Minimizing predation risk in a landscape of multiple predators: effects on the spatial distribution of African ungulates

Studies that focus on single predator-prey interactions can be inadequate for understanding antipredator responses in multi-predator systems. Yet there is still a general lack of information about the strategies of prey to minimize predation risk from multiple predators at the landscape level. Here we examined the distribution of seven African ungulate species in the fenced Karongwe Game Reserve (KGR), South Africa, as a function of predation risk from all large carnivore species (lion, leopard, cheetah, African wild dog, and spotted hyena). Using observed kill data, we generated ungulate-specific predictions of relative predation risk and of riskiness of habitats. To determine how ungulates minimize predation risk at the landscape level, we explicitly tested five hypotheses consisting of strategies that reduce the probability of encountering predators, and the probability of being killed. All ungulate species avoided risky habitats, and most selected safer habitats, thus reducing their probability of being killed. To reduce the probability of encountering predators, most of the smaller prey species (impala, warthog, waterbuck, kudu) avoided the space use of all predators, while the larger species (wildebeest, zebra, giraffe) only avoided areas where lion and leopard space use were high. The strength of avoidance for the space use of predators generally did not correspond to the relative predation threat from those predators. Instead, ungulates used a simpler behavioral rule of avoiding the activity areas of sit-and-pursue predators (lion and leopard), but not those of cursorial predators (cheetah and African wild dog). In general, selection and avoidance of habitats was stronger than avoidance of the predator activity areas. We expect similar decision rules to drive the distribution pattern of ungulates in other African savannas and in other multi-predator systems, especially where predators differ in their hunting modes.     

Understanding determinants of home range behaviour of feral cats as introduced apex predators in insular ecosystems: a spatial approach

The introduced feral cat (Felis catus) is a widespread generalist with flexible social behaviour and an apex predator without major interspecific competitors in insular ecosystems that evolved in the absence of predators. Mechanistic definitions consider an animal’s home range to be the spatial expression of a cognitive map that is kept up-to-date with the status of critical resources that contribute to animal fitness. We assumed there are two major determinants structuring the home range of cats as apex predators in insular ecosystems: the distribution of critical food resources and conspecific distribution. We hypothesized that cats structure their home ranges by optimizing the use of staple critical food resources and that as a consequence of the presence of rich resources cats tend to socialise, aggregate and share space. We carried out spatial analyses using location data for feral cats tracked using lightweight GPS collars in conjunction with the suitability value of rabbit patches and their associated ownership costs for cats within a New Zealand braided-river environment. Cat home ranges and spatial distribution, especially for females, were related to the inclusion of rabbit patches within home ranges with higher mean value than the average of neighbourhood patches in the landscape. Cats showed solitary behaviour but tolerance to conspecific presence by sharing high-use areas and high-value rabbit patches, mostly at different times, resulting in occasional encounters among males and females. Home range size and patterns of spatial overlap were dependant on sex and season. Solitary spacing patterns as consequence of innate preferences together with resource constraints may regulate feral cat population densities.     

Patterns of fluctuating asymmetry in beetle horns: an experimental examination of the honest signalling hypothesis

Recent theoretical arguments have claimed that negative relationships between the size and symmetry of secondary sexual traits are indicative of honest signalling of male quality. The patterns of fluctuating asymmetry in beetle horns have been proposed to support the honest signalling hypothesis. Here we examine three assumptions of the hypothesis, (1) that traits are costly to produce; (2) the levels of fluctuating asymmetry are indicative of stress imposed during development; and (3) that males with larger traits should have more symmetrical traits, using the horned beetle, Onthophagus taurus. Experimental manipulations of brood mass were used to manipulate horn size and asymmetry. The development of horns was found to be environmentally determined and costly in terms of delayed development and increased risk of pre-adult mortality. Decreasing resource availability increased relative horn asymmetry. However, horn height was positively related to absolute horn asymmetry. While the results do support the hypothesis that sexual selection on secondary sexual traits should increase levels of fluctuating asymmetry, they provide no support for the notion that the patterns of asymmetry honestly signal male quality. Horns are used in disputes between males and may be indicative of male parental investment. Thus, we conclude that while horn size may be an indication of male quality, the patterns of fluctuating asymmetry are not. Received: 16 December 1996 / Accepted after revision: 5 May 1997     

The Baltic Sea spring phytoplankton bloom in a changing climate: an experimental approach

The response of the Baltic Sea spring bloom was studied in mesocosm experiments, where temperatures were elevated up to 6°C above the present-day sea surface temperature of the spring bloom season. Four of the seven experiments were carried out at different light levels (32–202?Wh?m^?2 at the start of the experiments) in the different experimental years. In one further experiment, the factors light and temperature were crossed, and in one experiment, the factors density of overwintering zooplankton and temperature were crossed. Overall, there was a slight temporal acceleration of the phytoplankton spring bloom, a decline of peak biomass and a decline of mean cell size with warming. The temperature influence on phytoplankton bloom timing, biomass and size structure was qualitatively highly robust across experiments. The dependence of timing, biomass, and size structure on initial conditions was tested by multiple regression analysis of the y-temperature regressions with the candidate independent variables initial light, initial phytoplankton biomass, initial microzooplankton biomass, and initial mesozooplankton (=copepod) biomass. The bloom timing predicted for mean temperatures (5.28°C) depended on light. The peak biomass showed a strong positive dependence on light and a weaker negative dependence on initial copepod density. Mean phytoplankton cell size predicted for the mean temperature responded positively to light and negatively to copepod density. The anticipated mismatch between phytoplankton supply and food demand by newly hatched copepod nauplii occurred only under the combination of low light and warm temperatures. The analysis presented here confirms earlier conclusions about temperature responses that are based on subsets of our experimental series. However, only the comprehensive analysis across all experiments highlights the importance of the factor light.     

Potential allelopathic effect of Pinus halepensis in the secondary succession: an experimental approach

Summary. Recent economic and social changes in north Mediterranean regions have led to an important rural depopulation. Consequently, meadows developed on abandoned agricultural lands (characterized by high species richness) undergo reforestation. These former fields are mainly colonized by Pinus halepensis Miller, which is known to synthetize a wide range of secondary metabolites, among these, some could influence plant succession through allelopathy. The allelopathic potential of P. halepensis, was tested against two target species (Lactuca sativa L. and Linum strictum L.) with aqueous extracts obtained from different organs (root and needle) taking into account the individual age (±10, ±30 and > 60 years old). Root and needle extracts affected differently germination and growth of the two target species, the responses varying with concentration of extracts, age and organs tested. The strongest inhibitory effect was observed on the germination and growth of L. strictum, exposed to needle extracts of young P. halepensis (±10 years old), and root extracts of older P. halepensis (> 30 years old). These extracts contained several phenolic acids (e.g. 4-hydroxybenzoic acid and p-coumaric acid), which are known as allelochemicals and their concentrations vary with age and organ tested. Hence, P. halepensis could influence secondary succession through the release of potential allelochemicals in the environment by leaf leachates or root exudates.     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

Evaluating the relative importance of intrinsic and extrinsic factors on the foraging activity of top predators: a case study on female little penguins

A complex interaction of biotic and abiotic factors influences animal foraging activity. It is often difficult to understand which factors may affect animals’ foraging and how it is affected. For instance, whereas the effect of sexual dimorphism on foraging activity has been reported in several species, little is known of the complex interactions between variables acting at a finer scale, e.g. the variability of body mass within a sex. Evaluating the importance of these finer scale factors is also essential to the understanding of foraging behaviour. We propose here a simple approach by applying principal component analysis (PCA) in a novel way to examine relationships between biotic and abiotic factors affecting foraging behaviour of top predators. We studied female little penguins (Eudyptula minor) of known age, carrying miniature accelerometers during the guard stage of breeding. Surprisingly, the body mass of the females did not influence any of the foraging parameters, but females foraging later in the breeding season dived shallower and more often, showing a strong correlation with laying date. Similarly, the diving effort of females was greater with increasing chick age within the same breeding stage. These results indicate that for female little penguin, the relationship between changes in prey availability and hunting effort can change at a fine scale, within a breeding stage. Therefore, any analysis of little penguin foraging behaviour during breeding should consider the timing in relation to the breeding season. We encourage researchers to develop the use of this PCA approach as it could help clarify the complexity of the underlying mechanisms determining foraging activity and we propose that it should be used as a first step of foraging behaviour analysis, before examining a particular relationship.     

Influence of the mineralogical composition on microbial activities in marine sediments: an experimental approach

We investigated the influence of the mineralogical composition of marine sediments on bacterial activity in experimental microcosms. Calcite and quartz were added to natural marine sediments and microbial response in terms of total bacterial abundance and biomass, β-D-glucosidase exo-enzymatic activity and bacterial incorporation of a radio-labelled (³H-leucine) substrate were investigated for a period of one month. We report here that after 15 days the mineralogical composition of the sediment (calcite vs. quartz) had an impact on bacterial abundance and activity (reduced for ca 15% and 56%, respectively). However, such impact was mitigated or even disappeared in high organic nutrient conditions.     

Influence of the mineralogical composition on microbial activities in marine sediments: an experimental approach

We investigated the influence of the mineralogical composition of marine sediments on bacterial activity in experimental microcosms. Calcite and quartz were added to natural marine sediments and microbial response in terms of total bacterial abundance and biomass, β-D-glucosidase exo-enzymatic activity and bacterial incorporation of a radio-labelled (³H-leucine) substrate were investigated for a period of one month. We report here that after 15 days the mineralogical composition of the sediment (calcite vs. quartz) had an impact on bacterial abundance and activity (reduced for ca 15% and 56%, respectively). However, such impact was mitigated or even disappeared in high organic nutrient conditions.     

Revisiting the carbon emissions hypothesis in the developing and developed countries: a new panel cointegration approach

Environmental and Ecological Statistics - Since global warming worsens with economic development and emitted CO2 is one of the main greenhouse gases, it is important to understand...     

Application of polysaccharide-based biopolymers as supports in photocatalytic treatment of water and wastewater: a review

Environmental Chemistry Letters - Rising health issues of Worldwide pollution by fossil fuel products are Fostering the development of safer materials such as biopolymers in many sectors such as...     

Eggshell strength of an obligate brood parasite: a test of the puncture resistance hypothesis

Eggs of several brood parasites have thicker and stronger shells than expected for their size. The present study evaluated the puncture resistance hypothesis for the occurrence of thick-shelled eggs in common cuckoos Cuculus canorus by investigating costs of cuckoo egg ejection in four Acrocephalus warblers—the great reed warbler A. arundinaceus, reed warbler A. scirpaceus, marsh warbler A. palustris and sedge warbler A. schoenobaenus. The three latter species all suffered ejection costs, while ejection was not costly in the larger great reed warbler. The occurrence of ejection costs was negatively related to host bill size. In the marsh warbler, we compared ejection costs in naturally parasitized nests and two experimental treatments, in which broods were parasitized artificially with great reed warbler and conspecific eggs. Hosts damaged their own eggs significantly more often when ejecting the thick-shelled cuckoo eggs than when ejecting the similarly sized but thinner-shelled great reed warbler eggs, providing some support for the puncture resistance hypothesis. Ejection of conspecific eggs did not involve any costs. Furthermore, contrary to predictions derived from the laying damage hypothesis, there was no evidence that egg damage was associated with cuckoo egg laying. Hosts damaging their own eggs during ejection were more likely to subsequently desert their clutches than those that did not. The frequency of clutches smeared with the contents of the ejected egg were positively related to the hypothesized difficulty of foreign egg puncturing. Potential advantages of thicker shells in common cuckoo eggs are discussed.     

The role of microbial mats during primary succession in calcareous dune slacks: an experimental approach

Laminated microbial mats from a sandy beach plain were grown in water-saturated pots in a glass house for six months and then used to assess their effect on the establishment of juveniles of three plant species representing different successional stages in dune slack development. The selected species wereSamolus valerandi, characteristic of pioneer stages,Calamagrostis epigejos, characteristic of more productive, late successional stages, andJuncus alpinoarticulatus, which occurs in a wide range of successional stages. Juveniles of all three species that were placed on top of intact living microbial mats established themselves in the mat.C. epigejos andJ. alpinoarticulatus survived for several weeks but later on their numbers decreased and the total biomass production of the species after six months was poor.S. valerandi, in contrast, grew profusely in intact microbial and algal mats. Heating of the microbial mat by heat sterilization, prior to the experiment, did not improve the performance of the species. When the juveniles were planted in the microbial mats after breaking the surface of the mat, the survival of juveniles ofC. epigejos andJ. alpinoarticulatus was much higher and so was the biomass of surviving plants after six months. Planting ofSamolus in the mats had some positive effect on the survival percentage of the juveniles, but not on the total biomass at the end of the experiment. Slightly lower water tables had a negative effect on the performance of all species. Measurements of the pH in the pots revealed that there were no significant differences in the top layer. Sulphide concentrations were very low in all the pots where juveniles had been planted and also in the pots withS. valerandi. Relatively high concentrations (30–50 μmol/l) were found in pots with poor growth ofJuncus andCalamagrostis plants. These values may exceed toxic levels for these species. Although oxygen concentrations in the pots were generally low, no relation existed between plant biomass and oxygen content, indicating that plant growth was not primarily limited by oxygen stress. These experiments support the idea that microbial mats may assist in extending the life span of early pioneer stages during dune slack succession by inhibiting the growth of species of later successional stages.     

Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Conflict of interest between a nematode and a trematode in an amphipod host: test of the "sabotage" hypothesis

Microphallus papillorobustus is a manipulative trematode that induces strong behavioural alterations in the gamaridean amphipod Gammarus insensibilis, making the amphipod more vulnerable to predation by aquatic birds (definitive hosts). Conversely, the sympatric nematode Gammarinema gammari uses Gammarus insensibilis as a habitat and a source of nutrition. We investigated the conflict of interest between these two parasite species by studying the consequences of mixed infection on amphipod behaviour associated with the trematode. In the field, some amphipods infected by the trematode did not display the altered behaviour. These normal amphipods also had more nematodes, suggesting that the nematode overpowered the manipulation of the trematode, a strategy that would prolong the nematode's life. We hypothesize that sabotage of the trematode by the nematode would be an adaptive strategy for the nematode consistent with recent speculation about co-operation and conflict in manipulative parasites. A behavioural test conducted in the laboratory from naturally infected amphipods yielded the same result. However, exposing amphipods to nematodes did not negate or decrease the manipulation exerted by the trematode. Similarly, experimental elimination of nematodes from amphipods did not permit trematodes to manipulate behaviour. These experimental data do not support the hypothesis that the negative association between nematodes and manipulation by the trematode is a result of the "sabotage" hypothesis.     

Prior residency as a key determinant of social dominance in the willow tit (Parus montanus)

Summary We studied how age, body size and prior residency affected social dominance in the willow tit (Parus montanus) groups. The contribution of each variable was experimentally tested in unisexual two-bird trials, in which the birds were matched for all variables except the one studied. Large birds were dominant over smaller ones (Fig. 1). The effect of body size was more prominent in males than in females. Age had no influence on dominance. Residents became dominant more often than newcomers (Fig. 2). Adulthood or larger body size did not override the advantage of prior residency (Fig. 2). Therefore, the proximate reason for the age-dependent dominance seen in natural willow tit flocks is most likely the prior residency advantage of the adults. Factors connected with fighting ability (body size and age) seem to be less important than the time of establishment of rank, which may reflect the importance of resource value differences between residents and newcomers in this context. The advantage of residency might make it advantageous to be a member of a flock even as a subordinate, rather than being solitary.     

Proximate control of siblicide in cattle egrets: a test of the food-amount hypothesis

Avian sibling aggression plays an important role in controlling food distribution among nest mates. The food-amount hypothesis predicts that the amount of food obtained by siblings acts as a proximate cue controlling their level of aggression. The importance of this cue to cattle egrets (Bubulcus ibis) in Oklahoma was evaluated by studying parental feeding rates, food distribution among nestlings, and nestling aggression in three-chick broods [A-(oldest), B-, and C-chicks]. We compared nests in which siblicide occurred to nests where all three chicks survived (nonsiblicide nests). Relative to nonsiblicide nests, siblicide nests received decreasing amounts of food as the day of siblicide approached. Similarly, aggression levels of all three chicks in siblicide nests increased significantly compared with nonsiblicide nests, with most of the aggressive behavior involving B- and C-chicks. As a result, on a per-day basis, food amount and aggression exhibited a strong, negative correlation, thus providing support for the food-amount hypothesis. Though the total amount of food delivered to siblicide nests declined on the days prior to siblicide, the amount of food received by A-chicks in these nests did not. Relative to nonsiblicide nests, A-chicks in siblicide nests received an increased share (percent of volume) of the food delivered at the expense of C-chicks, with the percent remaining the same for B-chicks. The percentage of food B-chicks received the day after siblicide increased significantly from that received the day before siblicide, with no change occurring in either the amount or percentage of food received by A-chicks; clearly, B-chicks were the beneficiary of their aggressive actions that resulted in the death of C-chicks. We provide evidence supporting the hypothesis that aggression is proximately linked to food amount, possibly by a relatively simple cue such as hunger. Received: 25 November 1994/Accepted after revision: 10 February 1996