the subsequent start of foraging. These two hornbill species have also been observed to influence the start of foraging actively by means of two behaviour patterns termed chivvying and waking. Both the mongooses and birds are exposed to a high predator pressure from raptors with an overlap in the birds of prey predating the various species. This predator pressure is counteracted behaviourally by the mongooses by means of an altruistic behaviour pattern, guarding. Both mongooses and birds warn vocally and flee when a raptor is sighted. The mongooses modify their guarding behaviour to compensate for the warning behaviour of the birds in two ways: (a) fewer mongooses guard when large numbers of birds are present and vice versa, (b) the frequency of the mongooses' intraspecific warning calls is significantly reduced in cases where birds are present in comparison with those where they are absent. The birds also sight and respond to the raptor first on significantly more occasions than the mongooses. In addition, the birds also warn for raptor species which do not predate them but which are mongoose predators, not, however, for raptors which are not mongoose predators. This mutualistic association with its high degree of compensatory behaviour by both parties appears to be unique for free-living vertebrates and has its closest parallel in the trophobiosis described for ants and aphids.     

Flocking behaviour of passerines: a dynamic model for the non-reproductive season

Summary The behaviour of a small male passerine bird over a typical winter day is studied by a dynamic programming model. The bird can be either unpaired or paired; an unpaired bird can forage in a flock, forage alone or sing to attract a mate. Foraging increases his reserves, while singing reduces them. The optimal policy and the expected behaviour of birds depend both on time and reserves. The model predicts that birds will flock, especially in the morning, if flocking birds find more food (foraging efficiency), and also more flocking can be expected when the predation risk is lower in a flock (antipredator benefit). Where flocking gives lower variance in food intake, with the same mean (reduced variance benefit), birds with low reserves at the end of the day choose to forage alone (high variance option), while otherwise they are risk-averse and forage in a flock. The cost of flocking increases with time in a day and with the probability of mate attraction through singing. Decisions inevitably involve trade-offs. Where flocking results in antipredator benefit, but also lower foraging gain, birds with low reserves forage alone, but birds with high reserves flock. Offprint requests to: A.I. Houston     

Correlates of boldness in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)

Behavioural variation is known to occur between individuals of the same population competing for resources. Individuals also vary with respect to their boldness or shyness. An individuals position along the shy-bold axis may be defined as the extent to which it is willing to trade off potentially increased predation risks for possible gains in resources. Similarly, group living may be interpreted as a trade-off between anti-predatory tactics and foraging efficiency. The responses of three-spined sticklebacks (Gasterosteus aculeatus) were tested across four social contexts to assess relative boldness or shyness and to further examine whether their behaviour would be consistent within and between contexts. Individuals displayed consistent responses within and between the first two contexts: those individuals which resumed foraging rapidly after a simulated aerial predator attack also displayed low shoaling tendencies. Such fish were deemed to be bold, whilst those which displayed the converse behaviour, slow resumption of foraging and a high shoaling tendency, were deemed to be shy. In a third context, bold individuals out-competed shy conspecifics for food. Boldness was also positively correlated with growth over a 6-week period. The position adopted by an individual within a group is usually interpreted as a trade-off between predation risk and foraging efficiency—both are greater at the front of a mobile group. Bold individuals showed significantly stronger tendencies towards front positions than shy conspecifics. The results suggest that, contrary to some previous studies on other animals, bold or shy behaviour in sticklebacks is consistent between contexts.Communicated by T. Czeschlik     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.     

Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

Optimal foraging and territorial defence in the Great Tit (Parus major)

Summary We describe an experiment designed to investigate the trade-off between foraging and territorial vigilance in the Great Tit. Captive territorial male Great Tits were observed while foraging in a large indoor aviary. They obtained food from two operant patches in which the supply of food was gradually depleted during a visit. We predicted that during control sessions the birds would switch between patches in such a way as to maximise their overall feeding rate. In experimental sessions, we introduced briefly a rival male as an intruder at the start of the test. The foraging male could see the rival only when travelling between patches and not while feeding within patches. We predicted that during experimental sessions birds would switch between patches more often than in control tests, sacrificing food intake for territorial vigilance. Three of the four males tested behaved in approximately the predicted manner. We discuss the use of an inverse optimality argument to provide a calibration of feeding against the benefit resulting from territorial vigilance.     

Vibrational communication between hitchhikers and foragers in leaf-cutting ants (Atta cephalotes)

Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Communicated by P. Pamilo     

Studies on the nutritional requirements of the larval stages of Penaeus japonicus using microencapsulated diets

Fatty acid biosynthesis in the larval stages of Penaeus japonicus Bate was examined by feeding microencapsulated diets containing (1-¹⁴C) palmitic acid, and fat-free diets supplemented with defined fatty acids. Highest larval growth rates were achieved on diets containing Tapes philippinarum lipid and, when defined fatty acids were substituted, on diets containing 20:63 fatty acid. The radioactive tracer experiments indicate that 16:17, 18:0 and 18:19 fatty acids may be synthesised from palmitic acid and that P. japonicus larvae may possess the ability to elongate 18:33 to 20:53 and 22:63, and 18:26 to 20:46. However, the rates of these reactions appear to be too slow to meet the larval requirements for essential fatty acids and the 3 series of polyunsaturated fatty acids must be provided in the diet.     

Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Digestive carbohydrases from the hepatopancreas of two Antarctic euphausiid species (Euphausia superba and E. crystallorophias)

The hepatopancreatic extracts of Euphausia superba Dana and E. crystallorophias Holt and Tattersall collected from the Antarctic. Ocean during January 1985, are most effective in hydrolysing substrates containing (13)--glucosidic linkages. Three enzymes appear to be involved in the depolymerization of (13)--D-glucans in the euphausiid diet: (1) an exo-(13)--D-glucanase, (2) an endo-(13)--D-glucanase and (3) a -D-glucosidase. The glucanases have a pH optimum of 5.4, a temperature optimum of 50°C and are optimally extracted in bistripropane buffer, pH 7.2. Levels of (13)-D-glucanase in laboratory-cultured E. superba are inversely affected by food availability, and activities double after starvation for 12 d. The increase is due mainly to higher activities of exo-acting glucanases. -Amylase and endo-(14)--D-glucanase (cellulase) activity are also present in the extracts in addition to glycosidase activity against a range of p-nitrophenyl substrates (-and -D-glocose, - and -D-galactose, -D-xylose, - and -D-mannose). Digestive activity against several acidic polysaccharides, including the acidic mucilage polysaccharide of the ice diatom Stauroneis amphioxys, is minimal and is not induced when the polysaccharide is present in the diet of E. superba. These results indicate that some, but not all, components of the algal material in the euphausiid diet can be hydrolyzed and assimilated.     

Information transfer and gain in flocks: the effects of quality and quantity of social information at different neighbour distances

We assessed experimentally how the quality and quantity of social information affected foraging decisions of starlings (Sturnus vulgaris) at different neighbour distances, and how individuals gained social information as a function of head position. Our experimental set up comprised three bottomless enclosures, each housing one individual placed on a line at different distances. The birds in the extreme enclosures were labelled senders and the one in the centre receiver. We manipulated the foraging opportunities of senders (enhanced, natural, no-foraging), and recorded the behaviour of the receiver. In the first experiment, receivers responded to the condition of senders. Their searching rate and food intake increased when senders foraged in enhanced conditions, and decreased in no-foraging conditions, in relation to natural conditions. Scanning was oriented more in the direction of conspecifics when senders behaviour departed from normal. In the second experiment, responses were dose dependent: receivers increased their searching rate and orientated their gaze more towards conspecifics with the number of senders foraging in enhanced food conditions. In no-foraging conditions, receivers decreased their searching and intake rates with the number of senders, but no variation was found in scanning towards conspecifics. Differences in foraging and scanning behaviour between enhanced and no-foraging conditions were much lower when neighbours were separated farther. Overall, information transfer within starling flocks affects individual foraging and scanning behaviour, with receivers monitoring and copying senders behaviour mainly when neighbours are close. Information transfer may be related to predation information (responding to the vigilance of conspecifics) and foraging information (responding to the feeding success of conspecifics). Both sources of information, balanced by neighbour distance, may simultaneously affect the behaviour of individuals in natural conditions.Communicated by H. Kokko     

Optimal foraging: Some simple stochastic models

Summary Some simple stochastic models of optimal foraging are considered. Firstly, mathematical renewal theory is used to make a general model of the combined processes of search, encounter, capture and handling. In the case where patches or prey items are encountered according to a Poisson process the limiting probability distribution of energy gain is found. This distribution is found to be normal and its mean and variance are specified. This result supports the use of Holling's disc equation to specify the rate of energy intake in foraging models. Secondly, a model based on minimization of the probability of death due to an energetic shortfall is presented. The model gives a graphical solution to the problem of optimal choices when mean and variance are related. Thirdly, a worked example using these results is presented. This example suggests that there may be natural relationships between mean and variance which make solutions to the problems of energy maximization and minimization of the probability of starvation similar. Finally, current trends in stochastic modeling of foraging behavior are critically discussed.     

Ecdysial rhythms in megalopae and first instars of the Dungeness crab Cancer magister

Diel molting cycles of megalopae and first instar Dungeness crabs Cancer magister Dana captured in the Grays Harbor estuary (46° 55N; 124° 05 W) in May 1991 were studied under laboratory conditions. Sixtyone percent of the megalopae and 76% of the first instar crabs molted during periods of ambient darkness under a normal light — dark diel regime, and molting pattern was not affected by changes in the photoperiod (24 h daylight or 24 h darkness). Time until metamorphic molt increased as conspecific density increased. Habitat type (shell or mud) did not affect time until molt of megalopae and first instar crabs, nor did it affect daily molting rhythm of first instars. We hypothesize that nightly ecdysis of megalopae and first instar Dungeness crabs and density-dependent molting may be an adaptive response to predation and cannibalism among young-of-the-year.Contribution No. 875 from the School of Fisheries, WH-10, University of Washington, Seattle, Washington     

An instance of bacterial inhibition in oceanic surface water

Water from the 150 surface film 50 miles north of Puerto Rico at 19°30 N, 66°30W supported only a negligible microflora compared to that of adjacent stations. Bacteriostasis was indicated when a ten-fold dilution of the sample resulted in a ninety-fold increase in cultivable microorganisms. The surviving organisms, atypical pseudomonads, were biochemically inactive in comparison to isolates from the other stations. Net phytoplankters at this station were similar in relative composition and abundance to those at the adjacent station.     

Reproductive effort in Adélie penguins

Summary We estimated reproductive effort (energy expenditures for reproduction, as opposed to maintenance) in Adélie penguins breeding at Palmer Station, Antarctica. Data on body composition changes and metabolic rate were obtained using isotopic methods. Adelie breeding behavior consists of an initial courtship stage (during which both sexes fast), incubation, the guard stage (when chicks are 1 to 18–28 days old), and the creche stage (from the end of guarding until chicks are 28–45 days old). Both males and females lost considerable mass during the initial stages of the reproductive season, but males fasted longer and lost more mass. Mass losses of both sexes consisted of 66% depot fat and 34% lean tissue. Mass and body composition remained constant once birds resumed feeding. The metabolic expenditure for the foraging necessary to accumulate the mass lost while fasting — one component of reproductive effort —was about 63 MJ in males and 39 MJ in females. Field metabolic rates (FMR) were low during courtship and while incubating, increasing more than 2-fold when birds resumed foraging. Although mean FMR increased between incubation and the creche stage, differences between stages were small and not significant. We used FMR data and an energy balance model to estimate the cost of feeding chicks. Results suggest a maintenance FMR of about 2.7 × basal metabolism (BMR), increasing to 3.4–3.6 × BMR during the creche stage. The reproductive effort (as metabolic expenditures) associated with feeding chicks is 31 MJ (males) to 36 MJ (females). Cumulative reproductive effort is 94 MJ in males and 75 MJ in females, or 5.3–6.2% of the annual energy budget. The reproductive effort devoted to chick care does not appear to be constrained by physiological or time limitations. Instead, selection to reduce the risk of predation may prevent the evolution of increased parental care. Correspondence to: M.A. Chappell     

Sunset and the orientation of a nocturnal bird migrant: A mirror experiment

Summary If savannah sparrows, (Passerculus sandwichensis), a North American night migrant, select a migratory heading based upon the setting sun, a shift in the position of that cue should produce a predictable shift in the migrant's nocturnal orientation. I tested this hypothesis by shifting the sunst position with mirrors and by recording the bird's orientation in Emlen funnels. The control group displayed directionality appropriate for spring migration (=342°). The mean heading of experimentals (=272°), which were exposed to a cue-shifted situation, was in the expected westerly direction relative to the control mean (P<0.05, V-test). The setting sun appears to be a sufficient source of directional information for this avian migrant.     

Gross biochemical composition of the bivalve Macoma balthica from the Gulf of Gdańsk (Southern Baltic)

Seasonal changes in lipid, glycogen, protein (Nx6.25), and ash content of the meat of Macoma balthica (L.) from five sampling sites in the Gulf of Gdask were studied from 1981 to 1984. Lipid and glycogen were stored mainly during spring and at the beginning of summer. The lipid content of up to 36% of the dry weight observed in populations from the Gulf of Gdask are among the highest values reported for bivalve molluscs. Potential causes of this high value are discussed.     

Nomadic pastoralists adopt subsistence strategies that maximise long-term household survival

Summary Optimality approaches have been used to investigate the adaptiveness of human subsistence strategies mainly in hunter-gatherer societies. However the static optimality models used are not suitable for investigating the long-term costs and benefits of decisions, especially in societies where wealth is accumulated and hence resources gained in one year have an influence on future years. Here I use dynamic optimality modelling to investigate the adaptiveness of a subsistence strategy in the Gabbra, a nomadic pastoralist group. I show that herders forego short-term gain in favour of long-term household survival. This is done by herders sometimes manipulating the life histories of their sheep. In herds where the breeding rate has been slowed, by restricting ewes access to males, females have greater longevity, but fewer offspring per year, than in unmanipulated herds. Models maximising offtake from the herd predict that herd breeding-rate should never be slowed. Models maximising short-term household survival predict herd-breeding rate should always be slowed. Models of long-term household survival predict that herd-breeding rate should be slowed only by relatively wealthy households. This is the behaviour observed. Poor and wealthy house-holds adopt different behaviours, yet all are following optimal strategies for their own level of wealth, that maximise their long-term survival. This is the behaviour that would be predicted on the basis of evolutionary theory, given that a family takes many years to be raised. Correspondence to: R. Mace     

Do kin always make better neighbours?: The effects of territory quality

Summary The effects of territory quality on kin-biased territorial defence behaviour and fitness of juvenile rainbow trout (Oncorhynchus mykiss) were examined by manipulating food and predation risk levels in an artificial stream channel. Groups of related (full sibling) or unrelated fry were observed in the channel under one of four treatments: (1) high food-low predator (high territory quality); (2) high food-high predator; (3) low food-low predator; and (4) low food-high predator (low territory quality). Fish within kin groups always initiated fewer aggressive interactions, defended smaller territories, exhibited higher proportions of threat type territorial defence behaviours (as opposed to overtly aggressive behaviours) and had a higher mean weight increase than non-kin groups. Within both kin and non-kin groups, decreasing territory quality significantly increased the frequency of aggressive interactions, the size of territories and decreased growth. It is argued that while kin-biased territorial defence behaviours are always present, decreased availability of resources and increased predation risk serve to reduce the magnitude of this kin bias. However, even in low-quality territories, the benefit of associating with kin may increase the probability of overwintering survival (increased fitness) of the young fish by increasing body size.