Kinship and aggression: do house sparrows spare their relatives?

Kin-selection theory predicts that relatedness may reduce the level of aggression among competing group members, leading to indirect fitness benefits for kin-favoring individuals. To test this hypothesis, we investigated whether relatedness affects aggressive behavior during social activities in captive house sparrow (Passer domesticus) flocks. We found that sparrows did not reduce their aggression towards kin, as neither the frequency nor the intensity of fights differed between close kin and unrelated flock-mates. Fighting success was also unrelated to kinship and the presence of relatives in the flock did not influence the birds’ dominance rank. These results suggest that the pay-offs of reduced aggression towards kin may be low in non-breeding flocks of sparrows, e.g. due to competition among relatives as predicted by a recent refinement of kin-selection theory. Our findings indicate that the significance of kin selection may be restricted in some social systems such as winter aggregations of birds. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

How do dispersal costs and habitat selection influence realized population connectivity?

Despite the importance of dispersal for population connectivity, dispersal is often costly to the individual. A major impediment to understanding connectivity has been a lack of data combining the movement of individuals and their survival to reproduction in the new habitat (realized connectivity). Although mortality often occurs during dispersal (an immediate cost), in many organisms costs are paid after dispersal (deferred costs). It is unclear how such deferred costs influence the mismatch between dispersal and realized connectivity. Through a series of experiments in the field and laboratory, we estimated both direct and indirect deferred costs in a marine bryozoan (Bugula neritina). We then used the empirical data to parameterize a theoretical model in order to formalize predictions about how dispersal costs influence realized connectivity. Individuals were more likely to colonize poor-quality habitat after prolonged dispersal durations. Individuals that colonized poor-quality habitat performed poorly after colonization because of some property of the habitat (an indirect deferred cost) rather than from prolonged dispersal per se (a direct deferred cost). Our theoretical model predicted that indirect deferred costs could result in nonlinear mismatches between spatial patterns of potential and realized connectivity. The deferred costs of dispersal are likely to be crucial for determining how well patterns of dispersal reflect realized connectivity. Ignoring these deferred costs could lead to inaccurate predictions of spatial population dynamics.     

Bilateral symmetry and locomotion: do elliptical regular sea urchins proceed along their longer body axis?

Vagile animals usually have bilaterally symmetrical bodies and proceed with their mouth-end first. Regular sea urchins have, however, radially symmetrical bodies with their mouth facing the substratum and show no preference in which side of the body should be anterior in their locomotion. The elliptical sea urchins in the subfamily Echinometrinae are exceptional among regular sea urchins in having elongated bilateral bodies. We studied whether they showed the preference in the direction of locomotion using Echinometra sp. type A. Directional preference was not observed in the proceedings in an open space. However, they proceeded preferentially with their long axis coinciding with the direction of locomotion when they moved along the water surface or along the wall of the aquarium. The speed of locomotion was the same irrespective of the direction of proceedings and of whether sea urchins moved freely or moved along the water surface or along the walls. We suggest that the bilateral body form and the habit of long-axis lead of this sea urchin have adaptive significance to increase the protected body surfaces, not to facilitate the efficiency in locomotion.     

Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity?

Predators and prey assort themselves relative to each other, the availability of resources and refuges, and the temporal and spatial scale of their interaction. Predictive models of predator distributions often rely on these relationships by incorporating data on environmental variability and prey availability to determine predator habitat selection patterns. This approach to predictive modeling holds true in marine systems where observations of predators are logistically difficult, emphasizing the need for accurate models. In this paper, we ask whether including prey distribution data in fine-scale predictive models of bottlenose dolphin (Tursiops truncatus) habitat selection in Florida Bay, Florida, U.S.A., improves predictive capacity. Environmental characteristics are often used as predictor variables in habitat models of top marine predators with the assumption that they act as proxies of prey distribution. We examine the validity of this assumption by comparing the response of dolphin distribution and fish catch rates to the same environmental variables. Next, the predictive capacities of four models, with and without prey distribution data, are tested to determine whether dolphin habitat selection can be predicted without recourse to describing the distribution of their prey. The final analysis determines the accuracy of predictive maps of dolphin distribution produced by modeling areas of high fish catch based on significant environmental characteristics. We use spatial analysis and independent data sets to train and test the models. Our results indicate that, due to high habitat heterogeneity and the spatial variability of prey patches, fine-scale models of dolphin habitat selection in coastal habitats will be more successful if environmental variables are used as predictor variables of predator distributions rather than relying on prey data as explanatory variables. However, predictive modeling of prey distribution as the response variable based on environmental variability did produce high predictive performance of dolphin habitat selection, particularly foraging habitat.     

Disturbance cues in freshwater prey fishes: do juvenile convict cichlids and rainbow trout respond to ammonium as an ‘early warning’ signal?

Summary.  Several freshwater invertebrate and vertebrate prey species rely on chemosensory cues, including non-injury released disturbance cues, to assess and avoid local predation threats. The prevailing hypothesis is that a pulse of ammonia released by disturbed or stressed prey functions as the disturbance cue. Here, we test this hypothesis in two phylogenetically distant freshwater prey fishes, convict cichlids and rainbow trout. In our first experiment, we measured NH₄ ⁺ concentration in tanks containing shoals of cichlid or trout before and after exposure to a realistic model predator (or left undisturbed). We failed to find an increase in ambient NH₄ ⁺ concentration for either cichlids or trout. In our second experiment, we exposed cichlids or trout to NH₄ ⁺ at 0.1 or 0.5 mg L⁻¹ (or a distilled water control) and measured the change in antipredator behaviour (time moving, foraging rate and area use). We found no consistent increase in antipredator behaviour in response to NH₄ ⁺. In our third study, we exposed cichlids and trout to the disturbance cues of cichlids or trout (versus the odour of undisturbed donors). We found significant increases in antipredator behaviour, regardless of donor species, for both cichlids and trout. Thus, the results of our first two experiments do not support the hypothesis that ammonium functions as a disturbance cue in prey fishes. However, the results of our final experiment do confirm the use of disturbance cues in convict cichlids and rainbow trout and support that hypothesis that the disturbance cue is indeed some generalized metabolic byproduct.     

Specialized bees fail to develop on non-host pollen: do plants chemically protect their pollen?

Bees require large amounts of pollen for their own reproduction. While several morphological flower traits are known to have evolved to protect plants against excessive pollen harvesting by bees, little is known on how selection to minimize pollen loss acts on the chemical composition of pollen. In this study, we traced the larval development of four solitary bee species, each specialized on a different pollen source, when reared on non-host pollen by transferring unhatched eggs of one species onto the pollen provisions of another species. Pollen diets of Asteraceae and Ranunculus (Ranunculaceae) proved to be inadequate for all bee species tested except those specialized on these plants. Further, pollen of Sinapis (Brassicaceae) and Echium (Boraginaceae) failed to support larval development in one bee species specialized on Campanula (Campanulaceae). Our results strongly suggest that pollen of these four taxonomic groups possess protective properties that hamper digestion and thus challenge the general view of pollen as an easy-to-use protein source for flower visitors.     

Industry,incidents and casualties in South West England: what is their relationship and are there social inequalities in their distribution?

This ecological study aimed, through the analysis of 1,146 wards in the South West of England (1998–2002), firstly, to examine whether chemical incidents and public casualties are more likely near complex industry (emissions to land, air or water: Integrated Pollution Control industry, IPC) or industry with emissions to air only (Local Air Pollution Control industry, LAPC). Secondly, the study examined whether industry, incidents and casualties are found close to deprivation. Social inequalities were examined across quintiles of wards. Fifty-two wards (4.5%) contained an IPC industry and 712 (62.1%) an LAPC. Incidents occurred in 132 wards (11.5%), with casualties in 59 (5.1%). Chemical incidents occurred more frequently in wards with LAPC (152, IPC 20); the same was true of casualties (211, 12). With each additional LAPC site in a ward, the risk of an incident rose by 22% (95% confidence interval [CI] 8–38%), suggesting a dose–response relationship. No clear social inequalities were found. In the South West of England, the public are more likely to be affected by an incident occurring at a simple LAPC site rather than a complex IPC site. This has implications for emergency planning which, at present, focusses most attention on the larger, more complex IPC sites.     

Invasive species distribution modeling (iSDM): Are absence data and dispersal constraints needed to predict actual distributions?

Species distribution models (SDMs) based on statistical relationships between occurrence data and underlying environmental conditions are increasingly used to predict spatial patterns of biological invasions and prioritize locations for early detection and control of invasion outbreaks. However, invasive species distribution models (iSDMs) face special challenges because (i) they typically violate SDM's assumption that the organism is in equilibrium with its environment, and (ii) species absence data are often unavailable or believed to be too difficult to interpret. This often leads researchers to generate pseudo-absences for model training or utilize presence-only methods, and to confuse the distinction between predictions of potential vs. actual distribution. We examined the hypothesis that true-absence data, when accompanied by dispersal constraints, improve prediction accuracy and ecological understanding of iSDMs that aim to predict the actual distribution of biological invasions. We evaluated the impact of presence-only, true-absence and pseudo-absence data on model accuracy using an extensive dataset on the distribution of the invasive forest pathogen Phytophthora ramorum in California. Two traditional presence/absence models (generalized linear model and classification trees) and two alternative presence-only models (ecological niche factor analysis and maximum entropy) were developed based on 890 field plots of pathogen occurrence and several climatic, topographic, host vegetation and dispersal variables. The effects of all three possible types of occurrence data on model performance were evaluated with receiver operating characteristic (ROC) and omission/commission error rates. Results show that prediction of actual distribution was less accurate when we ignored true-absences and dispersal constraints. Presence-only models and models without dispersal information tended to over-predict the actual range of invasions. Models based on pseudo-absence data exhibited similar accuracies as presence-only models but produced spatially less feasible predictions. We suggest that true-absence data are a critical ingredient not only for accurate calibration but also for ecologically meaningful assessment of iSDMs that focus on predictions of actual distributions.     

Evaluating animal personalities: do observer assessments and experimental tests measure the same thing?

The animal personality literature uses three approaches to assess personality. However, two of these methods, personality ratings and experimentation, have been little compared in captivity and never compared in the wild. We assessed the boldness of wild chacma baboons Papio ursinus using both ratings and experimental methods. Boldness was experimentally assessed when individuals were presented with a novel food item during natural foraging. The boldness of the same individuals was rated on a five-point scale by experienced observers. The ratings and experimental assessments of boldness were found to correlate positively and in a linear fashion. When considered categorically the two approaches showed variable agreement depending on the number of categories assigned and the cut-off criteria adopted. We suggest that the variation between approaches arises because each method captures different aspects of personality; ratings consider personality in absolute terms (using predefined criteria) and multiple contexts, while experimental assessments consider personality in relative terms (using experimental scores relative to the population average) and in limited contexts. We encourage animal personality researchers to consider adopting both methodologies in future studies. We also propose that future studies restrict their analyses to continuous data, since the greatest comparability between methods was found with these data. However, if individuals must be categorised, we suggest that researchers either (a) analyse only those individuals categorised as bold or shy by both ratings and experimental approaches or, if these methods cannot be employed simultaneously, (b) do not use approach-specific criteria but choose a cut-off that can be compared by both approaches.     

Time budgets and foraging in a Malagasy primate: do sex differences reflect reproductive condition and female dominance?

Female mammals commonly employ behavioral tactics of modulating activity levels and foraging behavior to counter the energetic burden of reproduction; these behavioral changes are reflected as intersexual differences. Traditional views of Malagasy primates posit that high reproductive costs select for female dominance which guarantees to energetically stressed females priority of resource access. I tested predictions regarding reproductive influences on sex differences in time budgets and foraging behavior using two groups of Milne-Edwards' sifaka (Propithecus diadema edwardsi) in southeastern Madagascar. Compared to males, females increased neither feeding nor resting time during gestation or lactation. Sex differences were essentially absent in all foraging time variables examined (time, duration, rate). In contrast, dietary composition diverged between the sexes in some months. The possibility that females selected particular food items to boost nutrient and energetic intake to meet increased requirements during reproduction must be further clarified with nutritional analyses. Sex differences in plant part choices coincided with lactation in one of the two study groups. Thus, the timing of sex differences in feeding patterns of P. d. edwardsi only partially supports the prediction that sex differences are most pronounced during the period of greatest female energetic demand. A comparative review indicated no tight association between female dominance and sex differences in foraging among Malagasy primates. Traditional female dominance theory falls short of explaining the observed patterns. The results of my study coupled with recent evidence suggest that non-behavioral tactics involving energy conservation and storage require further attention as mechanisms by which female lemurs cope with reproductive costs. Received: 12 June 1998 / Accepted after revision: 10 October 1998     

Growth rates, seed size, and physiology: do small-seeded species really grow faster?

Relative growth rate (RGR) is currently the most commonly used method for measuring and comparing species' intrinsic growth potential. Comparative studies have, for example, revealed that small-seeded species have higher RGR, leading to the common belief that small-seeded species possess physiological adaptations for rapid growth that would allow them to outgrow large-seeded species, given sufficient time. We show that, because RGR declines as individual plants grow, it is heavily biased by initial size and does not measure the size-corrected growth potential that determines the outcome of competition in the long-term. We develop a daily growth model that includes a simple mechanistic representation of aboveground and belowground growth and its dependency on plant size and environmental factors. Intrinsic growth potential is encapsulated by the size-independent growth coefficient, G. We parameterized the model using repeated-harvest data from 1724 plants of nine species growing in contrasting nutrient and temperature regimes. Using information-theoretic criteria, we found evidence for interspecific differences in only three of nine model parameters: G, aboveground allocation, and frost damage. With other parameters shared between species, the model accurately reproduced above- and belowground biomass trajectories for all nine species in each set of environmental conditions. In contrast to conventional wisdom, the relationship between G and seed size was positive, despite a strong negative correlation between seed size and average RGR, meaning that large-seeded rather than small-seeded species have higher size-corrected growth potential. Further, we found a significant positive correlation between G and frost damage that, according to simulations, causes rank reversals in final biomass under daily temperature changes of +/- 5 degrees C. We recommend the wider use of this new kind of plant growth analysis as a better way of understanding underlying differences in species' physiology; but we recognize that RGR is still a useful metric if considering the potential rate of population increase in empty habitats.     

Making scents of defense: do fecal shields and herbivore-caused volatiles match host plant chemical profiles?

Many plant families have aromatic species that produce volatile compounds which they release when damaged, particularly after suffering herbivory. Monarda fistulosa (Lamiaceae) makes and stores volatile essential oils in peltate glandular trichomes on leaf and floral surfaces. This study examined the larvae of a specialist tortoise beetle, Physonota unipunctata, which feed on two M. fistulosa chemotypes and incorporate host compounds into fecal shields, structures related to defense. Comparisons of shield and host leaf chemistry showed differences between chemotypes and structures (leaves vs. shields). Thymol chemotype leaves and shields contained more of all compounds that differed than did carvacrol chemotypes, except for carvacrol. Shields had lower levels of most of the more volatile chemicals than leaves, but more than twice the amounts of the phenolic monoterpenes thymol and carvacrol and greater totals. Additional experiments measured the volatiles emitted from M. fistulosa in the absence and presence of P. unipunctata larvae and compared the flower and foliage chemistry of plants from these experiments. Flowers contained lower or equal amounts of most compounds and half the total amount, compared to leaves. Plants subjected to herbivory emitted higher levels of most volatiles and 12 times the total amount, versus controls with no larvae, including proportionally more of the low boiling point chemicals. Thus, chemical profiles of shields and volatile emissions are influenced by the amounts and volatilities of compounds present in the host plant. The implications of these results are explored for the chemical ecology of both the plant and the insect.     

The relative influence of habitat loss and fragmentation: do tropical mammals meet the temperate paradigm?

The relative influence of habitat loss vs. habitat fragmentation per se (the breaking apart of habitat) on species distribution and abundance is a topic of debate. Although some theoretical studies predict a strong negative effect of fragmentation, consensus from empirical studies is that habitat fragmentation has weak effects compared with habitat loss and that these effects are as likely to be positive as negative. However, few empirical investigations of this issue have been conducted on tropical or wide-ranging species that may be strongly influenced by changes in patch size and edge that occur with increasing fragmentation. We tested the relative influence of habitat loss and fragmentation by examining occupancy of forest patches by 20 mid- and large-sized Neotropical mammal species in a fragmented landscape of northern Guatemala. We related patch occupancy of mammals to measures of habitat loss and fragmentation and compared the influence of these two factors while controlling for patch-level variables. Species responded strongly to both fragmentation and loss, and response to fragmentation generally was negative. Our findings support previous assumptions that conservation of large mammals in the tropics will require conservation strategies that go beyond prevention of habitat loss to also consider forest cohesion or other aspects of landscape configuration.     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

Male mating history and female fecundity in the Lepidoptera: do male virgins make better partners?

In insects, large ejaculate and associated materials, including spermatophores, appear to have evolved via sexual selection acting on males to either delay female remating or to increase the rate of egg-laying. It is also possible, however, that females use nutrients transferred during mating to increase their lifetime fecundity. If so, male ejaculate size may also have evolved under natural selection as a form of paternal investment. In Lepidoptera, males with a greater number of prior matings tend to produce smaller spermatophores. However, the reported effects of male mating history on female fecundity vary widely among species. We therefore performed a meta-analysis using data from 29 studies of 25 species. Overall, the reproductive output of females mated to virgin males was significantly higher than that of females mated to sexually experienced males (Hedges d=0.33, P<0.01). A sample size of around 145 females per male mating type is required to detect an effect of this size with 80% statistical power at =0.05 (two-tailed). There was no difference in mean effect size between butterflies/skippers and moths. After controlling for any effect of taxonomic group, however, the mean effect size for polyandrous species was significantly greater than that for monandrous species (Hedges d=0.45 vs 0.25, P=0.01). We then discuss possible reasons why male mating history, presumably acting through its effect on spermatophore size, might have a stronger effect in polyandrous than monandrous species.Communicated by A. Cockburn     

Variation in growth rate and reproductive output in British populations of the dorid nudibranch Adalaria proxima: consequences of restricted larval dispersal?

 Growth and reproduction were compared among six geographically and genetically distinct intertidal populations of the annual, semelparous, dorid nudibranch Adalaria proxima (Alder & Hancock) to evaluate variation in fitness-related traits. The six populations spanned the geographic range in the northern British Isles: NE England (Cowling Scar), E Scotland (Kinkell Braes), NW Scotland (Loch Eriboll), W Scotland (Cuan Ferry), Northern Ireland (Portaferry), and N Wales (Menai Bridge). Nudibranchs from five sites were collected in July to August 1992 as post-metamorphic juveniles and were laboratory-reared under the same conditions of ambient temperature and photoperiod for up to 10 months and the completion of spawning. Individuals from the sixth site were added to the experiment in December 1992. Growth was monitored every 2 weeks, and reproductive performance was expressed as a weight-adjusted dimensionless index (ΣRI) of each individual's spawnings summed over the reproductive period. In general, larger nudibranchs produced larger first spawn masses and more total spawn than did smaller nudibranchs, but these size-related trends were observed only in some populations. The patterns of energy partitioning to spawnings varied significantly among populations, from allocations of a large number of eggs to few spawn masses (Loch Eriboll) to production of many small spawnings over a long spawning period (Portaferry). There was no relationship between maximum body size and the amount of spawn produced after the first spawning, nor to the length of the spawning period or the number of spawn produced. Both Menai Bridge and Kinkell Braes had low mean population ΣRI, reflecting a very poor reproductive performance, given their large maximum (pre-spawning) body sizes. By contrast, the Loch Eriboll, Cuan Ferry, and Portaferry populations all displayed high mean population ΣRI, albeit as a result of differing combinations of numbers and sizes of spawn masses and duration of the spawning period. This high variance of reproductive allocation among populations, and previous evidence of relatively stable among-population differences in allozyme frequencies, adult color, and embryo characteristics suggest very restricted larval transport of lecithotrophic larvae of A. proxima. Received: 10 December 1998 / Accepted: 23 March 2000     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.