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.     

The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

Circadian rhythmicity of locomotor behavior and oviposition in female Teleogryllus commodus

Summary Locomotor activity of the Teleogryllus commodus is under circadian control, with _LL=25.3 h and _DD=23.4 h. In LD 12:12 h, running occurs either exclusively during darkness (57%), mainly in the dark phase (35%), or with substantial activity peaks in both phases (8%). For oviposition, _LL>24 h and _DD<24 h; in LD 12:12 h, 80% of the studied deposited their eggs mainly during the light phase and 20% primarily during darkness. In the discussion, a temporal correlation between stridulation, spermatophore formation, and locomotion, and oviposition is established, which serves to make an encounter and mating between the sexes as likely as possible.Dedicated to Prof. Dr. M. Gersch on the occasion of his 70th birthday     

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.     

Reproductive effort in relation to maternal social rank in reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves body-mass change during their first winter). In the models, we included September females body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females social rank, whereas late preweaning daily growth rate of calves was not influenced by females rank. Calves preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.Communicated by R. Gibson     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

Lead-based paint in dwellings: The potential for contamination of the home environment during renovation

The quantity and particle size characteristics of lead in dust released during three different paint removal techniques was determined under controlled conditions and in situ in a dwelling. Air-lead and deposited dust-lead levels were highest after sanding but burning-off and hot-air removal methods also produced significant contamination. The importance of dust particle-size and lead is discussed in relation to the potential hazard to home renovators via inhalation and to children via the hand-to-mouth route.This paper is also being published in Proceedings of the Conference on Lead in the Home Environment and is one of several selected from SEGH sponsored conferences for simultaneous publication in this journal.     

Metabolism of a nonylphenol isomer by Sphingomonas sp. strain TTNP3

For elucidation of the metabolism of the endocrine disruptor nonylphenol by Sphingomonas sp. strain TTNP3, the degradation of an isomer of nonylphenol, 4(2,6-dimethyl-2-heptyl)-phenol, has been studied. As in the case of 4(3,5-dimethyl-3-heptyl)-phenol, the metabolism of this nonylphenol isomer leads to the formation of the NIH-shifted product 2(2,6-dimethyl-2-heptyl)-1,4-benzenediol (NIH: National Institute of Health), but also to the alkoxy derivative 4(2,6-dimethylheptan-2-yloxy)phenol as additional metabolite. To the best of our knowledge, this is the first report describing the formation of alkoxyphenol as a degradation product of nonylphenol. Additionally, these results provide for the first time evidence for slight differences in the biodegradation of the isomers of nonylphenol.     

Song as an attractant for male and female European starlings,and the influence of song complexity on their response

Summary We tested the hypothesis that song attracts females and repels males in the European starling. We broadcast recorded song from speakers attached to nestboxes, while paired boxes with silent speakers served as controls. As predicted, females were attracted to the song boxes. Contrary to prediction, males were also attracted to the `song' boxes. Singing by male starlings may be costly because it attracts competitors for limited nesting sites, but the cost cannot be avoided due to the need to attract a mate. In a second experiment simple song (composed of 20 different phrase types) and complex song (40 phrase types) were played simultaneously. More males were captured at boxes where simple song was played. Song complexity may function as an indicator of male quality and be used by male starlings to assess potential competitors.     

The erect ‘penis’ is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas

In East Africa, spotted hyenas live in large clans in a highly structured society dominated by females. A clan is a fission-fusion society where members are often solitary or in small groups. Spotted hyenas have a ritualized greeting during which two individuals stand parallel and face in opposite directions. Both individuals usually lift their hind leg and sniff or lick the anogenital region of the other. The unique aspect of greetings between individuals is the prominent role of the erect penis in animals of both sex. Female spotted hyenas have fused outer labiae and a pseudo-penis formed by the clitoris which closely resembles the male penis and can be erected. During greetings subordinates signalled submission with gestures which were not necessarily reciprocated by the dominant participant. Asymmetries were most pronounced in greetings between adult females where the probability of asymmetries increased with the divergence in rank between partners. Greetings between adult females and males were uncommon and restricted to males above median rank, principally the alpha male. Models of primate affiliative behavior assume that benefits derived from social relationships with different individuals are not equal and that individuals are selected to maximize the benefits they receive from social relationships with others. The observed distribution of greetings between partners of different rank matched the predictions of these models. An examination of non-adaptive hypotheses on the evolution of the pseudo-penis demonstrated that the conventional scenario linking (initial) virilization of female genitalia with selection for female dominance does not explain either the initial virilization, nor the evolution of the pseudo-penis to its current form and use. We sketch a new scenario that links (1) initial virilization to the occurrence of neonatal siblicide amongst members of a twin litter, and (2) costs of maintenance, pseudo-penile control over copulation and male submission. Our analysis confirms previous adaptive hypotheses on the function of the pseudo-penis in greetings and suggests new hypotheses to account for hitherto unexplained features. Correspondence to: M.L. East     

Effects of monospecific algal diets of varying biochemical composition on the growth and survival of Pacific oyster (Crassostrea gigas) larvae

Using monospecific diets of Thalassiosira pseudonana cells grown under different steady-state conditions, it was determined that higher growth rates of larval Crassostrea gigas Thunberg were obtained when fed T. pseudonana cells grown under high light. High light grown T. pseudonana cells consistently contained relatively more of the saturated fatty acids 14:0 and 16:0. Considered over three independent experiments, high light grown T. pseudonana cells were lower in protein and higher in carbohydrate than low light grown cells. Higher growth rates of larval C. gigas were obtained on diets with more of the essential fatty acid (EFA) 22:63, and less of the other EFA, 20:53. The relative requirements of C. gigas larvae for the essential fatty acids 20:53 and 22:63 are discussed. Faster growing larvae contained higher percentages of the fatty acids 14:0 and 16:0, and lower percentages of 22:2j. Oyster growth rates were correlated with their content of the fatty acids: 14:0, 16:0 and 22:2j in two experiments utilizing separately spawned batches of larvae. Fatty acid profiles are proposed as a technique for assessing larval condition. C. gigas larvae contained ten times the percent composition of the FAs 16:43, 18:17, 20:17 and 22:2j compared with their diet. Correlation analysis suggests that the dietary source of 18:17, 20:17 and 22:2j was 16:17. It is concluded that T. pseudonana cells grown under high light are a superior diet for C. gigas larvae in comparison with low light grown cells of the same species.     

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     

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     

Arsenic tissue concentration of immature mice one hour after oral exposure to gold mine tailings

A potentially high bioavailability of arsenic in gold mine tailings from a site in northern California has been suggested by solubility studies. To help address this issue, an in vivo dosing study was conducted using 12dayold Swiss Webster mouse pups (n=8/group). A sample of sizefractionated mine tailings from the site (<20m particle size, 691g g^–1 arsenic) was prepared as an aqueous suspension and administered by gavage in a volume that provided 4mg As/kg body weight. The control group received the same volume of a commercial soil (1g g^–1 As) of similar particle size (<60m). No mortality or toxic signs were noted in either group. Tissue samples were collected 1h after gavage, freezedried, microwavedigested and analysed for arsenic by ICP/MS (detection limit 2ng As g^–1 dry weight). Arsenic concentrations (ng As g^–1 dry weight) in tissues from the pups who received mine tailings were significantly higher than in control tissues. The mean elevation in arsenic concentration was highest in the liver (3364% of control, p<0.0001), followed by blood (818 of control, p<0.0001), skin (207% of control, p=0.07), and brain (143% of control, p<0.0001). The carcass arsenic concentration (excluding the GI tract, liver, brain and skin) was 138 of control (p=0.02). The data indicate uptake of arsenic from weathered mine tailings by the immature mouse pups after oral exposure.     

Importance of the fatty acids 20:5ω 3 and 22:6ω 3 in the diet of plaice (Pleuronectes platessa) larvae

The requirement of plaice (Pleuronectes platessa L.) larvae for the fatty acids 20:5 3 and 22:6 3 was studied. Larvae were reared from first feeding to beyond metamorphosis on Artemia sp. nauplii (EG brand), whose nutritional content had been manipulated by enrichment. Some larvae were fed Artemia rich in 20:5 3 and 22:6 3, and others were fed nauplii with no 22:6 3 and low levels of 20:5 3 (1.9% of the total fatty acids). The differences in diet had no significant difference on growth or survival of the plaice larvae. The results indicate that the plaice larvae in this study did not require 22:6 3 in their diet, even when the levels of 20:5 3 were low.     

Aggregation and foraging behavior of whirligig beetles (Gyrinidae)

Summary Whirligig beetles aggregate in the daytime into dense single-and multispecies groups (rafts) of hundreds or thousands of individuals. On the 22km shoreline of Lake Itasca in northern Minnesota, these aggregations were on the average 0.8 km apart, and they were usually found day after day in the same ocations.Most beetles apparently do not home to the aggregation of their origin after dispersing at night because (a) the species composition of some aggregations changed greatly, and (b) paint-marked beetles (Dineutus horni) moved overnight from one aggregation as far as 4km, joining 11 of the 14 large (>300 beetles) D. horni groups on the lake.Throughout the night, the largest concentrations of beetles remained within 100m of the diurnal aggregation sites. Beetles reconvened into the compact rafts before daybreak, in part by following each other in sometimes long single files or trains. Their forward motion stopped after they joined large number of other beetles. We infer that following behavior enables those individuals that have dispersed from their original aggregations (during their nocturnal foraging) to find and join other aggregations before daylight.Naive fish ate the beetles despite their noxious secretions. However, fish living near rafting sites and feeding on insects on the water surface in daylight should soon learn to avoid the beetles. The rafting sites would then become safe places. We observed fish attacking only those beetles that had been either dispersed from their rafts or released into open water away from raft sites in the daytime. We speculate that the evolutionary significance of the aggregation behavior is related to predator (fish) avoidance.     

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.     

Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

Low primary-productivity levels in high-nutrient environments of the Southern Ocean are not yet well understood. An accurate knowledge of nutrient assimilation by phytoplankton, at the base of the pelagic food-web, therefore appears to be essential. A study of the mesoscale hydrological structure and chemical observations in the upper layers of the Indian sector of the Antarctic Ocean (Cruise MD-25 FIBEX of M.S. Marion Dufresne, January–February 1981) permitted us to develop a method for estimating nutrient consumption by phytoplankton. After strong vertical mixing in winter, the homogeneous Antarctic Surface Water (ASW) is, in summer, divided into two parts by a well-formed pycnocline. In the upper layer, corresponding approximately to the photic zone, we observed nutrient depletion resulting from photosynthetic activity, while the underlaying thermal minimum layer displayed unaltered winter characteristics, including nutrient concentrations typical of winter surface-water conditions. Taking into account the nutrient depletion in summer, we calculated the assimilation ratios for Antarctic phytoplankton as follows: ([NO ₃ ^- lower]-[NO ₃ ^- upper]):([PO ₄ upper])=N:P=11.2±2.4 and N:Si=0.27±0.05. These ratios are lower than the classical Redfield ratios (N:P=16 and N:Si=1), but are in good agreement with the elementary composition of phytoplankton samples collected during the same cruise. Moreover, they agree with previously published data on stoechiometric determinations of particulate matter in surface-water samples. Besides, in the transition layer between the lower layer and Warm Deep Water (WDW), the calculated mineralization ratios were slightly higher than the assimilation ratios: N:P=14.6 and N:Si=0.37. From these ratios, it would appear that, in these surface layers of the Southern Ocean (down to 250 m), assimilation-regeneration mechanisms operate in such a way that phosphate and silicate contents decrease much more than nitrate content during the northward drift of the surface waters. The determination of assimilation ratios taking into account the nutrient depletion of the upper layer allowed us to evaluate the carbon net-production which integrates time-space variations in the photic layer over the whole early summer period. We estimated a production rate (0.4 g C m^-2 d^-1) of the same order of magnitude as the mean value obtained by the ¹⁴C method (0.2 g C m^-2 d^-1). The theoretic silicium consumption by phytoplankton along a meridian transect, calculated using an estimated assimilation ratio of C:Si=1,85, indicates that the marked south-north decrease in silicate concentration in the Antarctic Surface Water would mainly result from biological activity. The silicate concentrations thus calculated were in good agreement with concentrations measured along a previous transect across the Southern Ocean.

---

Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

     

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.     

Environmental influence on the locomotor activity rhythm of the sand-shore amphipod Talorchestia deshayesi

The precision and persistence of the nocturnal, surface locomotor activity rhythm of the sand-beach amphipod Talorchestia deshayesi is described. The rhythm is expressed as a circadian activity period with no evidence of a circa-tidal component associated with the time of high tide. The rhythm is light-sensitive and entrainment occurs in response to changes in the time of an experimentally simulated dusk transition. Locomotor activity is initiated at the end of the dusk transition with cessation always occurring prior to dawn. The point of activity initiation is taken to be the position of a synchronizing cue controlling entrainment. The cue appears to be an absolute irradiance level of approximately 1.0 to 2.0×10^-4 W m^-2