Brood neglect and contingent foraging behavior in a pelagic seabird

Among species where there is a risk to leaving offspring unattended, parents usually take alternating shifts guarding their young. However, they may occasionally exhibit brood neglect by leaving their offspring unattended at the nest. To investigate this phenomenon further, we examined the foraging behavior of the northern gannet (Morus bassanus) during chick-rearing. This species has a prolonged nestling period (13 weeks) and the single chick is usually guarded by one or other of its parents, because unattended chicks are frequently attacked by conspecifics. We tested the prediction that the foraging behavior of adults when they left their offspring alone at the nest (unattended trips) would differ in character to when adults left offspring with their partner (attended trips). Brood neglect typically occurred after a longer-than-average attendance period at the nest. Unattended trips were, on average, about half the duration of attended trips, and therefore much closer to the colony. There was also a difference in departure direction between attended and unattended trips, with unattended trips tending to be northeast of the colony. Chicks were fed by parents after both attended and unattended trips, but the frequency and the duration of unattended trips increased as chicks grew older whereas the duration of attended trips decreased as chicks grew. These results indicate that parents may be making a trade-off between risk of attack to their offspring and food-intake rate, and that the solution to this trade-off is dependent on chick age.Communicated by C. Brown     

Environmental determinants of distribution and foraging behaviour of cormorants (Phalacrocorax spp.) in temperate estuarine habitats

The distribution and behaviour of cormorants in estuarine environments was examined on the central coast of New South Wales, Australia, with respect to habitat associations at different spatial scales. No consistent variation in abundance was found for four species of cormorants (great Phalacrocorax carbo, pied P. varius, little black P. sulcirostris, and little pied P. melanoleucos) with state of tide (high and low) and time of day (early, middle, and late) at five estuarine locations. Differences in abundance were found among locations that were not confounded by short-term temporal variation (i.e. time of day and tide). Differences in abundance were detected among habitats (e.g. bays, creeks, and headlands) separated by hundreds of metres to kilometres in different estuaries. Cormorants of all species were rare on the open coast and near the entrance of estuaries. Abundances of cormorants varied greatly within and among creeks, bays, and river channels. Presence of seagrass beds explained much of this variation and most of the cormorants swimming and feeding were found near seagrass. Mapping of seagrass beds and the positions of cormorants at scales of metres to hundreds of metres showed a close relationship between the presence of swimming and roosting beds and the presence of seagrass beds for P. melanoleucos and for P. carbo. We argue that cormorants make decisions to visit particular estuarine habitats, especially those with seagrass, where many types of prey (e.g. fish and crustaceans) are probably most abundant. These choices must be interpreted in the context of decisions that cormorants make on scales of hundreds to thousands of kilometres during periodic excursions to the interior of Australia. Further, environmental threats to seagrass beds could impinge on these mobile visitors to the same extent as on more permanent residents. Received: 14 February 2000 / Accepted: 17 July 2000     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Directional change in a suite of foraging behaviors in tropical and temperate evolved honey bees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

The concept of a suite of foraging behaviors was introduced as a set of traits showing associative directional change as a characterization of adaptive evolution. I report how naturally selected differential sucrose response thresholds directionally affected a suite of honey bee foraging behaviors. Africanized and European honey bees were tested for their proboscis extension response thresholds to ascending sucrose concentrations, reared in common European colonies and, captured returning from their earliest observed foraging flight. Race constrained sucrose response threshold such that Africanized bees had significantly lower sucrose response thresholds. A Cox proportional hazards regression model of honey bee race and sucrose response threshold indicated that Africanized bees were 29% (P<0.01) more at risk to forage over the 30-day experimental period. Sucrose response threshold organized age of first foraging such that each unit decrease in sucrose response threshold increased risk to forage by 14.3% (P<0.0001). Africanized bees were more likely to return as pollen and water foragers than European foragers. Africanized foragers returned with nectar that was significantly less concentrated than European foragers. A comparative analysis of artificial and naturally selected populations with differential sucrose response thresholds and the common suite of directional change in foraging behaviors is discussed. A suite of foraging behaviors changed with a change in sucrose response threshold that appeared as a product of functional ecological adaptation.Communicated by R.F.A. Moritz     

Variation in zonal recovery in four Talitrus saltator populations from different coastlines: a comparison of orientation in the field and in an experimental arena

Under stressful conditions (e.g. finding themselves on dry or moisture-saturated substrates) littoral talitrids (Crustacea, Amphipoda) demonstrate zonal orientation, in which they must promptly reach the optimal zone of the beach, the wet fringe near the shoreline. A relationship might therefore exist between the use of orientation and the frequency of such stressful conditions in the natural environment. Moreover, the efficiency of orientation toward the sea could be related to the possibility of using strategies other than zonal orientation in order to avoid stress. This study analysed the actual use and efficiency of orientation under natural conditions of four Talitrus saltator (Montagu, 1808) populations from Mediterranean and northern European Atlantic coasts with different ecological features. Orientation tests were carried out on the beach with all natural cues available. Then the same individuals underwent control experiments to study their sun orientation far from the sea in an experimental arena. The following results emerge from the comparison of the circular distributions: (1) marked differences among populations in the precision of zonal recovery under natural conditions; (2) a common solar orientation capacity in the control tests far from the sea; (3) different orientation choices of the same individuals according to the test conditions, natural or controlled. The habitat diversity of the four populations (amount, distribution and kind of detritus and wrack on the beach, degree of coastal erosion, orientation of the shoreline, human use of the beach) provides an ecological interpretation for the differences in orientation observed among populations. Received: 13 October 1997 / Accepted after revision: 26 April 1998     

Differential responses of honeybee (<Emphasis Type="BoldItalic">Apis mellifera</Emphasis>) patrilines to changes in stimuli for the generalist tasks of nursing and foraging

Which task a social insect worker engages in is influenced by the worker’s age, genotype and the colony’s needs. In the honeybee, Apis mellifera, genotype influences both the age a worker switches tasks and its propensity of engaging in specialist tasks, such as water collecting, which only some workers will perform. In this study, we used colonies with natural levels of genetic diversity and manipulated colony age demography to drastically increase the stimuli for the generalist tasks of foraging and nursing, which all workers are thought to engage in at some point in their lives. We examined the representation of worker patrilines engaged in nursing and foraging before and after the perturbation. The representation of patrilines among foragers and nurses differed from that of their overall colony’s population. In the case of foraging, over- and underrepresentation of some patrilines was not simply due to differences in rates of development among patrilines. We show that replacement foragers tend to be drawn from patrilines that were overrepresented among foragers before the perturbation, suggesting that there is a genetic component to the tendency to engage in foraging. In contrast, the representation of patrilines in replacement nurses differed from that in the unperturbed nursing population. Our results show that there is a genetic influence on even the generalist tasks of foraging and nursing, and that the way patrilines in genetically diverse colonies respond to increases in task stimuli depends upon the task. The possible significance of this genetic influence on task allocation is discussed. Electronic supplementary material Supplementary material is available in the online version of this article at doi: and is accessible to authorized users.     

The impact of behavioral and physiological maternal effects on offspring sex ratio in the common snapping turtle,<Emphasis Type="Italic"> Chelydra serpentina</Emphasis>

Theory suggests that maternal effects are especially important in organisms with environmentally-sensitive sex-determining mechanisms. However, there is no substantive body of empirical evidence to confirm this conjecture. We integrated field and laboratory studies to jointly evaluate the significance of behavioral (nest-site choice) and physiological (yolk hormone allocation) maternal effects on offspring sex ratio in the common snapping turtle (Chelydra serpentina), a species with temperature-dependent sex determination (TSD). Of the 16 microhabitat variables measured, only three (south, east, and total overstory vegetation cover) were significantly correlated with nest temperature: cooler nests were located under more vegetation cover. In turn, these microhabitat predictors of nest temperature, and nest temperature itself, may influence nest sex ratio: shadier, cooler nests were more likely to produce a higher proportion of male offspring than less shady, warmer nests. Analysis of eggs from these same nests incubated in a common garden design in the laboratory revealed that clutch sex ratio was unaffected by levels of yolk estradiol, yolk testosterone, or their interaction. Examination of both behavioral and physiological maternal effects revealed no concordant impact on offspring sex ratio. However, eggs from nests that produced male-biased sex ratios in the field yielded higher proportions of males under constant-temperature conditions in the laboratory. Our study confirms the importance of behavioral maternal effects in nature on offspring sex ratios in species with TSD, while also revealing the potential presence of a predisposition for sex-ratio production underlying TSD in this system.Communicated by S. Krackow     

How resources and encounters affect the distribution of foraging activity in a seed-harvesting ant

We examined how the foraging ecology of the seed-harvesting ant Messor andrei depends upon the distribution of resources and the presence of conspecifics. Bait experiments showed that colonies can recruit to high-density patches of seeds. However, at the seasonal scale, natural resource distribution did not affect the distribution of foraging activity. We conducted the study in years of high rainfall and thus seed availability may not have been a limiting factor. Colonies always preferred to forage in areas closer to their nest, which may reduce travel time between the nest and foraging sites. On a day-to-day scale, encounters between neighboring colonies at a site increased the probability that colonies would return to forage at that site; this was true both for natural and experimental encounters. In the summer, this resulted in colonies foraging at the sites of intraspecific encounters on more days than in areas where no encounter had occurred. Encounters between colonies included fighting, and there was little overlap between the foraging areas of neighboring colonies: both results suggest that one function of encounters is to defend foraging space. The high probability of return to the site of an encounter between colonies suggests that encounters may have a second function: to indicate the presence of resources. Received: 28 June 1999 / Received in revised form: 12 October 1999 / Accepted: 16 October 1999     

Sex allocation in a monomorphic seabird with a single-egg clutch: test of the environment, mate quality, and female condition hypotheses

Sex allocation theory posits that mothers should preferentially invest in sons when environmental conditions are favorable for breeding, their mates are of high quality, or they are in good body condition. We tested these three hypotheses in rhinoceros auklets (Cerorhinca monocerata), monomorphic seabirds that lay a single-egg clutch, in 2 years that differed in environmental conditions for breeding. Results supported the environment and mate quality hypotheses, but these effects were interactive: offspring sex was independent of paternal traits in the poor year for breeding, while females mated to larger and more ornamented males reared more sons in the better year. Conversely, offspring sex was unrelated to female condition, as indexed by hatching date. We propose that good rearing conditions enable females to rear sons possessing the desirable phenotypic attributes of their mates. Results also supported two critical assumptions of sex allocation theory: (1) dimorphism in offspring condition at independence: daughters fledged with higher baseline levels of corticosterone than sons and (2) differential costs of rearing sons versus daughters: mothers rearing sons when environmental conditions were poor completed parental care in poorer condition than mothers rearing daughters in the same year and mothers rearing either sex when conditions were better. These novel results may help to explain the disparate results of previous studies of avian sex allocation.     

The role of migration and founder effect for the evolution of cooperation in a multilevel selection context

The idea that natural selection can be meaningfully applied at the group level may be more important than previously thought. This perspective, a modern version of group selection, is called multilevel selection. Multilevel selection theory could incorporate previous explanations for the evolution of cooperation including kin selection. There is general agreement that natural selection favors noncooperators over cooperators in the case of an unstructured population. Therefore, the evolution of cooperation by multilevel selection often requires positive assortment between cooperators and noncooperators. The question is how this positive assortment can arise in the ecological meaning. We constructed an individual-based model of multilevel selection and introduced migration and evolution. The results showed that positive assortment was generated especially when a migration strategy was adopted in which individuals respond specifically to bad environmental conditions. It was also shown that the founder effect in the evolutionary process could further facilitate positive assortment by working with migration. We analyzed assortment by using relatedness defined in group-structured populations. The fact that cooperation was achieved by such migration and by the founder effect highlights the importance of sensitiveness to the ecological environment and of fluctuations in group size, respectively.     

Cooperative breeding and immunity: a comparative study of PHA response in African birds

Cooperatively breeding birds might be expected to suffer from higher costs of parasitism than pair-breeding species because of two aspects of their ecology which should facilitate horizontal transmission and possibly select for higher parasite virulence: first, they interact regularly with more individuals than pair-breeding species, and second, these individuals are commonly close relatives that could share similar resistance alleles. This hypothesis predicts that cooperative breeders should invest relatively more in immune defence than closely related species which breed in pairs. I tested this prediction comparatively in African birds by examining the response of the immune system to the mitogenic lectin, phytoahemagglutinin (PHA response) in relation to cooperative breeding. Among 66 species, of which 18 breed cooperatively, PHA response was significantly higher in cooperatively breeding species. This association appeared not to be confounded by body size, clutch size, nest position, coloniality or similarity owing to common phylogenetic descent. These results suggest that cooperatively breeding birds may have been selected to invest more than pair-breeders in defences against parasites. If so, then additional costs of philopatry and helping behaviour might be imposed on breeders, helpers and offspring.     

Vicarious sampling: the use of personal and public information by starlings foraging in a simple patchy environment

Group foragers may be able to assess patch quality more efficiently by paying attention to the sampling activities of conspecifics foraging in the same patch. In a previous field experiment, we showed that starlings foraging on patches of hidden food could use the successful foraging activities of others to help them assess patch quality. In order to determine whether a starling could also use another individual’s lack of foraging success to assess and depart from empty patches more quickly, we carried out two experimental studies which compared the behaviour of captive starlings sampling artificial patches both when alone and when in pairs. Solitary starlings were first trained to assess patch quality in our experimental two-patch system, and were then tested on an empty patch both alone and with two types of partner bird. One partner sampled very few holes and thus provided a low amount of public information; the other sampled numerous holes and thus provided a high amount of public information. In experiment 1, we found no evidence of vicarious sampling. Subjects sampled a similar number of empty holes when alone as when with the low and high information partners; thus they continued to rely on their own personal information to make their patch departure decisions. In experiment 2, we modified the experimental patches, increasing the ease with which a bird could watch another’s sampling activities, and increasing the difficulty of acquiring accurate personal sampling information. This time, subjects apparently did use public information, sampling fewer empty holes before departure when with the high-information partner than when with the low-information partner, and sampling fewer holes when with the low-information partner than when alone. We suggest that the degree to which personal and public information are used is likely to depend both on a forager’s ability to remember where it has already sampled and on the type of environment in which foraging takes place. Received: 31 January 1995/Accepted after revision: 11 September 1995     

Relationships between nutrients and macroalgal biomass in a brazilian coastal lagoon: the impact of a lock construction

The Piratininga Lagoon is a coastal, choked, and brackish ecosystem in SE Brazil, where uncontrolled discharge of domestic sewage led to eutrophication, increasing massive proliferation of benthic macroalgae, and decrease of the lagoon mean depth. In 1992, a dam was constructed by the local Municipality at its tidal channel aiming at stabilising its water level. Main physico-chemical parameters, together with macroalgae biomass, nutrient concentrations in the water column, particulate organic carbon, and chlorophyll a were recorded monthly at 4 sampling stations within the Piratininga lagoon from April 1994 to April 1995. The data, compared with “before-lock” existing studies, show that nutrients and chlorophyll a concentrations significantly increased after the lock construction. Based on the functioning of the ecosystem, we propose to harvest the algal mats before their decomposition period in order to partly remove the nutrient stocks from the lagoon and the future ecosystem modelling to predict the impact of natural and anthropogenic eutrophication.     

Seasonal and spatial character of PCBs in a chemical industrial zone of Shanghai,China

As one of China’s great metropolises, Shanghai is suffering from the impact of manufacture and the use of chemical industrial products, and it faces serious pollution from polychlorinated biphenyls (PCBs). Therefore, in this study, in order to assess the seasonal and spatial character of contamination from chemical industrial zones, the concentrations of PCBs have been measured in various environmental media, including soil, leaves, and atmospheric particulate samples collected in a chemical industrial zone of Shanghai and compared with samples from presumably unpolluted sites of rural areas. In soils, the PCB concentrations ranged from 0.5 ng g⁻¹ (unpolluted site) to 586.85 ng g⁻¹ (chlor-alkali industry site). The concentrations of PCBs in evergreen leaves ranged from 0.3 ng g⁻¹ to 32.46 ng g⁻¹, and more chlorinated biphenyls congeners, such as penta-biphenyls and hexa-biphenyls, were the dominant contributors in winter and spring. Seasonal differences and the constitution patterns of congeners might be affected by the temperature and industrial activities. The PCB concentrations in the leaves of deciduous trees increase over time as the leaves grow. The PCB concentration in atmospheric particulates was in the range of 9.22–14.15 × 10³ pg m⁻³, which might be the result of influence from climate and industrial activities. The relativity of PCB contents among the environmental media was discussed. The results in this paper provide an important profile of the current contamination status of a key chemical industrial zone in China.     

The sensitivity of carbon sequestration to harvesting and climate conditions in a temperate cypress forest: Observations and modeling

The role of disturbance and climate factors in determining the forest carbon balance was investigated at a Japanese cypress forest in central Japan with eddy flux measurements, tree-ring analyses, and a terrestrial biosphere model. The forest was established as a plantation after intermittent harvesting and replanting between 1959 and 1977, and acted as a strong carbon sink of approximately 500 g C m⁻² year⁻¹ for the measurement years between 2001 and 2007. A terrestrial biosphere model, BIOME-BGC, was validated using the eddy flux data at daily to interannual timescales, and the tree-ring width data at interannual to decadal timescales. According to the model simulation, during the observation period 270 ± 55 g C m⁻² year⁻¹ was additionally sequestered due to the indirect effects of the harvesting and planting, whereas the increase of CO₂ concentration and the change in climate increased the sink of 110 ± 40 and 30 ± 80 g C m⁻² year⁻¹, respectively. The model simulation shows that the forest is now recovering from harvesting, and that harvesting is a more important determinant of the current carbon sink than either interannual climate anomalies or increased atmospheric CO₂ concentration. We found that harvesting with long rotation length could be effective management activity in order to increase carbon sequestration, if the harvested timber is converted into products with long lifecycles.     

地形因子对暖温带森林群落物种丰富度-地上生物量关系的影响

地形因子是森林群落物种丰富度与生产力的重要影响因素,可以修正物种丰富度-生产力两者之间的关系。为了厘清地形因子对物种丰富度-生产力关系的修正效应,以东灵山暖温带天然次生林为研究对象,基于20 hm2样地DEM地图,提取海拔、坡度、坡向、凹凸度等地形因子,用GLM模型探讨了地形因子对群落丰富度与地上生物量的影响。结果表明,群落地上生物量与物种丰富度分别受不同地形因子的影响,群落地上生物量受Ea、El2、El3、Co34个因子的显著影响（P〈0.05）,而物种丰富度则受Sl、El、El2、El34个因子的显著作用（P〈0.05）,两者受海拔相关因子的影响最大,分别解释了两者3.99%和11.09%的变异。地形因子修正了物种丰富度-地上生物量之间的单峰曲线关系,使其朝着单调与直线化趋势变化。以后在探讨森林群落物种丰富度-生产力关系时,地形因子的修正作用应该受到重视。     

Parasitism in a social wasp: effect of gregarines on foraging behavior, colony productivity, and adult mortality

Behavior in eusocial insects likely reflects a long history of selection imposed by parasites and pathogens because the conditions of group living often favor the transmission of infection among nestmates. Yet, relatively few studies have quantified the effects of parasites on both the level of individual colony members and of colony success, making it difficult to assess the relative importance of different parasites to the behavioral ecology of their social insect hosts. Colonies of Polybia occidentalis, a Neotropical social wasp, are commonly infected by gregarines (Phylum Apicomplexa; Order Eugregarinida) during the wet season in Guanacaste, Costa Rica. To determine the effect of gregarine infection on individual workers in P. occidentalis, we measured foraging rates of marked wasps from colonies comprising both infected and uninfected individuals. To assess the effect of gregarines on colony success, we measured productivity and adult mortality rates in colonies with different levels of infection prevalence (proportion of adults infected). Foraging rates in marked individuals were negatively correlated with the intensity of gregarine infection. Infected colonies with high gregarine prevalence constructed nests with fewer brood cells per capita, produced less brood biomass per capita, and, surprisingly, experienced lower adult mortality rates than did uninfected or lightly infected colonies. These data strongly suggest that gregarine infection lowers foraging rates, thus reducing risk to foragers and, consequently, reducing adult mortality rates, while at the same time lowering per-capita input of materials and colony productivity. In infected colonies, queen populations were infected with a lower prevalence than were workers. Intra-colony infection prevalence decreased dramatically in the P. occidentalis population during the wet season.An erratum to this article can be found at     

Achieving environmental goals in a world of trade and hidden action: the role of trade policies and eco-labeling

Importing countries often resort to trade policies and consumer actions in order to reduce negative environmental impacts of the products they consume. Traditionally, these policies were non-discriminatory, i.e., they treated all imports equally, without considering the actual damages caused by the product. More recently, there is a trend towards process-discriminatory policies which attempt to discriminate against environmentally unsound imports while encouraging sound alternatives. We develop a theoretical model of the consuming country's optimal trade policy, allowing for asymmetric information and costly monitoring, and analyze what type of policy is preferable under what conditions.     

温带阔叶红松林中不同树种和倒木对土壤性质的影响

以长白山温带阔叶红松林为研究对象,通过野外采集土壤样品,综合研究老龄树及倒木对周边表层土壤性质的影响。研究结果显示,在本研究样地条件下,不同树种周边的表层土壤性质存在显著性差异,红松（Pinus koraiensis）周边土壤中水溶性有机碳WSOC的质量分数和特征吸光系数SUVA254值都高于紫椴（乃砌amurensis）和水曲柳（Fraxinus mandshurica）,但是土壤水溶性总氮WSTN和交换性无机氮的质量分数却是红松低于紫椴和水曲柳。对倒木影响的森林表层土壤性质的研究结果显示,倒木下方森林表层土壤中WSOC、WSTN以及交换性矿质氮的质量分数都明显地高于林间土壤,说明在小尺度空间区域内,倒木可能是森林表层土壤活性碳、氮库的重要输入源。此外,本研究还利用主成分分析探讨了受树种影响的森林表层土壤不同性质之间的关联性,并对树种和倒木影响周边土壤性质的可能性机制进行分析。