Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Grazer avoidance may explain the invasiveness of the red alga Gracilaria vermiculophylla in Scandinavian waters

The invasive red alga Gracilaria vermiculophylla has quickly spread across Europe, but it is unclear whether its success is based on a high tolerance to variations in environmental conditions or to the absence of native grazers that feed on this alga. We tested whether native invertebrate grazers prefer native algae to G. vermiculophylla. Feeding preferences of three common herbivores were quantified when offered G. vermiculophylla and native Fucus vesiculosus, Ceramium virgatum, and Ulva intestinalis in no-, two- and multiple-choice trials. Herbivore growth was measured when fed each of the algae separately. Grazers consumed G. vermiculophylla in no-choice trials, but avoided generally this alga when having a choice. U. intestinalis was always preferred over G. vermiculophylla, and grazers fed with U. intestinalis grew faster than those fed with G. vermiculophylla. We conclude that grazers avoid G. vermiculophylla to most native algae, which may benefit G. vermiculophylla in northern European estuaries.     

New hypothesis helps explain elasmobranch "outburst" on Georges Bank in the 1980s.

Regime shifts are a feature of many ecosystems. During the last 40 years, intensive commercial exploitation and environmental changes have driven substantial shifts in ecosystem structure and function in the northwest Atlantic. In the Georges Bank-southern New England region, commercially important species have declined, and the ecosystem shifted to one dominated by economically undesirable species such as skates and dogfish. Aggregated abundance indices indicate a large increase of small and medium-sized elasmobranchs in the early 1980s following the decline of many commercial species. It has been hypothesized that ecological interactions such as competition and predation within the Georges Bank region were responsible for and are maintaining the "elasmobranch outburst" at the heart of the observed ecosystem shift. We offer an alternative hypothesis invoking population connectivity among winter skate populations such that the observed abundance increase is a result of migratory dynamics, perhaps with the Scotian Shelf (i.e., it is an open population). Here we critically evaluate the survey data for winter skate, the species principally responsible for the increase in total skate abundance during the 1980s on Georges Bank, to assess support for both hypotheses. We show that time series from different surveys within the Georges Bank region exhibit low coherence, indicating that a widespread population increase was not consistently shown by all surveys. Further, we argue that observed length-frequency data for Georges Bank indicate biologically unrealistic population fluctuations if the population is closed. Neither finding supports the elasmobranch outburst hypothesis. In contrast, survey time series for Georges Bank and the Scotian Shelf are negatively correlated, in support of the population connectivity hypothesis. Further, we argue that understanding the mechanisms of ecosystem state changes and population connectivity are needed to make inferences about both the causes and appropriate management responses to large-scale system change.     

Temporal patterns in reproduction may explain variationin mating frequencies in the green-veined white butterfly Pieris napi

There is genetic variation in the female mating rate in the green-veined white butterfly (Pieris napi), and females benefit from male ejaculates that contain both sperm and accessory gland substances. Although polyandry corresponds to higher lifetime fecundity than monandry, some females abstain from remating irrespective of the number of available mates. Explaining genetic variation in mating rates requires that monandrous females perform better than polyandrous ones under some conditions. We experimentally explored the reproductive performance of females either with a low or high intrinsic mating rate by allowing them to mate, feed, and lay eggs freely in a laboratory. Individual females followed different life histories: during the early days of reproduction, females with a low mating rate produced more eggs than females with a high mating rate. Hence, refraining from the benefits of multiple mating may be beneficial, if the time for reproduction is limited, or other female traits associated with polyandry are traded off against longevity. Given the day length of 10 h, a model shows that even if polyandrous females enjoy higher lifetime reproductive success, changeable and unpredictable weather will favor monandry if each period of suitable weather lasts, on average, less than 5 days. Thus, a combination of life history cost and unpredictability of fitness may explain the maintenance of monandry in the wild. Our results are also consistent with the observation that frequency of monandry increases with latitude.     

Inter-sexual conflicts over copulations in the European starling: evidence for the female mate-guarding hypothesis

Recent studies have suggested that conflicts of interests between the sexes may lead to variation in copulation behavior among pairs. We examined differences in the rate and timing of copulation solicitations and copulations among females of different mating status in the facultatively polygynous starling in an attempt to explain why females copulate repeatedly with their mate. All within-pair copulations were female-solicited indicating that females control copulations in the starling. Before egg laying, females solicited copulations at a high rate (usually more than 2 per hour). In contrast to most other species studied so far, females continued to solicit copulations throughout the egg laying period, and also solicited after egg laying (the latest solicitation occurred when the nestlings were 4 days old). Primary females solicited more copulations than monogamous females both during and after the fertile period. Many copulation solicitations of primary females occurred at the nestbox where their male was singing to obtain an extra female. Both primary and secondary females solicited more copulations after the egg-laying period than monogamous females. A large proportion of female copulation solicitations was refused by the male partner: female solicitation resulted in more male refusals in primary females than in monogamous females. Primary females were more likely to be chased aggressively when they solicited a copulation than monogamous females; most aggressive chases occurred when primary females flew towards their male to solicit copulation when he was singing at another nestbox. Overall, our results demonstrate that there is a conflict over copulation between males and females in polygynous pairs. The conflict presumably relates to the cost of sharing male parental investment: females use copulation solicitation behavior to interrupt their singing males apparently in an attempt to prevent them from becoming polygynous. We present the first empirical evidence that female songbirds use copulation solicitation behavior as a form of mate guarding, often in a non-reproductive context. We did not find a positive relationship between copulation rate during the fertile period and the amount of male parental care as is predicted by the paternity confidence hypothesis.     

Can the ratio of aromatic skeletons explain cross-species responses within evolutionarily conserved Ostariophysan alarm cues?: testing the purine-ratio hypothesis

Summary. While the response to damage-released chemical alarm cues within the superorder Ostariophysi appears to be highly conserved across species, it is generally observed that the intensity of response to heterospecific alarm cues decreases with increasing phylogenetic distance. Recent studies have demonstrated that purine-N-oxides function as chemical alarm cues within Ostariophyian fishes and that the nitrogen-oxide functional group is conserved as the chief molecular trigger. According to the purine-ratio hypothesis, these cross-species differences may be due to the relative proportion of different carrier compounds associated with the nitrogen-oxide molecular trigger. To test this hypothesis, we exposed glowlight tetras (Hemigrammus erythrozonus, Characidae, Ostariophysi) to one of five synthetic stimuli (hypoxanthine-3-N-oxide (H3NO), pyridine-N-oxide (PNO) or mixed stimuli of 75 % H3NO-25 % PNO, 50 % H3NO-50 % PNO, or 25 % H3NO-75 % PNO), natural conspecific chemical alarm cue or a distilled water control. We quantified changes in shoal cohesion and vertical area use as species typical indicators of an antipredator response. As predicted, response intensity decreased as the ratio of hypoxanthine-3-N-oxide to pyridine-N-oxide decreased and the strongest response was to natural alarm cue. These results suggest that species-specific carrier compounds may account for the well-documented cross-species differences in the response to heterospecific alarm cues within phylogenetically related taxa.     

Sex differences in the repeatability of boldness over metamorphosis

Consistent individual differences in boldness have been identified in many species and can have important effects on fitness. In most animals, juveniles face different costs and benefits of risk-taking behavior than do adults. Furthermore, profound changes in hormones, morphology and environment often occur when juveniles become adults. Therefore, the boldness of individuals might change with ontogeny. In field crickets, adult males call to attract sexually receptive females, and male calling increases predation risk. We measured the repeatability of boldness (latency to emerge from a safe refuge) in both male and female crickets. Each cricket was tested once as a small nymph and once as an adult. We found that boldness was repeatable across metamorphosis in females, but not in males. Males became less bold with maturation, a result that we predicted because of the risk associated with calling for mates. We also found that in general, nymphs were bolder than adults and that individuals spent more time immobile in response to a predator cue when they were nymphs, versus when they were adults.     

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.     

Changes in hourly precipitation may explain the sharp reduction of discharge in the middle reach of the Yellow River after 2000

The Hekou-Longmen reach, together with local floods, is the main source area for coarse sedimentations into the Yellow River. When total rainfall slightly increased in the area, discharge dramatically decreased by 40%–70% after the year of 2000, and attracting extensive attention in the context of global climate change. High temporal resolution precipitation (timescales between 1 and 4 h) data from the June to September period from 270 rain gauges over the past three decades was mined in order to help explain the phenomenon. Each rainfall event was classified as light/moderate rain, large rain, heavy rain or rainstorm by the event’s rainfall amount, and further classified as low intensity rain, medium intensity rain and high intensity rain by the event’s rainfall intensity. The Mann-Kendall trend test was applied to detect the presence and significance of monotonic trends, and to find the change points in the mean and variance of the precipitation characteristics series, including the amount, intensity, frequency and duration of each rainfall category. Results show that although the total amount of precipitation has slightly increased, the average rainfall intensity has significantly decreased. The larger change happened in light/moderate rain events and low/medium intensity rain events, and the intensity changes have a great extent occurred around the threshold of Non-Runoff Rainfall regime, which was proposed for the approximate calculation of initial losses. Changes in rainfall distribution between different classes of the Runoff Rainfall regime in the 2000s could lead to 0.9 mm less runoff depth (17.3% of the total reduction) than the 1980–1999 period. The study indicates that changes in hourly precipitation may be responsible for the sharp reduction of discharge.     

Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition

The effect of aggressive competition over food resources on energy intake rate is analyzed for individuals of three groups of 25–35 white-faced capuchin monkeys, Cebus capucinus, living in and near Lomas Barbudal Biological Reserve, Costa Rica. An individuals energy intake rate on a given food species was affected by its rank and the number of agonistic interactions within the feeding tree. Dominant group members had higher energy intake rates relative to subordinate group members whether or not there was agonism within the feeding tree. Low- and mid-ranked individuals had lower energy intake rates in trees with higher amounts of aggression, while energy intake rate of high-ranked individuals was not affected by the amount of aggression in the feeding tree. Energy intake was not influenced by the sex of the individual when rank was held constant statistically. Energy intake was positively correlated with total crown energy (measured in kilojoules) within the feeding tree for two of three study groups. This difference may be explained by the quality of each groups territory. Finally, high-ranked individuals are responsible for the majority of agonism within feeding trees and target middle- and low-ranked individuals equally. These findings fit the predictions of current socioecological models for within-group contest competition over food resources. The results of this study suggest that within-group competition affects energy intake rate in white-faced capuchin monkeys.     

Benthic polychaetes in the Ratnagiri bay, India: influence of anthropogenic factors

Study of changes in macrobenthic community structure is an intrinsic part of many environmental monitoring programmes. Hence, it is pivotal to distinguish the effects of natural and anthropogenic factors on these sensitive organisms for accurate assessment and management of coastal environment. Polychaete species diversity of five stations in the Ratnagiri bay was investigated during premonsoon and postmonsoon months in 2007. Though no spatial trends in polychaete diversity vis-à-vis the pollution was visible, the polychaete univariate indices were uniformily better in premonsoon indicating clear seasonal trends. Shannon diversity values ranged from 1.4-2.4 during premonsoon and 0.6-1.6 during postmonsoon. Station 5, which was most impacted by anthropogenic wastes as demonstrated by the anoxic conditions coupled with higher nutrient load, had comparatively higher H' values (1.9 and 1.6) and better evenness values (0.9 and 0.7) during both seasons indicating that the polychaetes were not influenced by anthropogenic contamination. The study indicates that the polychaete distribution and diversity in the bay were governed primarily by variations in sediment texture rather than the anthropogenic disturbances.     

Sperm competition or sperm selection: no evidence for female influence over paternity in yellow dung flies Scatophaga stercoraria

Recent studies of non-random paternity have suggested that sperm selection by females may influence male fertilization success. Here we argue that the problems originally encountered in partitioning variation in non-random mating between male competition and female choice are even more pertinent to interpreting patterns of non-random paternity because of intense sperm competition between males. We describe an experiment with the yellow dung fly, Scatophaga stercoraria, designed to partition variance in the proportion of offspring sired by the second male, P ₂, between males and females, and to control for sperm competition. Large males were shown to have a higher P ₂ than small males but P ₂ was independent of the size of the female’s first mate. This result might suggest an absolute female preference for large males via sperm selection. However, large males have a higher constant rate of sperm transfer and displacement. After controlling for this effect of sperm competition, large males did not achieve higher paternity than small males. We argue that a knowledge of the mechanism of sperm competition is essential so that male effects can be controlled before conclusions are made regarding the influence of sperm selection by females in generating non-random paternity. Received: 4 April 1995 / Accepted after revision: 17 October 1995     

The human footprint in the west: a large-scale analysis of anthropogenic impacts

Anthropogenic features such as urbanization, roads, and power lines, are increasing in western United States landscapes in response to rapidly growing human populations. However, their spatial effects have not been evaluated. Our goal was to model the human footprint across the western United States. We first delineated the actual area occupied by anthropogenic features, the physical effect area. Next, we developed the human footprint model based on the ecological effect area, the zone influenced by features beyond their physical presence, by combining seven input models: three models quantified top-down anthropogenic influences of synanthropic predators (avian predators, domestic dog and cat presence risk), and four models quantified bottom-up anthropogenic influences on habitat (invasion of exotic plants, human-caused fires, energy extraction, and anthropogenic wildland fragmentation). Using independent bird population data, we found bird abundance of four synanthropic species to correlate positively with human footprint intensity and negatively for three of the six species influenced by habitat fragmentation. We then evaluated the extent of the human footprint in relation to terrestrial (ecoregions) and aquatic systems (major rivers and lakes), regional management and conservation status, physical environment, and temporal changes in human actions. The physical effect area of anthropogenic features covered 13% of the western United States with agricultural land (9.8%) being most dominant. High-intensity human footprint areas (class 8-10) overlapped highly productive low-elevation private landholdings and covered 7% of the western United States compared to 48% for low-intensity areas (class 1-3), which were confined to low-productivity high-elevation federal landholdings. Areas within 1 km of rivers were more affected by the human footprint compared to lakes. Percentage human population growth was higher in low-intensity human footprint areas. The disproportional regional effects of the human footprint on landscapes in the western United States create a challenge to management of ecosystems and wildlife populations. Using footprint models, managers can plan land use actions, develop restoration scenarios, and identify areas of high conservation value at local landscapes within a regional context. Moreover, human footprint models serve as a tool to stratify landscapes for studies investigating floral and faunal response to human disturbance intensity gradients.     

Differential predation by males on moulted females may explain the competitive displacement of Gammarus duebeni by G. pulex (Amphipoda)

Summary Several studies have reported the ability of Gammarus pulex to successfully compete with and ultimately displace G. duebeni in freshwater. These studies have been unable, however, to provide adequate explanations for this displacement based on mechanisms of exploitative competition and/or differences in the environmental tolerances of the species. The present study investigates the role of predation as an extreme form of interference competition between G. pulex and G. duebeni. The data presented show that: (1) given the opportunity, males of both species prey on females of the other species, but only on females that have recently moulted; (2) mate guarding in both species significantly reduces the incidence of predation on moulted females; (3) mate guarding by male G. pulex is more successful in reducing predation on moulted females than is guarding by male G. duebeni. We propose that differential predation rate on moulted females may constitute the primary mechanism by which G. pulex displaces G. duebeni. Offprint requests to: J.T.A. Dick     

The race is not to the swift: long-term data reveal pervasive declines in California's low-elevation butterfly fauna

Understanding the ecology of extinction is one of the primary challenges facing ecologists in the 21st century. Much of our current understanding of extinction, particularly for invertebrates, comes from studies with large geographic coverage but less temporal resolution, such as comparisons between historical collection records and contemporary surveys for geographic regions or political entities. We present a complementary approach involving a data set that is geographically restricted but temporally intensive: we focus on three sites in the Central Valley of California, and utilize 35 years of biweekly (every two weeks) surveys at our most long-sampled site. Previous analyses of these data revealed declines in richness over recent decades. Here, we take a more detailed approach to investigate the mode of decline for this fauna. We ask if all species are in decline, or only a subset. We also investigate traits commonly found to be predictors of extinction risk in other studies, such as body size, diet breadth, habitat association, and geographic range. We find that population declines are ubiquitous: the majority of species at our three focal sites (but not at a nearby site at higher elevation) are characterized by reductions in the fraction of days that they are observed per year. These declines are not readily predicted by ecological traits, with the possible exception of ruderal/non-ruderal status. Ruderal species, in slightly less precipitous decline than non-ruderal taxa, are more dispersive and more likely to be associated with disturbed habitats and exotic hosts. We conclude that population declines and extirpation, particularly in regions severely and recently impacted by anthropogenic alteration, might not be as predictable as has been suggested by other studies on the ecology of extinction.     

Delayed maturation in plumage colour: Evidence for the female-mimicry hypothesis in the kestrel

Summary In many sexually dichromatic species, young males have female-like plumage during their first potential breeding year. The female-mimicry hypothesis (FMH) supposes that by possessing female-like plumage young males deceive older conspicuous males into believing that they are females, thus reducing competition from adult males. The status-signalling hypothesis (SSH) supposes that adult males can distinguish sex, but postulates that young males reduce competition from adult males by reliably signaling low status with their dull plumage. We tested these hypotheses in the European kestrel (Falco tinnunculus). Female-like young males settled to breed closer to adult males than did other adult males (Figs. 1a, b). By settling near adult males, young males seemed to increase their chance of mating with adult females. Adult female-young male pairs had better reproductive success than yearling-yearling pairs. These results suggest that there is an adaptive value in possessing a female-like plumage colour in the breeding season. To test the FMH, we measured sexual preference of adult males when adult females and young males were simultaneously shown in an aviary. Adult males were unable to recognize sex, because in half the cases they preferred young males (Fig. 3). However, when adult males and females were shown simultaneously, males preferred females (Fig. 2). Our results support the FMH rather than the SSH, because young males successfully deceived older males by their plumage.     

Trends over time in tree and seedling phylogenetic diversity indicate regional differences in forest biodiversity change

Changing climate conditions may impact the short-term ability of forest tree species to regenerate in many locations. In the longer term, tree species may be unable to persist in some locations while they become established in new places. Over both time frames, forest tree biodiversity may change in unexpected ways. Using repeated inventory measurements five years apart from more than 7000 forested plots in the eastern United States, we tested three hypotheses: phylogenetic diversity is substantially different from species richness as a measure of biodiversity; forest communities have undergone recent changes in phylogenetic diversity that differ by size class, region, and seed dispersal strategy; and these patterns are consistent with expected early effects of climate change. Specifically, the magnitude of diversity change across broad regions should be greater among seedlings than in trees, should be associated with latitude and elevation, and should be greater among species with high dispersal capacity. Our analyses demonstrated that phylogenetic diversity and species richness are decoupled at small and medium scales and are imperfectly associated at large scales. This suggests that it is appropriate to apply indicators of biodiversity change based on phylogenetic diversity, which account for evolutionary relationships among species and may better represent community functional diversity. Our results also detected broadscale patterns of forest biodiversity change that are consistent with expected early effects of climate change. First, the statistically significant increase over time in seedling diversity in the South suggests that conditions there have become more favorable for the reproduction and dispersal of a wider variety of species, whereas the significant decrease in northern seedling diversity indicates that northern conditions have become less favorable. Second, we found weak correlations between seedling diversity change and latitude in both zones, with stronger relationships apparent in some ecoregions. Finally, we detected broadscale seedling diversity increases among species with longer-distance dispersal capacity, even in the northern zone, where overall seedling diversity declined. The statistical power and geographic extent of such analyses will increase as data become available over larger areas and as plot measurements are repeated at regular intervals over a longer period of time.     

Condition-specific competition in container mosquitoes: the role of noncompeting life-history stages

Condition-specific competition, wherein competitive superiority varies with the abiotic environment, can determine species' distributions in a spatially heterogeneous environment. We investigated this phenomenon with two competing container-dwelling mosquitoes. We tested the hypothesis that habitat drying alters the outcome of interspecific competition, predicting that the competitive impact of Aedes albopictus on Aedes aegypti would be severe in wetter environments, but greatly reduced in drier environments. We tested these predictions in a laboratory experiment within cages, with aquatic larvae residing in water-filled cups within the cage, and adults emerging within the cage and ovipositing on the cups. We raised each species alone or with the competitor. Environmental treatments were fluctuating (water in cups evaporated to 50% of the original volume and then cups were refilled), and drying (water in cups evaporated completely and cups remained dry for two weeks before refilling). There was a significant interaction between treatment and species combination for adult populations of both species. Interspecific competition was highly asymmetrical. In the wetter fluctuating environment, interspecific competition had a large negative effect on A. aegypti, but in the drying environment, interspecific competition had a large negative effect on A. albopictus, and relatively little impact on A. aegypti. The main cause of the shift in competitive advantage appeared to be a greater increase in egg mortality for A. albopictus under dry conditions, compared to A. aegypti. Thus, mortality impinging on noncompeting life cycle stages can alter the population level impact of interspecific competition. The hypothesis that dry conditions shift competitive advantage away from A. albopictus is supported in this experiment.     

Males vs workers: testing the assumptions of the haploid susceptibility hypothesis in bumblebees

The haploid state of males in eusocial Hymenoptera—the ants, bees, and wasps—has been proposed as a driving force in the evolution of social behavior under the assumption that haploidy results in higher susceptibility to pathogens. In this study, we present the first test of the assumptions of the “haploid male susceptibility hypothesis”. We challenged males and workers of the bumblebee Bombus terrestris with its parasite Crithidia bombi but found no differences in either initial susceptibility or the intensity of infection between haploid males and diploid females. We reviewed observational studies on parasitism in haplodiploid insects and found that in 15 out of 26 cases, haploid males had lower parasite prevalence. However, the majority of available data related to nontransmissible parasites and thus any general statements about haploid susceptibility remain unclear. Using a simulation model, we studied how diverse genetic mechanisms could affect the values for resistance; results suggest that only a phenomenon that renders workers effectively haploid, e.g., imprinting, could explain our experimental results. A more likely explanation is that, in eusocial Hymenoptera with predominantly female populations, parasites may simply become more adapted to the more common female hosts and, thus, male haploid susceptibility may be hidden due to parasite adaptation. Our results do not support the idea that the haploid susceptibility hypothesis explains the origin or maintenance of social systems in the eusocial Hymenoptera.