Synchrony in egg-laying and reproductive success of neighboring common murres,Uria aalge

Common murres (Uria aalge) are highly colonial; pairs often breed at the highest possible densities, in bodily contact with neighbors. At Bluff, a colony in western Alaska, we tested for synchrony in egg laying at various spatial scales and found little evidence for higher synchrony, either within study plots of 15–195 pairs, or within subplots containing several pairs, than among plots in a 5-year study. Egg laying of neighbors generally was more synchronous than expected based on overall frequency distributions in laying dates, however. Breeding success was positively correlated with the number of breeding neighbors and the number of neighbors tending eggs or nestlings at the time of egg laying. Breeding success of pairs with neighbors was positively related to the breeding success of neighbors. Pairs that produced eggs synchronously with at least one neighboring pair had higher success than those that began breeding either before or after their neighbors. Most reproductive failures at Bluff are due to accidental egg loss and predation on eggs by common ravens, Corvus corax, soon after laying. By occupying space where a raven might otherwise land and defending their own eggs, active breeding neighbors locally reduce the probability of egg predation. Active breeding neighbors also are less likely to flush and accidentally dislodge nearby eggs when disturbed than are nonbreeders. Murres breeding synchronously with neighbors have the highest assurance of the presence of active breeding neighbors both at the time of egg laying and throughout their reproductive attempts. Groups of neighboring murres can be considered small “selfish herds,” demonstrating by-product mutualism through their continued presence and defense of their own eggs and nestlings. Despite the advantages of breeding synchronously with neighbors, early breeding may often be favored, however. Received: 22 January 1996/Accepted after revision: 16 June 1996     

Prey density and the behavioral flexibility of a marine predator: the common murre (Uria aalge)

Flexible time budgets allow individual animals to buffer the effects of variable food availability by allocating more time to foraging when food density decreases. This trait should be especially important for marine predators that forage on patchy and ephemeral food resources. We examined flexible time allocation by a long-lived marine predator, the Common Murre (Uria aalge), using data collected in a five-year study at three colonies in Alaska (USA) with contrasting environmental conditions. Annual hydroacoustic surveys revealed an order-of-magnitude variation in food density among the 15 colony-years of study. We used data on parental time budgets and local prey density to test predictions from two hypotheses: Hypothesis A, the colony attendance of seabirds varies nonlinearly with food density; and Hypothesis B, flexible time allocation of parent murres buffers chicks against variable food availability. Hypothesis A was supported; colony attendance by murres was positively correlated with food over a limited range of poor-to-moderate food densities, but independent of food over a broader range of higher densities. This is the first empirical evidence for a nonlinear response of a marine predator's time budget to changes in prey density. Predictions from Hypothesis B were largely supported: (1) chick-feeding rates were fairly constant over a wide range of densities and only dropped below 3.5 meals per day at the low end of prey density, and (2) there was a nonlinear relationship between chick-feeding rates and time spent at the colony, with chick-feeding rates only declining after time at the colony by the nonbrooding parent was reduced to a minimum. The ability of parents to adjust their foraging time by more than 2 h/d explains why they were able to maintain chick-feeding rates of more than 3.5 meals/d across a 10-fold range in local food density.     

Differential breeding investment in bridled and non-bridled common guillemots (Uria aalge): morph of the partner matters

Polymorphism is the coexistence of two or more phenotypically distinct and genetically determinate forms in a population and implies a selective balance between the alternative morphs to permanently exist. Common guillemots Uria aalge occur in two genetically distinct morphs—a non-bridled and a bridled—the latter with white eye rings and a well-defined stripe behind the eyes. In this study, we investigated differences between the morphs with regard to reproductive parameters. We used a detailed family-based sample providing data on mother, father, and chick over three breeding seasons. The mating between morphs was random but pure non-bridled and pure bridled pairs produced smaller chicks at age 15 days than the two mixed pair compositions. Body mass of the adults showed much the same pattern; pure pairs having lower body mass than mixed pairs. There were no differences between the two morphs in reproductive parameters without considering the morph of the partner. This suggests that reproductive decisions in some way not only depend on the morph but also on the tactic of the partner. Different reproductive strategies between morph family groups as found in this study may contribute to the understanding of the existence of a balanced polymorphism in common guillemots. The overall breeding conditions during the years of this study were good. However, over time in a variable environment, we suggest that tactics of different family groups may have different success stabilizing the frequency of non-bridled and bridled birds over time.     

Association of body mass with price of bushmeat in Nigeria and Cameroon

Spatially extensive patterns of bushmeat extraction (and the processes underlying these patterns) have not been explored. We used data from a large sample (n= 87) of bushmeat trading points in urban and rural localities in Nigeria and Cameroon to explore extraction patterns at a regional level. In 7,594 sample days, we observed 61,267 transactions involving whole carcasses. Rural and urban trading points differed in species for sale and in meat condition (fresh or smoked). Carcass price was principally associated with body mass, with little evidence that taxonomic group (primate, rodent, ungulate, or mammalian carnivore) affected price. Moreover, meat condition was not consistently associated with price. However, some individual species were more expensive throughout the region than would be expected for their size. Prices were weakly positively correlated with human settlement size and were highest in urban areas. Supply did not increase proportionally as human settlement size increased, such that per capita supply was significantly lower in urban centers than in rural areas. Policy options, including banning hunting of more vulnerable species (those that have low reproductive rates), may help to conserve some species consumed as bushmeat because carcass prices indicate that faster breeding, and therefore the more sustainable species, may be substituted and readily accepted by consumers.     

Environmental hypoxia and haemocyanin between-individual variability in Norway lobsters Nephrops norvegicus (L.)

The extent to which exposure to environmental hypoxia (PO₂=8, 6, and 4 kPa, approximately 40%, 30%, and 20% saturation, respectively) resulted in an increase in the concentration of the respiratory pigment haemocyanin ([Hc]) in individuals of the Norway lobster Nephrops norvegicus depended crucially on the initial [Hc] as well as the intensity of the hypoxia itself. While mean [Hc] did increase with decreasing PO₂ (and the variation decreased), for individuals with relatively high initial [Hc] there was no change or even a decrease observed. The greatest hypoxia-related increases in [Hc] were observed in the individuals with the lowest initial [Hc]. Consequently the notion of an 'optimum' [Hc] for a given level of oxygenation was postulated. The changes in [Hc] recorded took place over a considerably shorter time scale (hours rather than days) than has been observed in previous studies. It was not possible to correlate environmental PO₂ and median [Hc] in the field. However, it was interesting that the [Hc] frequency distribution curve for lobsters collected from a site that had been markedly hypoxic (PO₂=8.0 kPa, approximately 40% saturation) until a few days before sampling was almost exactly the same shape (i.e. leptokurtic) and position (i.e. similar median) to that obtained when lobsters were exposed to PO₂=8.0 kPa in the laboratory. Although it would have been desirable to investigate the effect of hypoxia on the frequency distribution of a physiological trait, in this case [Hc], it was not possible due to insufficient sample size, even though the sample sizes employed were considerably larger than those normally used in physiological investigations.     

Metabolic rates of epipelagic marine zooplankton as a function of body mass and temperature

The metabolic rates (oxygen uptake, ammonia excretion, phosphate excretion) of epipelagic marine zooplankton have been expressed as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using the multiple-regression method. Zooplankton data used for this analysis are from phylogenetically mixed groups (56 to 143 species, representing 7 to 8 phyla, body mass range: 6 orders of magnitude) from various latitudes (habitat temperature range:-1.4° to 30°C). The results revealed that 84 to 96% of variation in metabolic rates is due to body mass and habitat temperature. Among the various body-mass units, the best correlation was provided by carbon and nitrogen units for all three metabolic rates. Oxygen uptake, ammonia excretion and phosphate excretion are all similar in terms of body-mass effect, but differ in terms of temperature effect. With carbon or nitrogen body-mass units, calculated Q₁₀ values are 1.82 to 1.89 for oxygen uptake, 1.91 to 1.93 for ammonia excretion and 1.55 for phosphate excretion. The effects of body mass and habitat temperature on the metabolic quotients (O:N, N:P, O:P) are insignificant. The present results for oxygen-uptake rate vs body mass do not differ significantly from those reported for general poikilotherms by Hemmingsen and for crustaceans by Ivleva at a comparable temperature (20°C). The importance of a body-mass measure for meaningful comparison is suggested by the evaluation of the habitat-temperature effect between mixed taxonomic groups and selected ones. Considering the dominant effects of body mass and temperature on zooplankton metabolic rates, the latitudinal gradient of community metabolic rate for net zooplankton in the ocean is estimated, emphasizing the non-parallelism between community metabolic rates and the standing stock of net zooplankton.     

Metabolic rates of epipelagic marine copepods as a function of body mass and temperature

Metabolic rates (oxygen consumption, ammonia excretion, phosphate excretion) have been calculated as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using multiple regression. The metabolic data used for this analysis were species structured, collected from Arctic to Antarctic seas (temperature range: -1.7°C to 29.0°C). The data were further divided into geographical and/or seasonal groups (35 species and 43 data sets for oxygen consumption; 38 species and 58 data sets for ammonia excretion; 22 species and 31 data sets for phosphate excretion). The results revealed that the variance attributed to body mass and temperature was highest (93-96%) for oxygen consumption rates, followed by ammonia excretion rates (74-80%) and phosphate excretion rates (46-56%). Among the various body mass units, the best correlation was provided by the nitrogen unit, followed by the dry weight unit. The calculated Q₁₀ values varied slightly according to the choice of body mass units; overall ranges were 1.8-2.1 for oxygen consumption rates, 1.8-2.0 for ammonia excretion rates and 1.6-1.9 for phosphate excretion rates. The effects of body mass and temperature on the metabolic quotients (O:N, N:P, O:P) were insignificant in most cases. Although the copepod metabolic data used in the present analysis were for adult and pre-adult stages, possible applications of the resultant regression equations to predict the metabolic rates of naupliar and early copepodite stages are discussed. Finally, global patterns of net growth efficiency [growth (growth+metabolism)^-1] of copepods were deduced by combining the present metabolic equation with Hirst and Lampitt's global growth equation for epipelagic marine copepods.     

Environmental variability and allocation trade-offs maintain species diversity in a process-based model of succulent plant communities

Ecological theory suggests that environmental variability can promote coexistence, provided that species occupy differential niches. In this study, we focus on two questions: (1) Do allocation trade-offs provide a sufficient basis for niche differentiation in succulent plant communities? (2) What is the relative importance of different forms of environmental variability on species diversity and community composition? We approach these questions with a generic, individual-based simulation model. In our model, plants compete for water in a spatially explicit environment. Species differ in their size at maturity and in the allocation of carbon to roots, leaves and storage tissue. The model was fully specified with independent literature data. Model output was compared to characteristics of a species-rich community in the semi-arid Richtersveld (South Africa). The model reproduced the coexistence of plants with different sizes at maturity, the dominance of succulent shrubs, and the level of vegetation cover. We analyzed the effects of three forms of environmental variability: (a) temporal fluctuations in precipitation (rain and fog), (b) spatial heterogeneity of water supply due to run-on and run-off processes and (c) ‘rock pockets’ that limit root competition in space. The three types of variability had differential effects on diversity: diversity exhibited a strong hump-shaped response to temporal variation. Spatial variability increased diversity, with the strongest increase occurring at intermediate levels of temporal variability. Finally, rock pockets had the weakest effect, but contributed to diversity by providing refuges for small species, particularly at low temporal variability. The model thus shows that spatio-temporal variation of resource supply can maintain diversity over long time scales even in small systems, as is the case in the Richtersveld succulent communities. Trade-offs in allocation provide the basis for necessary niche differentiation. By describing resource competition between individual plants, our model provides a mechanistic basis for the link from species traits to community composition at given environmental conditions. It thereby contributes to an understanding of the forces shaping plant communities. Such an understanding is critical to reduce the threats environmental change poses to biodiversity and ecosystem services.     

Patterns of body mass senescence and selective disappearance differ among three species of free-living ungulates

Declines in survival and reproduction with age are prevalent in wild vertebrates, but we know little about longitudinal changes in behavioral, morphological, or physiological variables that may explain these demographic declines. We compared age-related variation in body mass of adult females in three free-living ungulate populations that have been the focus of long-term, individual-based research: bighorn sheep (Ovis canadensis) at Ram Mountain, Canada; roe deer (Capreolus capreolus) at Trois Fontaines, France; and Soay sheep (Ovis aries) on St. Kilda, Scotland. We use two recently proposed approaches to separate contributions to age-dependent variation at the population level from within-individual changes and between-individual selective disappearance. Selective disappearance of light individuals in all three populations was most evident at the youngest and oldest ages. In later adulthood, bighorn sheep and roe deer showed a continuous decline in body mass that accelerated with age while Soay sheep showed a precipitous decrease in mass in the two years preceding death. Our results highlight the importance of mass loss in explaining within-individual demographic declines in later adulthood in natural populations. They also reveal that the pattern of senescence, and potentially also the processes underlying demographic declines in late life, can differ markedly across related species with similar life histories.     

Effects of temperature and body mass on metabolic rates of sprat, Sprattus sprattus L.

Sprat, Sprattus sprattus L., is a small schooling clupeid forming large stocks in several ecosystems. Despite its high trophodynamic impact, little is known about its energy consumption rates. As a central component of a bioenergetic budget, metabolic rates of sprat from 3.11 to 9.71 g wet weight (WW) were measured at nine different temperatures (T) ranging from 9 to 21°C using a computer-controlled intermittent-flow respirometer. Routine metabolism (R _R) was related to T (°C) and WW (g) by R _R = 0.074 WW^1.077 e^{0.080 T}. Standard metabolic rates (R _S) as calculated from the 10% percentiles of the repeated measurements were on average 12% lower and still influenced by continuous swimming activity: R _S = 0.069 WW^1.073 e^{0.078 T}. We interpret the deviation of the scaling exponent b from typically found exponents of b ~ 0.8 as a consequence of permanently elevated activity level. The high permanent swimming activities also indicated that the concept of standard metabolism may not be meaningful in schooling planktivorous fish. These results suggest that generally in bioenergetic models for clupeid schooling fish the activity multipliers should be chosen very conservatively.     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Setting population targets for mammals using body mass as a predictor of population persistence

Conservation planning and biodiversity assessments need quantitative targets to optimize planning options and assess the adequacy of current species protection. However, targets aiming at persistence require population‐specific data, which limit their use in favor of fixed and nonspecific targets, likely leading to unequal distribution of conservation efforts among species. We devised a method to derive equitable population targets; that is, quantitative targets of population size that ensure equal probabilities of persistence across a set of species and that can be easily inferred from species‐specific traits. In our method, we used models of population dynamics across a range of life‐history traits related to species’ body mass to estimate minimum viable population targets. We applied our method to a range of body masses of mammals, from 2 g to 3825 kg. The minimum viable population targets decreased asymptotically with increasing body mass and were on the same order of magnitude as minimum viable population estimates from species‐ and context‐specific studies. Our approach provides a compromise between pragmatic, nonspecific population targets and detailed context‐specific estimates of population viability for which only limited data are available. It enables a first estimation of species‐specific population targets based on a readily available trait and thus allows setting equitable targets for population persistence in large‐scale and multispecies conservation assessments and planning.     

Diving behaviour of Little Penguins from four colonies across their whole distribution range: bathymetry affecting diving effort and fledging success

Little Penguins, Eudyptula minor, breed in several small colonies in New Zealand and Australia. In this study, we compare the birds’ diving performances at different sites situated throughout their breeding range. Environmental conditions and breeding success vary drastically amongst colonies, but all birds feed on similar types of prey and face similar limitations on their foraging range. We examined several diving parameters and calculated the proportion of foraging zone available during breeding to examine whether oceanographic and geographic factors in the foraging zone can explain variations in diving behaviour and fledging success among the different colonies. In colonies with high fledging success, Penguin Island and Oamaru, penguins made shallow dives <50 m depth and had lower diving effort. More than 90% of the foraging zone was in waters <50 m depth in these colonies. Motuara Island also has shallow waters with 95% <50 m depth, but the fledging success was low. Phillip Island has only 42% of waters <50 m and comparatively low fledging success. Thus, penguins dived deeper and showed a higher diving effort in colonies with lower fledging success (Motuara Island and Phillip Island), indicating that they were disadvantaged compared to conspecifics from other colonies that dived shallower and with a lesser diving effort. We concluded that bathymetry is an important factor, but not the only one, which influences fledging success.     

The relative role of age and experience in determining variation in body mass during the early breeding career of the Common Tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>)

Body mass is a consistent individual trait that characterises the state of adult birds and mammals and is positively related with long-term reproductive success. However, whether and to what extent body mass changes over lifetime in long-lived birds is poorly studied. In this paper, we investigate how individual body mass varies with age. Furthermore, we try to separate possible effects of age and experience on body mass. This study was conducted in a Common Tern colony on the German Wadden Sea coast. Transponders allowed registration of individuals throughout the breeding season and consecutive years with an antenna system combined with electronic balances for recording individual body-mass changes within and between years. Individual body masses of males and females were measured during three stages of the breeding season: at arrival, during incubation and chick rearing when mass was lowest in both sexes. Individual-based longitudinal analyses clearly showed that body mass during arrival, incubation and chick rearing increased up to an age of 5 or 6 years. First-time breeders had lower body mass than experienced breeders. Experience had stronger effects on incubation mass than age. Recruiting age also affected body-mass development of breeders: Three-year-old recruits showed a stronger increase in mass with experience than 4-year-old recruits. We assume that increasing experience enables birds to cope better with the physiological challenge of reproduction. To explain the general phenomenon of higher body mass in older birds, our results support the constraint hypothesis rather than either the selection or restraint hypothesis.     

Interannual variability of growth and reproduction in Bursera simaruba: the role of allometry and resource variability

Plants are expected to differentially allocate resources to reproduction, growth, and survival in order to maximize overall fitness. Life history theory predicts that the allocation of resources to reproduction should occur at the expense of vegetative growth. Although it is known that both organism size and resource availability can influence life history traits, few studies have addressed how size dependencies of growth and reproduction and variation in resource supply jointly affect the coupling between growth and reproduction. In order to understand the relationship between growth and reproduction in the context of resource variability, we utilize a long-term observational data set consisting of 670 individual trees over a 10-year period within a local population of Bursera simaruba (L.) Sarg. We (1) quantify the functional form and variability in the growth-reproduction relationship at the population and individual-tree level and (2) develop a theoretical framework to understand the allometric dependence of growth and reproduction. Our findings suggest that the differential responses of allometric growth and reproduction to resource availability, both between years and between microsites, underlie the apparent relationship between growth and reproduction. Finally, we offer an alternative approach for quantifying the relationship between growth and reproduction that accounts for variation in allometries.     

环境基准研究与环境标准制定进展及展望

环境基准是一个非常棘手且具有挑战性的科学难题,环境标准制定与修订是一项长期的、艰巨的任务。根据国际上该方向近年来的发展,对环境基准和环境标准的概念与内涵进行了论述,对过去的认识进行了纠正,指出:环境基准不等于环境质量基准,环境基准应该包括环境质量基准和污染环境修复基准2个方面;相应地,环境标准则应包括环境质量标准和污染环境修复标准2个部分。在此基础上,概述了国际和国内有关环境质量基准研究方面取得的显著成绩,列举了法国(作为西欧的代表)、加拿大(作为美洲的代表)和意大利(作为中欧的代表)在环境质量标准制定过程中的先进做法,介绍了美国、英国和俄罗斯等国家在污染环境修复基准研究与污染环境修复标准制定等方面的进展。最后,对我国今后环境基准的研究进行了展望。     

邻苯二甲酸二乙基己酯(DEHP)污染及其毒性研究进展

邻苯二甲酸二乙基己酯(DEHP)作为重要的增塑剂被广泛应用于涂料、食品包装、医疗器材、儿童玩具等产品中。研究表明DEHP在水、土壤、空气等各个环境要素以及食物、饮用水中已被普遍检出,并对环境产生潜在的危害。本文通过分析国内外DEHP的环境暴露和毒性效应的研究成果,总结了DEHP在室外大气、室内空气、土壤、地表水、地下水、食品和饮用水中的污染现状,并对DEHP的替代产品进行总结;此外,本文深入探讨了DEHP对水生生物和陆生生物的生态毒性效应,以及对生殖发育、肝脏、呼吸系统和神经系统等的健康毒性效应。最后,结合DEHP在环境暴露调查和毒性效应研究方面的不足,指出需要进一步加强我国DEHP的环境暴露调查,制定相关环境基准值与标准限值,开展慢性生态毒性效应和人体健康毒性效应等方面的研究。