The cost of host egg damage caused by a brood parasite: experiments on great spotted cuckoos (Clamator glandarius) and magpies (Pica pica)

Adult great spotted cuckoos, Clamator glandarius, frequently damage one or more eggs of their magpie host, Pica pica, without removing or eating them. The presence of damaged host eggs could signal parasitism thereby increasing the probability that the parasitic egg is ejected. This hypothesis was tested by experimentally introducing a model cuckoo egg with or without damaged host eggs. Magpie responses to experimental parasitism did not differ significantly between treatments implying that damaged host eggs are not used by magpies to assess parasitism. We followed the fate of magpie eggs naturally damaged by the great spotted cuckoo or experimentally damaged by us. Host response was very similar for naturally or experimentally damaged host eggs, but varied significantly according to the type of egg damage, eggs being removed more frequently when pecked than crushed, while cracked eggs were never removed. However, the egg damage that most readily causes egg removal is albumen leakage. Received: 30 November 1998 / Received in revised form: 7 June 1999 / Accepted: 12 June 1999     

Chick recognition and acceptance: a weakness in magpies exploited by the parasitic great spotted cuckoo

Hosts of brood parasites have evolved the ability to discriminate non-mimetic and even mimetic eggs, but not non-mimetic chicks. Here we demonstrate that the great spotted cuckoo Clamator glandarius does not provide its magpie Pica pica host with a super-normal stimulus that helps to avoid recognition, because single cuckoo chicks introduced into otherwise unparasitized magpie nests are not fed at a higher frequency than single magpie chicks introduced to parasitized magpie nests. Another series of experiments demonstrated that magpies have the ability to discriminate cuckoo chicks, mainly when these are introduced at the end of the nestling period, and especially when the cuckoo chick together with a magpie chick is presented to adult magpies outside the nest. This supports the idea that cuckoos exploit the obligatory reaction of magpies to feed all young that have been hatched in their nests and whose signatures they have learnt. Furthermore, the experimental cuckoo chicks in parasitized magpie nests were more likely to be accepted than they were in non-parasitized nests. This supports the hypothesis that magpies may learn to recognise their own nestlings as those present in the nest and may indicate that a comparison between cuckoo and magpie nestlings is the basis of discrimination.     

Preferential allocation of food by magpies Pica pica to great spotted cuckoo Clamator glandarius chicks

Adult magpies Pica pica provide parasitic great spotted cuckoo Clamator glandarius nestlings with a diet very similar to that fed to their own chicks. In both naturally and experimentally parasitized nests, great spotted cuckoo chicks were fed at a higher rate than magpie chicks in the same nest. This preferential allocation of food by magpie parents to great spotted cuckoo chicks is consistent with the supernormal stimulus hypothesis, because this result implies that cuckoo chicks provide stronger stimuli for parental care than host chicks. Great spotted cuckoo chicks receive most of the food brought to the nest by the foster parents, because they exploit a series of stimuli which jointly (or sometimes individually) operate as a supernormal stimulus. This hypothesis predicts that if any stimulus is masked, the efficiency of the cuckoo in eliciting parental care will decrease. Here, we analyze experimentally the effects of two of these stimuli, preferential feeding of large nestlings and of nestlings with conspicuous palatal papillae. Firstly, when we experimentally introduced one medium-sized (7–9 days) cuckoo chick into an unparasitized magpie nest where the largest magpie chick was 12–15 days old, the cuckoo did not receive significantly more food than the average or the largest magpie chick. Secondly, when unparasitized nests were experimentally parasitized with a cuckoo chick that had its gape painted to mimic that of magpie chicks, the parasitic cuckoo received less food than the average magpie chick.     

Predation risk and the territory value of cover: an experimental study

In an experiment on territorial brown trout (Salmo trutta), we addressed the novel hypothesis that protective cover increases the value of a territory in relation to the perceived level of predation risk. We predicted that territory holders should invest more resources defending territories with cover than territories without cover, and that defence should increase as predation risk increases. First, trout were allowed to establish ownership in territories with or without overhead cover. Second, predation risk was manipulated by simulating aerial predator attacks in half of the territories of each type, whereupon the preference for cover was estimated. Third, owners of the four types of territories were staged in dyadic contests against size-matched intruders. Territory owners showed a preference for cover, which increased further after simulated predator attacks. In subsequent contests, conflicts over territories with cover were settled faster than conflicts over territories without cover, which may suggest that the value of cover increases the motivational asymmetry between owner and intruder. Consistent with our hypothesis, owners of territories with cover were much more aggressive if they had been subjected to predator attacks the day before the conflict. These results suggest that territory owners are able to estimate the value of protective cover in response to variation in the level of predation risk in the habitat.Communicated by J. Krause     

Developing an empirical model of phytoplankton primary production: a neural network case study

We describe the development of a neural network model for estimating primary production of phytoplankton. Data from an enriched estuary in the eastern United States, Chesapeake Bay, were used to train, validate and test the model. Two error backpropagation multilayer perceptrons were trained: a simpler one (3-5-1) and a more complex one (12-5-1). Both neural networks outperformed conventional empirical models, even though only the latter, which exploits a larger suite of predictive variables, provided truly accurate outputs. The application of this neural network model is thoroughly discussed and the results of a sensitivity analysis are also presented.     

Competitor assessment by scent matching: an experimental test

Summary If males assess potential opponents by comparing their odour with that of scent marks in the vicinity (the so called scent matching hypothesis) they should be more reluctant to fight when these odours are the same. Under these circumstances the potential opponent is thus identified as a resource holder (a male monopolising a territory or group of females) that would escalate encounters to defend its resources. We tested this prediction by comparing the fighting behaviour of male house mice, Mus domesticus, when on a scent-marked substrate that matched the odour of an opponent with their behaviour on a substrate that did not match. As predicted, the delay before the first fight was longer and the rate of subsequent fighting lower when the scents matched. This result is consistent with the interpretation that resource holders scent mark to provide opponents with a means of competitor assessment and that they thus reduce the costs of resource defence. Offprint requests to: L.M. Gosling     

Simulation of inclined dense jets in stagnant environments: an LES and experimental study

Environmental Fluid Mechanics - Inclined dense jets are commonly used to mitigate the environmental impacts of brine discharge in coastal desalination plants. Numerous studies have been performed...     

Siblicide in Serengeti spotted hyenas: a long-term study of maternal input and cub survival

In the Serengeti National Park, Tanzania, large fluctuations of prey abundance alters the frequency at which spotted hyena (Crocuta crocuta) cubs are nursed and thus the total level of maternal input available to them. Maternal input is high when mothers feed on high densities of locally available migratory herbivores and low when mothers travel up to 70 km to forage. Using data from 19 cub cohorts on the incidence of siblicide (from monitoring the survival of 609 cubs in twin litters) and cub growth rates (from 195 cubs in twin litters) as a measure of maternal input, we demonstrate that the incidence of siblicide increased as average cohort growth rate declined. In total, there were 37 siblicides in 384 litters (9% of litters). When both cubs were alive, total maternal input in siblicidal litters was significantly lower than in non-siblicidal litters and the mean share of the dominant sib of 64.6% was significantly higher than the mean of 52.1% for dominant sibs of non-siblicidal litters. After siblicide, growth rates of siblicide victors substantially increased, demonstrating that mothers did not reduce maternal input after litter reduction. As a result, siblicide victors achieved a long-term growth rate similar to that of singletons and thus significantly increased their expected survival. We conclude that in spotted hyenas, high maternal input in lactation has favoured the evolution of facultative siblicide in populations inhabiting areas with low or fluctuating food resources.     

Estimating the number of active and successful bald eagle nests:an application of the dual frame method

Monitoring the number of animals in a wildlife population is an important task. In this paper, we estimate the number of active and successful bald eagle nests in a specific region using dual frame sampling techniques. The dual frame method combines independent samples from an incomplete list frame and an area frame that is assumed to be complete. Hartley's screening estimator is used to combine sample information from both frames. This methodology will be useful for monitoring any wildlife population where the breeding individuals have highly visible nesting territories which tend to be stable over many years.     

Courtship display by great frigatebirds, Fregata minor: an energetically costly handicap signal?

We used detailed time-budget observations, mark-resight data, and doubly labeled water estimates of energy expenditure to test whether energy spent on courtship display by male frigatebirds functions as a sexually selected handicap signal. During a 2-day period of time budget observations, males displayed on average 29.5% of the time (range 0–100% for 15 different males), and this value was correlated with an index of season-long display effort. Season-long display effort was strongly predictive of pairing success. Average field metabolic rate (FMR) during the 2-day time budget period was 676.5 kJ/day (range 464.8–1035.0), substantially lower than the mass-specific FMR predicted from studies of other seabirds during incubation or chick-rearing. Despite a low overall FMR, display effort could function as an energetic handicap, either if FMR correlates positively with the amount of courtship performed or if high-quality males display at a high rate because they pay a lower energy cost per unit of display than do low-quality males. We found no relationship between FMR and display rate, despite good power for doing so. We also did not find a significant difference in energy spent divided by courtship time for males that were or were not chosen by females (inferred to be high- and low-quality males, respectively), though the medians differed in the predicted direction (preferred males having lower relative costs than non-preferred males) and the confidence interval on the difference between groups was very wide. Thus, we found evidence that male courtship effort is predictive of pairing success, we rejected one mechanism by which energetic cost of display could function as a handicap, and our test of the alternate energetic handicap mechanism was equivocal.     

Contaminants in drinking water and its mitigation using suitable adsorbents: an overview

Various options are applicable for the removal of water pollutants included reverse osmosis, ion exchange, coagulation, co-precipitation, catalytic reduction, herbal filtration, electrodialysis and adsorption. This paper deals with the sorption phenomena for the removal of pollutants from drinking water. Attempts have been made to use low cost sorbents developed by pretreatment/activation/impregnation with alkalis, acids, iron oxide, manganese dioxide, ferric chloride, alum, lime, aluminum salts with natural products/indigenous minerals viz. activated alumina, activated carbon, groundnut husk, saw dust, chemically coated sand, fly ash, zeolites, clay minerals and other plant products. Application of Freundich and Langmuir isotherms were used to assess the adsorption capacity. Equilibrium isotherms were determined at optimum temperature and pH to characterize the sorption process. Statistical parameters such as mass transfer coefficients, multiple regression analysis were applied to establish the mechanism. It is suggested that the characterization of suitable, and exhausted sorbent through the application of fourier transformed infrared spectroscopy (FTIR), X-ray diffraction (XRD), X-ray fluorescence (XRF) is essential to establish its surface bonding. Scope for safety evaluation and risk assessment to human and biosphere may provide the guideline and predication to the regulatory agencies for its sustainable use and safe disposal The ecotoxicological assessment of the leachates and low cost removal technology are discussed in this paper.     

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

Economic growth,public welfare and sustainability: an empirical system analysis

SUMMARY

This paper explores the relationships between economic activity and sustainability, growth, and welfare. The issues regarding the nature of the relationships were framed by Ekins in questions which are addressed empirically across countries and the states of the US. The sustainability issue is addressed by comparing nonrenewable energy throughput, a surrogate for impacts, to the renewable energy throughput in the associated ecosystem. The effect of a changing energy intensity, in developing and developed countries, on socioeconomic variables provides clues to social welfare. In developing countries a rising energy throughput (economic growth) and energy intensity is positively related to public welfare while impacts are generally moderate. The reverse appears to be true in developed countries, i.e. as energy throughput and intensity rise in developed countries and the US states, pollution and impacts also rise while socioeconomic variables worsen, so public welfare begins to decline. Economic development appears to pass through three phases which can be characterized by the nature and extent of its energy use. Each phase exhibits characteristic energy, pollution, structural and socioeconomic trends.     

Fifteen-spined stickleback (Spinachia spinachia) females prefer males with more secretional threads in their nests: an honest-condition display by males

In the fifteen-spined stickleback (Spinachia spinachia), the male alone builds a nest and provides care for the eggs until they hatch. The nest is made of filamentous algae held together with shiny secretional threads of a glycoprotein, here called tangspiggin. Tangspiggin is produced by transformed kidney cells. I investigated how tangspiggin production was related to food intake by giving males low or high food rations. Males in the high-food group produced significantly more tangspiggin than those in the low-food group. Since low food rations did not lead to a significant drop in resting metabolic rate (oxygen consumption), tangspiggin production appears to be one of the first energy-demanding processes to be dropped during food deprivation. Among males given equal food rations, females preferred to mate with those with elevated levels of tangspiggin in their nests. Tangspiggin threads may signal safety for the offspring by keeping the eggs within the nest and protecting them from egg predators. Moreover, the threads become less shiny with age, due to particle deposition, and older nests that may house older (more valuable) eggs are thereby well camouflaged. Tangspiggin provides an honest signal of male condition, acting as an extra-bodily ornament, signaling direct benefits to females or possibly also indirect benefits of "good genes" from a male who is a successful forager.     

Short- and long-term consequences of prenatal testosterone for immune function: an experimental study in the zebra finch

Hormone-mediated maternal effects play an important role in the formation of a differentiated phenotype. They have been shown to influence a wide array of offspring traits, both early in life and in adulthood. One important offspring trait that is under the influence of maternal androgens is the immune system. In birds, a growing number of studies show that yolk androgens modulate immune function during the chick stage. However, there is a lack of knowledge regarding long-term effects of prenatal androgens on offspring immunity. In this study, we therefore investigated the influence of prenatal testosterone (T) on several measures of immunity in fledgling and adult zebra finches (Taeniopygia guttata). Cell-mediated immune response (towards phytohaemagglutinin, PHA) of fledglings hatching from control eggs was negatively related to brood size, whereas there was no such association for fledglings hatching from eggs with experimentally elevated T levels (T fledglings). Male control fledglings showed reduced mass gain compared to female control fledglings within 24 h after the PHA injection. This pattern was reversed in T fledglings. Total antibody levels in fledglings were not affected by egg treatment. Neither cell-mediated immunity nor total antibody levels in sexually mature zebra finches were influenced by egg treatment. However, there was an immuno-enhancing effect of elevated egg T on both primary and secondary humoral immune responses toward diphtheria and tetanus antigens in ca 5 and 7 month old zebra finches. In addition, the covariation between different immune components differed between T and control offspring, suggesting that egg treatment may have altered the potential trade-offs between different parts of the immune system. Our results suggest that prenatal androgens could be an important factor contributing to individual variation in immune function even in adulthood.     

Permanent exposure versus facultative concealment of sexual traits: an experimental study in the house sparrow

In many bird species the cryptic winter plumage is due to the presence of light feather tips that conceal conspicuous colorations. The gradual abrasion of these tips that makes sexual traits visible has been interpreted as a strategy to improve mating success (here referred as the permanent exhibition hypothesis). However, under some circumstances, the maintenance of a full plumage that facultatively enables the bird to exhibit or cover aggression-inducing traits has proved to be advantageous (the coverable badge hypothesis). In a population of house sparrows where black throat patches (here called badges) are used in intrasexual competition, the degree of abrasion of dull feather tips that conceal bright colour early in the breeding season correlated neither with badge size nor with traits indicating morphology and body condition. These results are only in accordance with predictions of the coverable badge hypothesis. In 1992 experimental clipping of badge feather tips, which forced permanent exposure of badges, negatively affected birds with the largest badges in terms of nest acquisition, but an opposite trend was indicated for birds with the smallest badges. In 1993, when more novel birds in the study area were competing for nesting sites than in 1992, only badge size, but not the experimental manipulation, affected nesting success. These results suggest that the loss of the ability to conceal badges was disadvantageous, and more so if most competing individuals had already been resident in the colony in previous years and the larger their badges were. There is a striking contrast between the results reported for this study population, where badges mainly signal fighting ability, and those reported for another house sparrow population, where badges are mainly used in mate choice. In this last population, tip abrasion is advantageous for dominant individuals showing the largest badges. This suggests that the trade-off between conserving versus wearing off the feathers that conceal ornaments could have different optimal resolutions depending on the relative importance of inter and intrasexual selection on indicator traits. Received: 29 September 1995/Accepted after revision: 14 July 1996     

Protandry in western cicada killer wasps, (Sphecius grandis,Hymenoptera: Sphecidae): an empirical study of emergence time and mating opportunity

Summary Sphecius grandis is a univoltine, colonial wasp. Females mate once and are sexually receptive when they emerge in July and early August. Males generally emerge earlier in the summer than females. The opportunity for each male to acquire mates is a function of the number of females emerging during his lifetime and the number of competitiors that are active when the females emerge. I determined a mating opportunity index (MOI) for each male in an aggregation of wasps for three separate summers, and correlated the MOI of individual males with their emergence date. The MOI of a male estimates the potential contribution that timing of emergence makes to his reproductive success. In 1984 males emerging near the mean of the male emergence distribution had the highest MOI. These males emerged between one and two weeks prior to the mean female emergence date. However, in 1981 late emerging males had the highest MOI. In 1983 there was no significant difference in MOI among males. As a result of between-year variation in female emergence schedules and in the duration of male lifetime, the selection pressures influencing male emergence time vary between years.     

Resource partitioning among savanna grazers mediated by local heterogeneity: an experimental approach

Recent theoretical studies predict that body size-related interspecific differences in spatial scale of perception and resource use may contribute to coexistence of species that compete for the same class of resources. These studies provide a new theoretical framework for explaining resource partitioning patterns among African ungulates that coexist in spatially heterogeneous savanna grasslands. According to these studies, different-sized ungulates can coexist because larger species forage at a coarser scale but can tolerate lower quality food, whereas smaller species need higher quality food but forage at a finer scale. To test this hypothesis in an African savanna, we created an experimental mosaic with variation in grain (spatial detail) and quality of short-grass patches and directly observed the visitation of naturally occurring grazers to this mosaic over a two-year period (total of 903 observation hours). Of the seven species that visited our experiment, warthog, impala, zebra, and white rhino visited long enough to allow data analysis. We showed that warthog and impala avoided plots with a finer grain of short grass and that warthog preferred fertilized plots to unfertilized plots. Zebra and white rhino did not avoid the finer grain plots. Our results suggest that differences in grain and quality of a resource might indeed contribute to partitioning of this resource by savanna ungulates. Although four focal species is unusually high for an experimental study on resource partitioning among naturally occurring savanna ungulates, this number is too low to evaluate the allometric basis of our hypothesis. Our results, however, encourage wider experimental testing of the role of spatial heterogeneity in facilitating the coexistence of potentially competing savanna herbivores.