A historical perspective on drift foraging models for stream salmonids

Thirty years ago, Fausch (Can J Zool 62:441–451, 1984) proposed a simple model of optimal positions for drift-feeding salmonids in streams, whereby fish maximize their net energy intake (NEI) by selecting focal points in low water velocity near faster currents that deliver abundant drifting invertebrates. The theory was based on earlier observations in artificial and natural streams describing characteristics of salmonid positions and a conceptual model by Chapman (Am Nat 100:345–357, 1966). A test of this simple drift foraging model in a laboratory stream showed that the growth rate of juvenile trout and salmon increased with NEI, and that the rank of NEI at positions held by coho salmon (Oncorhynchus kisutch) correlated nearly perfectly with their rank in the dominance hierarchy. Fausch (1984) inferred from these findings that positions that optimize NEI, within the constraints of the dominance hierarchy, are the resource for which these stream salmonids compete. In turn, the model was used to test the effects of interspecific competition by coho salmon on the foraging positions held by brook trout (Salvelinus fontinalis) and brown trout (Salmo trutta), and these results were used to infer potential effects of the introduced salmon on resident trout in Lake Michigan tributaries. Though the goals for this model were originally modest, it was tested in the field and further refined by Hughes and Dill (Can J Fish Aquat Sci 47:2039–2048, 1990) and others. During the last 20 years, the general theory has been incorporated into other models, which have been applied widely to analyze salmonid distribution and abundance in streams and rivers and used for management and restoration of habitat and flow regimes to benefit these fishes.     

Plant phenotypic plasticity belowground: a phylogenetic perspective on root foraging trade-offs

Many plants proliferate roots in nutrient patches, presumably increasing nutrient uptake and plant fitness. Nutrient heterogeneity has been hypothesized to maintain community diversity because of a trade-off between the spatial extent over which plants forage (foraging scale) and their ability to proliferate roots precisely in nutrient patches (foraging precision). Empirical support for this hypothesis has been mixed, and some authors have suggested that interspecific differences in relative growth rate may be confounded with measurements of foraging precision. We collected previously published data from numerous studies of root foraging ability (foraging precision, scale, response to heterogeneity, and relative growth rate) and phylogenetic relationships for >100 plant species to test these hypotheses using comparative methods. Root foraging precision was phylogenetically and taxonomically conserved. Using a historical and phylogenetically independent contrast correlations, we found no evidence of a root foraging scale-precision trade-off, mixed support for a relative growth rate-precision relationship, and no support for the widespread assumption that foraging precision increases the benefit gained from growth in heterogeneous soil. Our understanding of the impacts of plant foraging precision and soil heterogeneity on plants and communities is less advanced than commonly believed, and we suggest several areas in which further research is needed.     

Incubation patterns in a central-place forager affect lifetime reproductive success: scaling of patterns from a foraging bout to a lifetime

Background

Long-lived seabirds face a conflict between current and lifelong reproductive success. During incubation shifts, egg neglect is sometimes necessary to avoid starvation, but may compromise the current reproductive attempt. However, factors underlying this decision process are poorly understood. We focus on the ancient murrelet, Synthliboramphus antiquus, an alcid with exceptionally long incubation shift lengths, and test the impact of environmental factors on incubation shift length in relation to reproductive success.

Methodology/Principal Findings

Using an information theoretic approach, we show that incubation shift length was a strong predictor of reproductive success for ancient murrelets at Reef Island, Haida Gwaii, British Columbia, Canada during the 2007 and 2008 breeding seasons. The most important factors explaining an individual''s shift length were egg size, wind speed and the length of the mate''s previous shift. Wind speed and tide height were the two most important factors for determining foraging behavior, as measured by dive frequency and depth.

Conclusions/Significance

Our study demonstrates that (i) species-specific reproductive strategies interact with environmental conditions such as wind speed to form multiple incubation patterns and (ii) maintaining regular incubation shifts is an essential component of reproductive success.     

Physiological tipping points in the relationship between foraging success and lifetime fitness of a long-lived mammal

Although anthropogenic change is often gradual, the impacts on animal populations may be precipitous if physiological processes create tipping points between energy gain, reproduction or survival. We use 25 years of behavioural, diet and demographic data from elephant seals to characterise their relationships with lifetime fitness. Survival and reproduction increased with mass gain during long foraging trips preceding the pupping seasons, and there was a threshold where individuals that gained an additional 4.8% of their body mass (26 kg, from 206 to 232 kg) increased lifetime reproductive success three-fold (from 1.8 to 4.9 pups). This was due to a two-fold increase in pupping probability (30% to 76%) and a 7% increase in reproductive lifespan (6.0 to 6.4 years). The sharp threshold between mass gain and reproduction may explain reproductive failure observed in many species and demonstrates how small, gradual reductions in prey from anthropogenic disturbance could have profound implications for animal populations.     

Natural 'poor start' does not increase mortality over the lifetime

Poor nutrition and other challenges during infancy can impose delayed costs, and it has been proposed that expression of costs during adulthood should involve increased mortality rather than reduced reproduction. Demonstrations of delayed costs come mostly from experimental manipulations of the diet and hormones of captive infants of short-lived species, and we know very little about how natural poor starts in life affect wild animals over their lifetimes. In the blue-footed booby, sibling conflict obliges younger brood members to grow up suffering aggressive subordination, food deprivation and elevated stress hormone, but surviving fledglings showed no deficit in reproduction over the first 5-10 years. A study of 7927 individuals from two-fledgling and singleton broods from 20 cohorts found no significant evidence of a higher rate of mortality nor a lower rate of recruitment in younger fledglings than in elder fledglings or singletons at any age over the 20 year lifespan. Development of boobies may be buffered against the three challenges of subordination. Experimental challenges to neonates that result in delayed costs have usually been more severe, more prolonged and more abruptly suspended, and it is unclear which natural situations they mimic.     

Honeyeater foraging: A test of optimal foraging theory

Honeyeaters (Meliphagidae) were observed foraging for nectar from Lambertia formosa inflorescences, each of which has seven flowers. The frequency distribution of numbers of flowers probed per visit to an inflorescence was found to be bimodal, with one peak at two and the other at seven. It is hypothesized that this frequency distribution results from a rule of departure from inflorescences that maximizes the net rate of energy gain. Patterns of nectar distribution were determined for a large sample of inflorescences. In addition the extent to which the honeyeaters re-probe flowers during a visit to an inflorescence was estimated. From these data and from field measurements of the times required by the honeyeaters to perform the various foraging behaviours, computer simulations of honeyeater foraging were constructed. These simulations led in turn to optimal frequency distributions of numbers of flowers probed per inflorescence that were bimodal but had peaks at 1 and 7 instead of 2 and 7. Although the observed and predicted behaviour were consequently similar, the difference between them was nevertheless significant. This difference could have been due to the birds' transient occupancy of the study area.     

Mixotrophy of a photosynthetic flagellate viewed from an optimal foraging perspective

Mixotrophy, a combination of phototrophic and phagotrophic nutrition, has been found in several classes of phytoplankton (Booras et al. 1988, Jones 2000) and appears to be a successful evolutionary strategy. Heterotrophic nutrition of phytoplankton has been suggested to be an important source of mineral nutrients (Nygaard and Tobiesen 1993). Potentially limiting mineral nutrients, particularly phosphorus (P), are often several orders of magnitude more concentrated in the biomass of food organisms of mixotrophs (e.g. in bacteria) than in the dissolved phase (Vadstein 2000). We used radioactive tracer experiments to show that the simultaneous uptake of P from dissolved inorganic and particular P sources by the marine phytoflagellate Chrysochromulina polylepis followed basic predictions of optimal foraging theory (Stephens and Krebs 1986). Chrysochromulina takes up its P rather unselectively from both bacterial P and dissolved P sources at low dissolved P concentrations, while it becomes more selective at higher dissolved inorganic P (DIP) concentrations. The onset of mixotrophic processes was dependent on DIP concentrations. These findings support the view of mixotrophy as a strategy of nutrient uptake in nutrient poor (oligotrophic) pelagic environments (Nygaard and Tobiesen 1993) and show that ideas of optimal foraging can be applied to unicellular organisms.     

The consequences of adult foraging success on the components of lifetime fitness in a semelparous,sit and wait predator

Summary We evaluated the role of adult foraging success in the lifetime fitness of female crab spidersMisumena vatia. Misumena are semelparous, sit and wait predators that hunt for insect prey on flowers, in this study primarily on inflorescences (umbels) of milkweedAsclepias syriaca. We used path analysis to integrate previously performed experimental and observational studies, thereby establishing the magnitude, correlations and causal relationships of key foraging and life history variables and their roles in lifetime fitness. A path proceeding from maternal hunting patch choice through maternal mass, clutch mass and number of dispersal-age young was the dominant element and explained a large part of the variation. Other paths that incorporated parasitism of the egg mass and predation of young leaving the nests made only small impacts on variation. No trade-offs were found, primarily because a single factor, maternal mass (a maternal effect) resulting from foraging success, provided major benefits for successive life history stages. Since differences in the numbers of eggs, egg loss and mortality at dispersal resulted almost entirely from differences in maternal mass, they are controlled by the maternal generation and, thus, are appropriately attributed to the lifetime fitnesses of the mothers, rather than to those of their offspring.     

The scale-precision trade-off in spacial resource foraging by plants: restoring perspective

BACKGROUND AND AIMS: From the results of a comparative study using quantitative standardized assays of the scale and precision of responses of root and shoot systems to resource patchiness, Campbell et al. (1991; Oecologia 87: 532-538) proposed a mechanism of species coexistence in herbaceous communities involving a dynamic equilibrium between, respectively, the coarse- and fine-scale foraging of dominant and subordinate species. The purpose of this paper is to reject a recent assertion that with respect to root systems the scale-precision hypothesis has been falsified. DISCUSSION AND CONCLUSION: Reference to the original papers confirms that the scope of the hypothesis was confined to circumstances (eg. mown meadows) where the vigour of potential dominants is restricted by intermittent removal of biomass. This qualification in the original hypothesis is a crucial omission from the meta-analysis conducted by Kembel and Cahill (2005; American Naturalist 166: 216-230). The original papers also contain examples that illustrate the operation of forms of selection that prevent the development of precise foraging below ground; these also appear to have escaped the attention of recent participants in this field of research.     

Linking foraging behavior to lifetime reproductive success for an insect parasitoid: adaptation to host distributions

European and American populations of the parasitoid Cotesiaglomerata show pronounced differences in foraging behavior acrossplants and leaves. This variation in spatial aspects of foragingbehavior was observed about 350 generations after the introductionof C. glomerata from Europe to North America. We used a simulationmodel to study how these behavioral differences affect lifetimereproductive success in environments that differ in the spatialdistribution of hosts. The preferred gregarious host Pierisbrassicae occurs in rare large clusters in Europe but is absentin North America. The solitary caterpillars of Pieris rapaeare negative binomially distributed across plants during summerin North America, whereas they are Poisson-distributed in Europe,and early and late in the season in North America. Simulationsshowed that the foraging strategy of American C. glomerata resultedin a higher lifetime reproductive success than did the strategyof European C. glomerata on a Poisson P. rapae distribution,but did not differ on the more clustered negative binomial distribution.American parasitoids spend less time on exploration flights,focusing on the exploitation of P. rapae patches. This suggeststhat C. glomerata has adapted to the North American environmentthrough the loss of exploration traits necessary for the locationof rare clusters of P. brassicae. Lifetime reproductive successof the European strategy was most sensitive to an increase inthe giving up time on infested leaves. This behavioral parameterwas more than twice as high in the American parasitoids comparedwith their European conspecifics.     

Behaviourally mediated indirect effects: interference competition increases predation mortality in foraging redshanks

1. The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an example of this. 2. We show that the foraging success of redshanks Tringa totanus (L.) foraging on active prey was highest in the front of flocks, whereas this was not the case for redshanks foraging on inactive prey. Also, when foraging on active prey, foraging success in a flock decreased as more birds passed through a patch, while overall foraging success was not lower on subsequent visits to the same patch. Thus, redshanks foraging on active prey suffered from interference competition, whereas this was not the case for redshanks foraging on inactive prey. 3. This interference competition led to differences in activity: redshanks attaining a lower foraging success had a higher walking rate. Greater activity was associated with wider flock spacing and shorter distances to cover, which has previously been shown to increase predation risk and mortality from sparrowhawks Accipiter nisus (L.). 4. We conclude that behavioural adaptations of prey species can lead to interference competition in foraging redshanks, and thus can affect their predation risk and mortality through increased activity. This study is one of the first to show how interference competition can be a mechanism for behaviourally mediated indirect effects, and provides further evidence for the suggestion that a single species occupying an intermediate trophic level may be simultaneously top-down controlled by a predator and bottom-up controlled by a behavioural response of its prey.     

Critical thermal limits in Mediterranean ant species: trade-off between mortality risk and foraging performance

1. In Mediterranean ant communities, a close relationship has been found between activity rhythm in the period of maximum activity and position in the dominance hierarchy: subordinate species are active during the day, when conditions are more severe, while dominants are active during the afternoon and the night.
2. Results obtained in this study confirmed that the species foraging at higher temperatures were closer to their critical thermal limits than the species foraging at lower temperatures.
3. This enabled two extreme strategies of foraging in relation to temperature to be distinguished: (1) heat-intolerant ant species behaved as risk-averse species, foraging at temperatures very far from their critical thermal limits; and (2) heat-tolerant ant species behaved as risk-prone species, foraging very near their critical thermal limits and running a high heat mortality risk.
4. Heat-tolerant species benefited from this strategy by having better foraging performance at high temperatures.
5. This wide range of thermal niches may be one reason why Mediterranean ant faunas are so diverse in the face of limited diversity in vegetation and habitat structure: the daily range of temperature may be sufficiently great to meet the requirement both of heat-adapted and cold-adapted species as well as a spectrum of intermediate forms.     

Size and temperature dependent foraging capacities and metabolism: consequences for winter starvation mortality in fish

The foraging related capacities, energy requirements and the ability of individuals to withstand starvation are strongly dependent on body size and temperature. In this study, we estimated size-dependent foraging rates and critical resource density (CRD) in small Arctic char ( Salvelinus alpinus ) under winter conditions and compared these with previous observations under summer conditions. We investigated if starvation mortality is size-dependent in the laboratory, and we assessed the potential for winter growth and the occurrence and size dependency of winter mortality both in a large scale pond experiment and in natural lakes. The efficiency of foraging on macroinvertebrates increased with size but was lower at 4 than 12°C, still CRD was lower at 4 than 12°C as metabolic rates decreased faster than foraging efficiency with temperature. When starved, small char died before large and at rates which suggest that YOY char need to feed during winter to avoid starvation. Results from both our pond experiment and field study indicate that survival of YOY char over winter is high, despite severe winter conditions, because YOY char are able to feed and grow during winter. In seasonal environments with declining resources, the size scaling and temperature dependency of foraging and metabolic demands may provide conditions which can favour either small or large individuals. This size advantage dichotomy relates to that larger individual's by having a higher CRD are more likely to start starving, but once resource levels are below CRD for all size classes, small individuals starve to death at a higher rate. Negative size-dependent winter mortality from starvation is suggested to be more pronounced in species that are not adapted to feed at low temperatures and in species feeding on zooplankton, since zooplankton abundance, in contrast to macroinvertebrate abundance, is generally low during winter.