A predator's perspective of nest predation: predation by red squirrels is learned, not incidentalPelech, Shawna A.; Smith, James N.M.; Boutin, Stan
doi: 10.1111/j.1600-1706.2009.17786.xpmid: N/A
Nest predation has been used to explain aspects of avian ecology ranging from nest site selection to population declines. Many arguments rely on specific assumptions regarding how predators find nests, yet these predatory mechanisms remain largely untested. Here we combine artificial nest experiments with behavioural observations of individual red squirrels Tamiasciurus hudsonicus to differentiate between two common hypotheses: predation is incidental versus learned. Specifically, we tested: 1) whether nest survival could be explained solely by a squirrel's activity patterns or habitat use, as predicted if predation was incidental; or 2) if predation increased as a squirrel gained experience preying on a nest, as predicted if predation was learned. We also monitored squirrel activity after predation to test for evidence of two search mechanisms: area‐restricted searching and use of microhabitat search images. Contrary to incidental predation and in support of learning, squirrels did not find nests faster in areas with high use (e.g. forest edges). Instead, survival of artificial nests was strongly related to a squirrel's prior experience preying on artificial nests. Experience reduced nest survival times by over half and increased predation rates by 150–200%. Squirrels returned to and doubled their activity at the site of a previously preyed on nest. However, neither area‐restricted searching nor microhabitat search images can explain how squirrels located artificial nests more readily with experience. Instead, squirrels likely used cues associated with the nests or eggs themselves. Learning implies that squirrels could be increasingly effective predators as the density or profitability of nests increases. Our results add support to the view that nest predation is complex and broadly influenced (e.g. by predator experience, motivation), and is unlikely to be predicted consistently by simple relationships with predator activity, abundance or habitat.
Leaf mines as visual defensive signals to herbivoresYamazaki, Kazuo
doi: 10.1111/j.1600-0706.2009.18300.xpmid: N/A
Leaf‐mining insects produce conspicuous and distinct leaf mines on various types of plant leaves. The diversity of leaf‐mine morphology has typically been explained by several factors, such as selective feeding on plant tissues, improvement of microclimate, faecal disposal, reduction in the efficiency of parasitoid search behaviour and leafminer phylogeny. Although these factors are certainly associated with mining patterns, masking the mines, rather than making them conspicuous, appears to be more advantageous for deterring parasitoids and predators of leafminers. However, here, I propose that prominent leaf mines may serve to signal or cue herbivores to avoid feeding on the mined leaves. Because most leafminers are sessile and complete their development within a single leaf, herbivory of mined leaves is detrimental to leafminer survival. Other herbivores appear to avoid consuming mined leaves for a variety of reasons: leaf mines mimic leaf variegation or mottling; mined leaves induce chemical and physical defences against herbivores; and leaf mines mimic fungal infection, animal excrement, and necrosed plant tissues. Hence, natural selection may have favoured leafminers that produce conspicuous mines because of the increased survival and fecundity of thereby reducing herbivory on mined leaves.
To be or not to be what you eat: regulation of stoichiometric homeostasis among autotrophs and heterotrophsPersson, Jonas; Fink, Patrick; Goto, Akira; Hood, James M.; Jonas, Jayne; Kato, Satoshi
doi: 10.1111/j.1600-0706.2009.18545.xpmid: N/A
Homeostasis of element composition is one of the central concepts of ecological stoichiometry. In this context, homeostasis is the resistance to change of consumer body composition in response to the chemical composition of consumer's food. To simplify theoretical analysis, it has generally been assumed that autotrophs exhibit flexibility in their composition, while heterotrophs are confined to a constant (strictly homeostatic) body composition. Yet, recent studies suggest that heterotrophs are not universally strictly homeostatic. We examined the degree to which autotrophs and heterotrophs regulate stoichiometric homeostasis (P:C, N:C, N:P, or %P and %N). We conducted a quantitative review and meta‐analysis using 132 datasets extracted from 57 literature sources which examined the dependence of organismal stoichiometry on resource stoichiometry. Among individual datasets, there was a wide range of responses from strictly homeostatic to non‐homeostatic. Even within heterotrophic organisms, varying levels of homeostasis were observed. Comparing the degree of homeostasis between organisms based on large‐scale habitat types using meta‐analysis indicated some significant differences between groups. For example, aquatic macroinvertebrates were significantly more homeostatic in terms of P:C than terrestrial invertebrates. Our meta‐analysis also confirmed that, with regard to N:P, heterotrophs are significantly more homeostatic than autotrophs. Furthermore, our analysis indicated that the homeostasis parameter 1/H, despite being a potentially useful predictive metric, has to be utilized with caution since it oversimplifies some important aspects of the responses of organisms to elemental imbalances. This critical evaluation of stoichiometric homeostasis contributes to a better understanding of many food‐web interactions, which are commonly driven by elemental imbalances between consumers and their resources.
Integrating elements and energy through the metabolic dependencies of gross growth efficiency and the threshold elemental ratioDoi, Hideyuki; Cherif, Mehdi; Iwabuchi, Tsubasa; Katano, Izumi; Stegen, James C.; Striebel, Maren
doi: 10.1111/j.1600-0706.2009.18540.xpmid: N/A
Metabolic theory proposes that individual growth is governed through the mass‐ and temperature‐dependence of metabolism, and ecological stoichiometry posits that growth is maximized at consumer‐specific optima of resource elemental composition. A given consumer's optimum, the threshold elemental ratio (TER), is proportional to the ratio of its maximum elemental gross growth efficiencies (GGEs). GGE is defined by the ratio of metabolism‐dependent processes such that GGEs should be independent of body mass and temperature. Understanding the metabolic‐dependencies of GGEs and TERs may open the path towards a theoretical framework integrating the flow of energy and chemical elements through ecosystems. However, the mass and temperature scaling of GGEs and TERs have not been broadly evaluated. Here, we use data from 95 published studies to evaluate these metabolic‐dependencies for C, N and P from unicells to vertebrates. We show that maximum GGEs commonly decline as power functions of asymptotic body mass and exponential functions of temperature. The rates of change in maximum GGEs with mass and temperature are relatively slow, however, suggesting that metabolism may not causally influence maximum GGEs. We additionally derived the theoretical expectation that the TER for C:P should not vary with body mass and this was supported empirically. A strong linear relationship between carbon and nitrogen GGEs further suggests that variation in the TER for C:N should be due to variation in consumer C:N. In general we show that GGEs may scale with metabolic rate, but it is unclear if there is a causal link between metabolism and GGEs. Further integrating stoichiometry and metabolism will provide better understanding of the processes governing the flow of energy and elements from organisms to ecosystems.
Sex in a material world: why the study of sexual reproduction and sex‐specific traits should become more nutritionally‐explicitMorehouse, Nathan I.; Nakazawa, Takefumi; Booher, Christina M.; Jeyasingh, Punidan D.; Hall, Matthew D.
doi: 10.1111/j.1600-0706.2009.18569.xpmid: N/A
Recent advances in nutritional ecology, particularly arising from Ecological Stoichiometry and the Geometric Framework for nutrition, have resulted in greater theoretical coherence and increasingly incisive empirical methodologies that in combination allow for the consideration of nutrient‐related processes at many levels of biological complexity. However, these advances have not been consistently integrated into the study of sexual differences in reproductive investment, despite contemporary emphasis on the material costs associated with sexually selected traits (e.g. condition‐dependence of exaggerated ornaments). Nutritional ecology suggests that material costs related to sex‐specific reproductive traits should be linked to quantifiable underlying differences in the relationship between individuals of each sex and their foods. Here, we argue that applying nutritionally‐explicit thought to the study of sexual reproduction should both deepen current understanding of sex‐specific phenomena and broaden the tractable frontiers of sexual selection research. In support of this general argument, we examine the causes and consequences of sex‐specific nutritional differences, from food selection and nutrient processing to sex‐specific reproductive traits. At each level of biological organization, we highlight how a nutritionally‐explicit perspective may provide new insights and help to identify new directions. Based on predictions derived at the individual level, we then consider how sex‐specific nutrient limitation might influence population growth, and thus potentially broader patterns of life history evolution, using a simple population dynamics model. We conclude by highlighting new avenues of research that may be more accessible from this integrative perspective.
Can ecological stoichiometry help explain patterns of biological invasions?González, Angélica L.; Kominoski, John S.; Danger, Michael; Ishida, Seiji; Iwai, Noriko; Rubach, Anja
doi: 10.1111/j.1600-0706.2009.18549.xpmid: N/A
Several mechanisms for biological invasions have been proposed, yet to date there is no common framework that can broadly explain patterns of invasion success among ecosystems with different resource availabilities. Ecological stoichiometry (ES) is the study of the balance of energy and elements in ecological interactions. This framework uses a multi‐nutrient approach to mass‐balance models, linking the biochemical composition of organisms to their growth and reproduction, which consequently influences ecosystem structure and functioning. We proposed a conceptual model that integrates hypotheses of biological invasions within a framework structured by fundamental principles of ES. We then performed meta‐analyses to compare the growth and production performances of native and invasive organisms under low‐ and high‐nutrient conditions in terrestrial and aquatic ecosystems. Growth and production rates of invasive organisms (plants and invertebrates) under both low‐ and high‐nutrient availability were generally larger than those of natives. Nevertheless, native plants outperformed invasives in aquatic ecosystems under low‐nutrient conditions. We suggest several distinct stoichiometry‐based mechanisms to explain invasion success in low‐ versus high‐nutrient conditions; low‐nutrient conditions: higher resource‐use efficiency (RUE; C:nutrient ratios), threshold elemental ratios (TERs), and trait plasticity (e.g. ability of an organism to change its nutrient requirements in response to varying nutrient environmental supply); high‐nutrient conditions: higher growth rates and reproductive output related to lower tissue C:nutrient ratios, and increased trait plasticity. Interactions of mechanisms may also yield synergistic effects, whereby nutrient enrichment and enemy release have a disproportionate effect on invasion success. To that end, ES provides a framework that can help explain how chemical elements and energy constrain key physiological and ecological processes, which can ultimately determine the success of invasive organisms.
Like moths to a street lamp: exaggerated animal densities in plot‐level global change field experimentsMoise, Eric R. D.; Henry, Hugh A. L.
doi: 10.1111/j.1600-0706.2009.18343.xpmid: N/A
Many recent field experiments have examined plant responses to global change factors such as climate warming, elevated atmospheric CO2, increased nitrogen addition or altered precipitation simulated at the plot level, yet the mechanisms underlying these responses can be difficult to isolate. One concern has been that the infrastructure used in these experiments can restrict the access of influential herbivores, detritivores or pollinators to the plots, and the absence of these animals is confounded with the treatment effects. However, in this paper we describe why free access by animals to experimental plots does not ensure realistic animal densities in response to global change treatments. On the contrary, much like moths swarm around streetlamps, animals that prefer the local conditions in treated plots may congregate at artificially high densities, or conversely, those that are repelled by the treatments may choose to avoid them. Therefore, animal densities or herbivore damage in the plots of global change experiments may grossly exaggerate or underestimate the contributions of animals to primary productivity or plant species composition under future environmental conditions. We describe how these potential animal congregation and avoidance artifacts may have been overlooked in the interpretation of results from many plot‐level global change field experiments. We also provide suggestions for how to best interpret the results of these experiments and how to isolate the effects of animal density artifacts.
Clonal mobility and its implications for spatio‐temporal patterns of plant communities: what do we need to know next?Zobel, Martin; Moora, Mari; Herben, Tomáš
doi: 10.1111/j.1600-0706.2010.18296.xpmid: N/A
Patterns of clonal growth and their controls on the level of individuals have been studied thoroughly, but little is known about the actual clonal mobility of plant individuals in vegetation and about its role in generating vegetation patterns and influencing species coexistence. Current evidence shows that communities are composed of spatially nonmobile ‘matrix‐forming species’ and mobile ‘inter‐matrix’ species, while local between‐species variation in clonal mobility has been shown to be positively correlated to small‐scale richness. We identify two major gaps in the knowledge. (1) Clonal mobility has a strong species‐specific component, but the existing information is mainly qualitative and describes the potential mobility of species the best. Also, species may respond by their clonal growth in a plastic way to some environmental stimuli, such as neighbors or abiotic environment, but this data comes almost exclusively from artificial conditions. We know very little of the actual spatial mobility of clonal plant individuals in the field and of the factors that determine it. (2) Theoretical research indicates that localized dispersal plays prime role in determination of community structure. While clonal mobility shares many important features with the seed dispersal, it also shows important differences to it, such as in dispersal kernel (non‐monotonic in clonal dispersal), role of microsite limitation, and role of plasticity. We have little information how systematic are these differences, and whether these differences in dispersal can play any role in shaping community dynamics. We conclude that clonal mobility has an important role in structuring plant communities in a small scale and propose further studies to address specific mechanisms, as well as community context of evolution of clonality.