The influence of mercury on the burrowing behaviour ofCorophium volutator

The burrowing behaviour ofCorophium volutator (Pallas) has been recorded in sediments treated with seawater solutions of 0.001, 0.1, 10, and 1000 ppm HgCl₂. Heterotrophic sediment bacteria have also been counted. Non-choice and choice experiments have been conducted. In non-choice experiments no individuals burrowed in the sediment treated with 1000 ppm HgCl₂ solutions. The same numbers of individuals burrowed in the sediments treated with 0.001, 0.1, and 10 ppm HgCl₂ solutions as in the control. Bacterial numbers were lower in the sediments treated with 10 and 1000 ppm HgCl₂ solutions. Hence, a mercury concentration that killed bacteria (10 ppm) did not inhibit burrowing. In choice experiments, individuals avoided all the mercury-treated sediments, particularly that treated with 1000 ppm HgCl₂ solution. Bacterial numbers were only lower than the control in the sediments treated with 10 and 1000 ppm HgCl₂ solutions. Hence, individuals avoided treated sediments (0.001, 0.1 ppm HgCl₂) in which no bacterial mortality had occurred.     

The influence of odour on the oviposition behaviour of the cabbage root fly

Summary. Host plant odours are known to be important in long-range host location by the cabbage root fly, whereas at short distances orientation is mainly visual. We show that olfaction also plays a significant role after a fly lands on a plant and before it moves down onto the soil to oviposit. Host plant acceptance by the cabbage root fly seems to result from a synergistic response to simultaneously perceived olfactory and contact chemostimulation. Received 12 May 1999; accepted 24 July 1999     

The influence of drainage networks on patterns of soil respiration in a desert catchment

Hydrologic flow and connectivity act as important determinants of ecological pattern and process in heterogeneous landscapes. Here we examine how the routing of water through the drainage network of an upper Sonoran Desert basin influences landscape patterns of soil respiration (SR) at both seasonal and event-based timescales. At seasonal timescales, SR varied up to 13-fold with downstream position in the drainage network, and annual estimates of CO2 efflux ranged from 185 g C x m(-2) x yr(-1) to 1190 g C x m(-2) x yr(-1) for sites arrayed along the same flow path. Spatial patterns of SR were unrelated to the carbon and water content of surface soils, but rather tracked changes in plant size and productivity, which in turn reflect downstream increases in groundwater availability. The relative importance of precipitation and temperature as drivers of SR also changed with landscape position, with the latter becoming more important in downstream locations. At the scale of individual precipitation events, SR increased up to 30-fold upon rewetting but typically returned to background levels within 24 h, even when soil moisture remained elevated. Unlike patterns observed at seasonal scales, event-based losses of CO2 varied across the landscape as a function of the organic-matter content in surface soils. Results from labile carbon amendments confirm that CO2 losses following precipitation pulses are initially constrained by substrate availability, not soil drying. By mediating spatial patterns of vegetation structure and soil resource availability, drainage networks represent an important physical template upon which belowground processes are organized in desert basins.     

The feeding behaviour of Schistocerca gregaria,the desert locust,on two starch mutants of Arabidopsis thaliana

Summary. Schistocerca gregaria, the desert locust, has been shown to regulate its dietary intake with respect to specific macronutrients in synthetic foods. This study examined the nutrients in the leaves of two starch mutants of Arabidopsis thaliana, and then compared the feeding behaviour of locusts on the two starch mutants. The high-starch mutant had c. 25 times more starch than the no-starch mutant. Newly molted 5th stadium locusts were preconditioned for 3 days on one of the mutants, and then observed for 90 min while exposed to the same or the alternative mutant. Locusts pretreated with the no-starch mutant fed longer during the first meal on high-starch mutants, spent more time feeding, and had the smaller latency to begin a meal when compared to the locusts pretreated on the high-starch mutant. The results of the study are interpreted in light of an integrative model of nutrient balancing. Received 16 November 1999; accepted 23 December 1999     

The influence of refuge sharing on social behaviour in the lizard Tiliqua rugosa

Refuge sharing by otherwise solitary individuals during periods of inactivity is an integral part of social behaviour and has been suggested to be the precursor to more complex social behaviour. We compared social association patterns of active versus inactive sheltering individuals in the social Australian sleepy lizard, Tiliqua rugosa, to empirically test the hypothesis that refuge sharing facilitates social associations while individuals are active. We fitted 18 neighbouring lizards with Global Positioning System (GPS) recorders to continuously monitor social associations among all individuals, based on location records taken every 10 min for 3 months. Based on these spatial data, we constructed three weighted, undirected social networks. Two networks were based on empirical association data (one for active and one for inactive lizards in their refuges), and a third null model network was based on hypothetical random refuge sharing. We found patterns opposite to the predictions of our hypothesis. Most importantly, association strength was higher in active than in inactive sheltering lizards. That is, individual lizards were more likely to associate with other lizards while active than while inactive and in shelters. Thus, refuge sharing did not lead to increased frequencies of social associations while lizards were active, and we did not find any evidence that refuge sharing was a precursor to sleepy lizard social behaviour. Our study of an unusually social reptile provides both quantitative data on the relationship between refuge sharing and social associations during periods of activity and further insights into the evolution of social behaviour in vertebrates.     

Foraminiferan (Protozoa) epizoites on Arctic isopods (Crustacea) as indicators of isopod behaviour?

A total of 38 219 specimens representing 63 species of marine isopods (Crustacea) from deep and shallow Arctic waters were studied in a search for epizoic foraminifers (Protozoa). Foraminifers occurred on 21 species, and their frequency was generally low. A total of 290 foraminifer individuals were found, of which 289 belonged to Cibicides wuellerstorfi, C. refulgens and Cibicides spp. (juveniles) (Cibicidae), while only a single individual belonged to Cornuspira sp. (Cornuspiridae). The foraminifers were most frequent on species of the families Munnidae, Ischnomesidae (suborder Asellota) and on Gnathia stygia (suborder Gnathiidea), but were totally absent from the asellote families Janiridae, Haploniscidae, Nannoniscidae and from the suborder Epicaridea. The foraminifers were mainly located on the legs (Munna acanthifera), the anterior part of the body (Haplomesus quadrispinosus, heteromesus frigidus), or on the head (G. stygia adults). The epizoic foraminifers occur mainly on epibenthic isopods, which do not or only rarely clean themselves. The foraminifers are known to prefer elevated substrata, and in this the habitat of the isopods and the foraminifers coincide. The size of individual isopods was not related to the presence or absence of foraminifers.     

Ecological and social determinants of spacing behaviour in desert baboon groups

This paper investigates the determinants of individual spacing behaviour in a desert baboon population (Papio cynocephalus ursinus). Patterns of neighbour proximity and neighbour density were examined among adults in four groups under different ecological and social conditions (through instantaneous sampling during focal follows). Initial analysis of these data shows that (1) the use of vertical substrates (refuges such as tall trees and cliff faces) can confound patterns of spacing, and (2) individual differences in spacing can depend on the spatial scale over which it is measured. To minimise these substrate and scale effects, this analysis focuses on animals which are off refuges and examines spacing behaviour through its underlying statistical `dimensions' (identified through factor analysis). Analysis of these dimensions indicates that sex, group size, activity-habitat and female reproductive state can all have independent effects on spacing: (1) males are more dispersed than females in smaller groups, (2) male and female dispersion increases with time spent in foraging habitats, and (3) female dispersion is reduced during lactation. According to the hypotheses tested, these results indicate that feeding competition only affects spacing behaviour during foraging while predation risk plays little or no role in spacing. Most aspects of spacing behaviour are best explained by male reproductive strategies and their social repercussions. Received: 25 May 1998 / Accepted after revision: 18 July 1998     

The tongue-replacing isopod Cymothoa borbonica reduces the growth of largespot pompano Trachinotus botla

Cymothoa borbonica prevalence in the buccal cavity of Trachinotus botla was high, with 45 % of all fish sampled being infected. Smaller fish were more susceptible to infection with no parasites found in fish over 400 mm FL. The detrimental effects of parasite infection on their hosts include basihyal (the bone commonly known as the “tongue”) damage, a loss in buccal cavity volume as a result of female parasite attachment, and a severe impact on host growth. By combining short-term dietary analysis and medium-term stable isotope analysis, there was little evidence to suggest a modification in either the diet or feeding habits of infected fish where infected and uninfected fish occupied the same trophic niche. Inhibited growth in infected fish is hypothesized to be from respiratory distress from long-term oxygen deficiency through buccal obstruction.     

Population energetics of the intertidal isopod Cirolana harfordi

Laboratory experiments and field measurements generated a population energy-budget estimate for the isopod Cirolana harfordi (Lockington). Assimilation of food energy averaged 88% on a diet of fish. About 35% of assimilated energy is allocated for growth and reproduction, while the bulk of the remainder is used for maintenance and activity (respiration). The high growth efficiency of C. harfordi is discussed with respect to ecological efficiency and energy transfer in marine systems.     

Reproductive biology of a small isopod symbiont living on a large isopod host: from the maternal marsupium to the protective grip of guarding males

Mating systems of many symbiotic crustaceans are characterised by a high degree of mate guarding. A peculiar case of mate guarding has been reported for small symbiotic janirid isopods where males mate with immature females. Field samples of individual hosts and laboratory experiments were conducted to reveal the mating behaviour of the symbiont in a natural environment, that is, on their hosts. Along the coast of the Magellan Strait, Chile, the janirid isopod Iais pubescens was frequently found on the shore-living isopod Exosphaeroma gigas. Symbiont prevalence (percent hosts occupied) was high at eight of the nine sampling sites. Mean symbiont intensity was very low at one site (<<1 individual host^-1), intermediate at two sites (1-10 individuals host^-1) and high at the other sites (10-40 individuals host^-1). The mean sex ratio (males:females) was male biased at most sampling sites (n=7). Females of I. pubescens reached substantially larger sizes (1.5-3.0 mm body length, BL) than males (1.1-1.9 mm BL). The majority of males were carrying small juveniles (66.15%), and males with juveniles were significantly larger than males without juveniles - this suggests that males prefer virgin juveniles to adult females and that they compete for small juveniles. In laboratory observations, males were seen to manipulate the marsupium of adult females that were about to release small juveniles. Males obtained virgin juveniles in this manner. Juveniles were carried for ~7 days, and they moulted shortly before being fertilised and released by males. The high proportion of juveniles carried by males in the field (68.2%) supports previous observations that males initially are not able to distinguish male and female juveniles. It is suggested that the mating system of symbiotic janirid isopods with long-term sperm storage and continuous receptivity in females and male mating with virgin females has evolved in response to highly unpredictable encounter probabilities between the sexes. Mate guarding and manipulation of small virgin juveniles may be favoured on the highly mobile hosts of symbiotic janirid isopods. Furthermore, adult females may gain by leaving their emerging offspring in the protective grip of guarding males, thereby reinforcing the maintenance of this peculiar mating system.     

Individual-based modelling of fishermen search behaviour with neural networks and reinforcement learning

A model to mimic the search behaviour of fishermen is built with two neural networks to cope with two separate decision-making processes in fishing activities. One neural network deals with decisions to stay or move to new fishing grounds and the other is constructed for the purpose of finding prey within the fishing areas. Some similarities with the behaviour of real fishermen are found: concentrated local search once a prey has been located to increase the probability of remaining near a prey patch and the straightforward movement to other fishing grounds. The artificial fisherman prefers areas near the port when conditions in different fishing grounds are similar or when there is high uncertainty in its world. In the latter case a reluctance to navigate to other areas is observed. The artificial fisherman selects areas with higher concentration of prey, even if they are far from the port of departure, unless a high uncertainty is related to the fishing ground. Connected areas are preferred and followed in orderly fashion if a higher catch is expected. The observed behaviour of the artificial fisherman in uncertain scenarios can be described as a risk-averse attitude. The approach seems appropriate for an individual-based modelling of fishery systems, focusing on the learning and adaptive characteristics of fishermen and on interactions that take place at a fine scale.     

Assimilation efficiency of the antarctic marine isopod Glyptonotus antarcticus

When fed shrimp every 2 days, the isopod Glyptonotus antarcticus Eights assimilated over 90% of the ingested food. Errors in this estimate due to loss of food material during maceration by G. antarcticus were investigated and allowed for. These results are discussed in relation to data for other isopods and marine invertebrates.     

Effect of temperature on the microdistribution of the isopod Sphaeroma rugicauda from a saltmarsh habitat

The saltmarsh isopod Sphaeroma rugicauda (Leach) is subjected to widespread diurnal and seasonal temperature fluctuations under natural conditions. Laboratory studies on its activity show that there is a relationship between behaviour activity and exposure temperature between 2.5° and 25°C. Although S. rugicauda has no complex metabolic adaptations, this isopod is able to maintain a rate of aerial oxygen consumption which is similar to that in water within the temperature range 5° to 25°C. The responses of S. rugicauda to changes in environmental temperature are discussed in relation to the seasonal microdistribution of the isopod in the salt-marsh habitat.     

Health and population-dependent effects of ocean acidification on the marine isopod Idotea balthica

Three populations of the grazing isopod Idotea balthica were exposed to high CO₂ treatment for a period of 20 days to investigate the effect of ocean acidification (OA) on animal health and immunocompetence. The results of the populations from more saline habitats were comparable and showed a 60–80 % decrease in immune response as a result of the high CO₂ treatment. Analysis of protein carbonyls showed no treatment effect, indicating that short-term OA does not increase oxidative protein damage. Meanwhile, the third tested population from the lower saline Baltic Sea had higher background protein carbonyl levels. Ocean acidification in addition to this resulted in 100 % mortality. The results of this study show that OA reduced immunocompetence of this marine isopod. In addition, populations and individuals in poor health are potentially at greater risk to succumb under OA.     

The influence of social experience on cooperative behaviour of rats (Rattus norvegicus): direct vs generalised reciprocity

Cooperation among non-kin has been attributed sometimes to reciprocal altruism: Two or more individuals exchange behaviour that benefits the respective partner. According to direct reciprocity, cooperation is based on past behaviour of a known partner. In contrast, in generalised reciprocity, cooperation is based on anonymous social experience where the identity of the partner is irrelevant. In a previous study, female Norway rats (Rattus norvegicus) were found to cooperate according to a generalised reciprocity mechanism. In this study, we tested whether Norway rats would also cooperate as predicted by a direct reciprocity mechanism and whether direct reciprocity would cause a higher propensity to cooperate than generalised reciprocity. Focal animals were experimentally manipulated to receive social experience from known or unknown, helpful or defecting partners in an instrumental cooperative task. Our first experiment shows that rats are more helpful towards a partner from which they had received help before than towards a partner that had not helped (i.e. direct reciprocity). Our second experiment revealed that after receiving help by others, rats were more helpful towards a partner from which they had received help before than towards a new partner (i.e. direct reciprocity generated a higher cooperation propensity than generalised reciprocity). We conclude that in female Norway rats, the tendency to cooperate is influenced by partner-specific information. To our knowledge, this is the first study to demonstrate direct reciprocity in rodents, and it is the first study testing direct vs generalised reciprocity in animals.     

Biology and population dynamics of the intertidal isopod Cirolana harfordi

The distribution of Cirolana harfordi (Lockington) populations is determined largely by the availability of loose boulders on sandy beaches. The isopods swim out from under the rocks at high tide to feed. Their diet consists primarily of minute polychaetes and crustaceans. In addition, the isopods locate and utilize any available dead animal matter in the surf zone. Breeding occurs throughout most of the year, except for a brief lull in the fall. Females produce 1 or 2 broods of 18 to 68 young during their 2-year life-span. Marsupial incubation lasts 3 to 4 months. The population size and structure remained relatively constant from one year to the next. Mortality rates estimated for juveniles, males and females showed that newly emergent young and post-reproductive females suffer the greatest losses (up to 75% mortality per month).     

The effects of size-dependent predation risk on the interaction between behavioral and life history traits in a stream-dwelling isopod

In field surveys, laboratory observations and field-based assays of behavior, I examined the effects of size-dependent predation risk on the interaction between size at reproductive maturity and maternal care behavior in the stream-dwelling isopod, Lirceus fontinalis. L. fontinalis exhibit population-specific sizes at reproductive maturity which result in population differences in predation risk during the adult phase. Females from streams containing salamander larvae (that prefer small prey) mature at large sizes and then become relatively safe from predation. Females from streams containing fish (that consume all size classes of prey equally) mature at small sizes and remain at risk. I tested whether these differences in expected survival were reflected in the behavior of females during the maternal phase (i.e., the period during which females exhibit maternal care). Female L. fontinalis carry developing juveniles inside a brood pouch. I simulated predatory attacks on gravid female L. fontinalis from the different population types and found that female behavior correlated with population differences in risk. When “attacked”, females from streams with predatory fish (that experience high risk to adult females) released juveniles from the brood pouch, whereas females from populations with predatory salamander larvae (that pose relatively little risk to adult females) did not release juveniles. I discuss the results with reference to the joint evolution of behavioral and life history traits. Received: 6 March 1996 /Accepted after revision: 12 August 1996     

The superseded female's dilemma: ultimate and proximate factors that influence guarding behaviour of the carpenter bee Xylocopa pubescens

Both solitary and primitively social nests of the facultatively social carpenter bee Xylocopa pubescens can be found throughout most of the breeding season. In social nests there is reproductive division of labour between a dominant forager and a guarding female. Two types of guarding females can be discerned: the young pre-reproductive guards, and older, formerly reproductive guards. The latter type of guard is found when, after a take-over of reproductive dominance either by a nestmate (mostly a daughter) or an intruder, the defeated female stays in the nest instead of leaving to try and found or usurp another nest. She is then manipulated into the role of a guard. The dominant female profits from the presence of the guard since she protects the nest against pollen robbery by conspecifics (Hogendoorn and Velthuis 1993). We have studied why superseded females might prefer to remain as a guard, rather than try their luck somewhere else. The hypotheses investigated pertain to (1) the difficulty for the defeated female of finding a new nest and of restarting reproductive activities due to (a) ecological constraints (nest and pollen shortage) and (b) the effect of age and wear on the defeated female; (2) the effects of guarding in terms of inclusive fitness. We found that superseded females remained as guards significantly more often when a nestmate (not necessarily close kin) took over reproductive dominance than when an intruder did so. Other factors associated with the decision of the defeated female to stay or leave were her age and the number of her own young still present after the supersedure. The probability of finding or constructing a new nest was lower for old than for young females. After finding a nest, old females produced less brood than young foundresses. As a result of these two factors old superseded females gained, in terms of inclusive fitness, by staying as guards, whereas young females profited from leaving the nest. We interpret these results as an indication that guarding behaviour has evolved due to kin selection. However, kin discrimination apparently did not occur. Therefore we conclude that in this species kin selection is not, in the proximate frame of reference, based on kin recognition and preference for helping kin. Correspondence to: K. Hogendoorn