Rethinking megafauna.

Concern for megafauna is increasing among scientists and non-scientists. Many studies have emphasized that megafauna play prominent ecological roles and provide important ecosystem services to humanity. But, what precisely are 'megafauna'? Here, we critically assess the concept of megafauna and propose a goal-oriented framework for megafaunal research. First, we review definitions of megafauna and analyse associated terminology in the scientific literature. Second, we conduct a survey among ecologists and palaeontologists to assess the species traits used to identify and define megafauna. Our review indicates that definitions are highly dependent on the study ecosystem and research question, and primarily rely on ad hoc size-related criteria. Our survey suggests that body size is crucial, but not necessarily sufficient, for addressing the different applications of the term megafauna. Thus, after discussing the pros and cons of existing definitions, we propose an additional approach by defining two function-oriented megafaunal concepts: 'keystone megafauna' and 'functional megafauna', with its variant 'apex megafauna'. Assessing megafauna from a functional perspective could challenge the perception that there may not be a unifying definition of megafauna that can be applied to all eco-evolutionary narratives. In addition, using functional definitions of megafauna could be especially conducive to cross-disciplinary understanding and cooperation, improvement of conservation policy and practice, and strengthening of public perception. As megafaunal research advances, we encourage scientists to unambiguously define how they use the term 'megafauna' and to present the logic underpinning their definition.

Variation in herbivore space use: comparing two savanna ecosystems with different anthrax outbreak patterns in southern Africa.

BACKGROUND: The distribution of resources can affect animal range sizes, which in turn may alter infectious disease dynamics in heterogenous environments. The risk of pathogen exposure or the spatial extent of outbreaks may vary with host range size. This study examined the range sizes of herbivorous anthrax host species in two ecosystems and relationships between spatial movement behavior and patterns of disease outbreaks for a multi-host environmentally transmitted pathogen. METHODS: We examined range sizes for seven host species and the spatial extent of anthrax outbreaks in Etosha National Park, Namibia and Kruger National Park, South Africa, where the main host species and outbreak sizes differ. We evaluated host range sizes using the local convex hull method at different temporal scales, within-individual temporal range overlap, and relationships between ranging behavior and species contributions to anthrax cases in each park. We estimated the spatial extent of annual anthrax mortalities and evaluated whether the extent was correlated with case numbers of a given host species. RESULTS: Range size differences among species were not linearly related to anthrax case numbers. In Kruger the main host species had small range sizes and high range overlap, which may heighten exposure when outbreaks occur within their ranges. However, different patterns were observed in Etosha, where the main host species had large range sizes and relatively little overlap. The spatial extent of anthrax mortalities was similar between parks but less variable in Etosha than Kruger. In Kruger outbreaks varied from small local clusters to large areas and the spatial extent correlated with case numbers and species affected. Secondary host species contributed relatively few cases to outbreaks; however, for these species with large range sizes, case numbers positively correlated with outbreak extent. CONCLUSIONS: Our results provide new information on the spatiotemporal structuring of ranging movements of anthrax host species in two ecosystems. The results linking anthrax dynamics to host space use are correlative, yet suggest that, though partial and proximate, host range size and overlap may be contributing factors in outbreak characteristics for environmentally transmitted pathogens.

Black rhinoceros (Diceros bicornis) natural diets: comparing iron levels across seasons and geographical locations.

Although excessive iron storage in black rhinoceros (Diceros bicornis) has been a cause for continuous concern over the last four decades and differences in the iron content of diet items fed in captivity and in the wild have been documented, no reports exist on the iron content of the total diet ingested by free-ranging animals. Here, the results of field studies using backtracking to record the ingested diets of black rhinoceros from three habitats across three seasons are reported. Levels of iron and of condensed tannins, which might reduce iron availability, averaged at 91 +/- 41 ppm dry matter and 3.0 +/- 1.0% dry matter, respectively, across all habitats and seasons. Although geographic and seasonal variation was significant, these differences are of a much lower magnitude than differences between the averages of these diets and those fed to black rhinoceros in captivity. The results can provide guidelines for the iron content of diets designed for black rhinoceros and suggest that the effect of tannins in these species should be further investigated.

Accommodating environmental variation in population models: metaphysiological biomass loss accounting.

1. There is a pressing need for population models that can reliably predict responses to changing environmental conditions and diagnose the causes of variation in abundance in space as well as through time. In this 'how to' article, it is outlined how standard population models can be modified to accommodate environmental variation in a heuristically conducive way. This approach is based on metaphysiological modelling concepts linking populations within food web contexts and underlying behaviour governing resource selection. Using population biomass as the currency, population changes can be considered at fine temporal scales taking into account seasonal variation. Density feedbacks are generated through the seasonal depression of resources even in the absence of interference competition. 2. Examples described include (i) metaphysiological modifications of Lotka-Volterra equations for coupled consumer-resource dynamics, accommodating seasonal variation in resource quality as well as availability, resource-dependent mortality and additive predation, (ii) spatial variation in habitat suitability evident from the population abundance attained, taking into account resource heterogeneity and consumer choice using empirical data, (iii) accommodating population structure through the variable sensitivity of life-history stages to resource deficiencies, affecting susceptibility to oscillatory dynamics and (iv) expansion of density-dependent equations to accommodate various biomass losses reducing population growth rate below its potential, including reductions in reproductive outputs. Supporting computational code and parameter values are provided. 3. The essential features of metaphysiological population models include (i) the biomass currency enabling within-year dynamics to be represented appropriately, (ii) distinguishing various processes reducing population growth below its potential, (iii) structural consistency in the representation of interacting populations and (iv) capacity to accommodate environmental variation in space as well as through time. Biomass dynamics provide a common currency linking behavioural, population and food web ecology. 4. Metaphysiological biomass loss accounting provides a conceptual framework more conducive for projecting and interpreting the population consequences of climatic shifts and human transformations of habitats than standard modelling approaches.

Contrasting capabilities of two ungulate species to cope with extremes of aridity.

Southern Africa is expected to experience increased frequency and intensity of droughts through climate change, which will adversely affect mammalian herbivores. Using bio-loggers, we tested the expectation that wildebeest (Connochaetes taurinus), a grazer with high water-dependence, would be more sensitive to drought conditions than the arid-adapted gemsbok (Oryx gazella gazella). The study, conducted in the Kalahari, encompassed two hot-dry seasons with similar ambient temperatures but differing rainfall patterns during the preceding wet season. In the drier year both ungulates selected similar cooler microclimates, but wildebeest travelled larger distances than gemsbok, presumably in search of water. Body temperatures in both species reached lower daily minimums and higher daily maximums in the drier season but daily fluctuations were wider in wildebeest than in gemsbok. Lower daily minimum body temperatures displayed by wildebeest suggest that wildebeest were under greater nutritional stress than gemsbok. Moving large distances when water is scarce may have compromised the energy balance of the water dependent wildebeest, a trade-off likely to be exacerbated with future climate change.

How free-ranging ungulates with differing water dependencies cope with seasonal variation in temperature and aridity.

Large mammals respond to seasonal changes in temperature and precipitation by behavioural and physiological flexibility. These responses are likely to differ between species with differing water dependencies. We used biologgers to contrast the seasonal differences in activity patterns, microclimate selection, distance to potential water source and body temperature of the water-independent gemsbok (Oryx gazella gazella) and water-dependent blue wildebeest (Connochaetes taurinus), free-living in the arid Kalahari region of Botswana. Gemsbok were more active nocturnally during the hot seasons than in the cold-dry season, while wildebeest showed no seasonal difference in their nocturnal activity level. Both species similarly selected shaded microclimates during the heat of the day, particularly during the hot seasons. Wildebeest were further than 10 km from surface water 30% or more of the time, while gemsbok were frequently recorded >20 km from potential water sources. In general, both species showed similar body temperature variation with high maximum 24-h body temperature when conditions were hot and low minimum 24-h body temperatures when conditions were dry, resulting in the largest amplitude of 24-h body temperature rhythm during the hot-dry period. Wildebeest thus coped almost as well as gemsbok with the fairly typical seasonal conditions that occurred during our study period. They do need to access surface water and may travel long distances to do so when local water sources become depleted during drought conditions. Thus, perennial water sources should be provided judiciously and only where essential.

Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator-prey web.

Shifting prey selection has been identified as a mechanism potentially regulating predator-prey interactions, but it may also lead to different outcomes, especially in more complex systems with multiple prey species available. We assessed changing prey selection by lions, the major predator for 12 large herbivore species in South Africa's Kruger National Park. The database was provided by records of found carcasses ascribed to kills by lions assembled over 70 years, coupled with counts of changing prey abundance extending over 30 years. Wildebeest and zebra constituted the most favored prey species during the early portion of the study period, while selection for buffalo rose in the south of the park after a severe drought increased their vulnerability. Rainfall had a negative influence on the proportional representation of buffalo in lion kills, but wildebeest and zebra appeared less susceptible to being killed under conditions of low rainfall. Selection by lions for alternative prey species, including giraffe, kudu, waterbuck, and warthog, was influenced by the changing relative abundance and vulnerability of the three principal prey species. Simultaneous declines in the abundance of rarer antelope species were associated with a sharp increase in selection for these species at a time when all three principal prey species were less available. Hence shifting prey selection by lions affected the dynamics of herbivore populations in different ways: promoting contrasting responses by principal prey species to rainfall variation, while apparently being the main cause of sharp declines by alternative prey species under certain conditions. Accordingly, adaptive responses by predators, to both the changing relative abundance of the principal prey species, and other conditions affecting the relative vulnerability of various species, should be taken into account to understand the interactive dynamics of multispecies predator-prey webs.

Predator-prey size relationships in an African large-mammal food web.

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.

Ecological and evolutionary legacy of megafauna extinctions.

For hundreds of millions of years, large vertebrates (megafauna) have inhabited most of the ecosystems on our planet. During the late Quaternary, notably during the Late Pleistocene and the early Holocene, Earth experienced a rapid extinction of large, terrestrial vertebrates. While much attention has been paid to understanding the causes of this massive megafauna extinction, less attention has been given to understanding the impacts of loss of megafauna on other organisms with whom they interacted. In this review, we discuss how the loss of megafauna disrupted and reshaped ecological interactions, and explore the ecological consequences of the ongoing decline of large vertebrates. Numerous late Quaternary extinct species of predators, parasites, commensals and mutualistic partners were associated with megafauna and were probably lost due to their strict dependence upon them (co-extinctions). Moreover, many extant species have megafauna-adapted traits that provided evolutionary benefits under past megafauna-rich conditions, but are now of no or limited use (anachronisms). Morphological evolution and behavioural changes allowed some of these species partially to overcome the absence of megafauna. Although the extinction of megafauna led to a number of co-extinction events, several species that likely co-evolved with megafauna established new interactions with humans and their domestic animals. Species that were highly specialized in interactions with megafauna, such as large predators, specialized parasites, and large commensalists (e.g. scavengers, dung beetles), and could not adapt to new hosts or prey were more likely to die out. Partners that were less megafauna dependent persisted because of behavioural plasticity or by shifting their dependency to humans via domestication, facilitation or pathogen spill-over, or through interactions with domestic megafauna. We argue that the ongoing extinction of the extant megafauna in the Anthropocene will catalyse another wave of co-extinctions due to the enormous diversity of key ecological interactions and functional roles provided by the megafauna.

New insights into the physiology of natural foraging.

The purpose of this symposium was to examine how foraging physiology is studied in the field across a diversity of species and habitats. While field studies are constrained by the relatively poor ability to control the experiment, the natural variability in both the environment and animal behavior provides insights into adaptation to change that are usually not tested in the laboratory. Talks in this session examined how foraging energy (both costs and gains) is partitioned over time. "Time," in this case, ranged from evolutionary time (how different animals are designed to most efficiently forage), to long, lifetime periods (development of foraging ability and growth), to short-duration feeding bouts, and ultimately to the minutes to hours following ingestion (metabolic and biochemical changes). From this diversity, two core themes emerged: that foraging strategies and behaviors are limited by physiology and biochemical processes and that time plays a central role in the organization of foraging behaviors and the physiological processes that underlie those behaviors.

Evidence of reduced individual heterogeneity in adult survival of long-lived species.

The canalization hypothesis postulates that the rate at which trait variation generates variation in the average individual fitness in a population determines how buffered traits are against environmental and genetic factors. The ranking of a species on the slow-fast continuum - the covariation among life-history traits describing species-specific life cycles along a gradient going from a long life, slow maturity, and low annual reproductive output, to a short life, fast maturity, and high annual reproductive output - strongly correlates with the relative fitness impact of a given amount of variation in adult survival. Under the canalization hypothesis, long-lived species are thus expected to display less individual heterogeneity in survival at the onset of adulthood, when reproductive values peak, than short-lived species. We tested this life-history prediction by analysing long-term time series of individual-based data in nine species of birds and mammals using capture-recapture models. We found that individual heterogeneity in survival was higher in species with short-generation time (< 3 years) than in species with long generation time (> 4 years). Our findings provide the first piece of empirical evidence for the canalization hypothesis at the individual level from the wild.

Identifying Space Use at Foraging Arena Scale within the Home Ranges of Large Herbivores.

An intermediate spatiotemporal scale of food procurement by large herbivores is evident within annual or seasonal home ranges. It takes the form of settlement periods spanning several days or weeks during which foraging activity is confined to spatially discrete foraging arenas, separated by roaming interludes. Extended by areas occupied for other activities, these foraging arenas contribute towards generating the home range structure. We delineated and compared the foraging arenas exploited by two African large herbivores, sable antelope (a ruminant) and plains zebra (a non-ruminant), using GPS-derived movement data. We developed a novel approach to specifically delineate foraging arenas based on local change points in distance relative to adjoining clusters of locations, and compared its output with modifications of two published methods developed for home range estimation and residence time estimation respectively. We compared how these herbivore species responded to seasonal variation in food resources and how they differed in their spatial patterns of resource utilization. Sable antelope herds tended to concentrate their space use locally, while zebra herds moved more opportunistically over a wider set of foraging arenas. The amalgamated extent of the foraging arenas exploited by sable herds amounted to 12-30 km2, compared with 22-100 km2 for the zebra herds. Half-day displacement distances differed between settlement periods and roaming interludes, and zebra herds generally shifted further over 12h than sable herds. Foraging arenas of sable herds tended to be smaller than those of zebra, and were occupied for period twice as long, and hence exploited more intensively in days spent per unit area than the foraging arenas of zebra. For sable both the intensity of utilization of foraging arenas and proportion of days spent in foraging arenas relative to roaming interludes declined as food resources diminished seasonally, while zebra showed no seasonal variation in these metrics. Identifying patterns of space use at foraging arena scale helps reveal mechanisms generating the home range extent, and in turn the local population density. Thereby it helps forge links between behavioural ecology, movement ecology and population ecology.

Coping with spatial heterogeneity and temporal variability in resources and risks: adaptive movement behaviour by a large grazing herbivore.

Movement is a key mean for mobile species to cope with heterogeneous environments. While in herbivorous mammals large-scale migration has been widely investigated, fine-scale movement responses to local variations in resources and predation risk remain much less studied, especially in savannah environments. We developed a novel approach based on complementary movement metrics (residence time, frequency of visits and regularity of visits) to relate movement patterns of a savannah grazer, the blue wildebeest Connochaetes taurinus, to fine-scale variations in food availability, predation risk and water availability in the Kruger National Park, South Africa. Wildebeests spent more time in grazing lawns where the grass is of higher quality but shorter than in seep zones, where the grass is of lower quality but more abundant. Although the daily distances moved were longer during the wet season compared to the dry season, the daily net displacement was lower, and the residence time higher, indicating a more frequent occurrence of area-concentred searching. In contrast, during the late dry season the foraging sessions were more fragmented and wildebeests moved more frequently between foraging areas. Surprisingly, predation risk appeared to be the second factor, after water availability, influencing movement during the dry season, when resources are limiting and thus expected to influence movement more. Our approach, using complementary analyses of different movement metrics, provided an integrated view of changes in individual movement with varying environmental conditions and predation risk. It makes it possible to highlight the adaptive behavioral decisions made by wildebeest to cope with unpredictable environmental variations and provides insights for population conservation.

How Rainfall Variation Influences Reproductive Patterns of African Savanna Ungulates in an Equatorial Region Where Photoperiod Variation Is Absent.

In high temperate latitudes, ungulates generally give birth within a narrow time window when conditions are optimal for offspring survival in spring or early summer, and use changing photoperiod to time conceptions so as to anticipate these conditions. However, in low tropical latitudes day length variation is minimal, and rainfall variation makes the seasonal cycle less predictable. Nevertheless, several ungulate species retain narrow birth peaks under such conditions, while others show births spread quite widely through the year. We investigated how within-year and between-year variation in rainfall influenced the reproductive timing of four ungulate species showing these contrasting patterns in the Masai Mara region of Kenya. All four species exhibited birth peaks during the putative optimal period in the early wet season. For hartebeest and impala, the birth peak was diffuse and offspring were born throughout the year. In contrast, topi and warthog showed a narrow seasonal concentration of births, with conceptions suppressed once monthly rainfall fell below a threshold level. High rainfall in the previous season and high early rains in the current year enhanced survival into the juvenile stage for all the species except impala. Our findings reveal how rainfall variation affecting grass growth and hence herbivore nutrition can govern the reproductive phenology of ungulates in tropical latitudes where day length variation is minimal. The underlying mechanism seems to be the suppression of conceptions once nutritional gains become insufficient. Through responding proximally to within-year variation in rainfall, tropical savanna ungulates are less likely to be affected adversely by the consequences of global warming for vegetation phenology than northern ungulates showing more rigid photoperiodic control over reproductive timing.

Ecology of grazing lawns in Africa.

Grazing lawns are a distinct grassland community type, characterised by short-stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co-evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low-biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall-grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short-grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall-grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.

Circularity in linear programming models of optimal diet.

The linear programming model of optimal diet for herbivores has been criticized for being biologically unrealistic, for being too successful given statistical realities, and for being circular. I try to clarify the issue of circularity. Circularity arises if constraint lines are estimated from average values for governing parameters, when the assumed constraints are not effective. This may occur (1) under benign season conditions when consumers do not maximize their food intake, because of costs associated with food processing and storage, or (2) when an unidentified constraint limits intake. To evaluate hypotheses about the factors controlling diet composition, it must be shown that (1) consumers respond appropriately to variation in the parameters controlling constraint settings, and (2) the assumed constraints are close to their maximum (or minimum) settings.

Effects of plant spinescence on large mammalian herbivores.

Plant thorns and spines had these effects on the feeding behaviour of the three species of browsing ungulate that we studied, kudu, impala and domestic goats: (i) bite sizes were restricted, in most cases to single leaves or leaf clusters; (ii) hooked thorns retarded biting rates; (iii) the acceptability of those plant species offering small leaf size in conjunction with prickles was lower, at least for the kudus, than those of other palatable plant species; (iv) the inhibitory effect of prickles on feeding was much less for the smaller impalas and goats than for the larger kudus; (v) from certain hook-thorned species the kudus bit off shoot ends despite their prickles; (vi) for certain straight-thorned species the kudus compensated partially for the slow eating rates obtained by extending their feeding durations per encounter. Most spinescent species were similar in their acceptability to the ungulates to unarmed palatable species, despite higher crude protein contents in their foliage than the latter. Such structural features furthermore reduce the tissue losses incurred by plants per encounter by a large ungulate herbivore, by restricting the eating rates that the animals obtain. In this way prickles function to restrict foliage losses to large herbivores below the levels that might otherwise occur.

Foliage acceptability to browsing ruminants in relation to seasonal changes in the leaf chemistry of woody plants in a South African savanna.

We investigated seasonal changes in food selection by hand-reared kudus and impalas in savanna vegetation in northern Transvaal, South Africa. The acceptability of the leaves of woody plants to these animals was compared with leaf concentrations of nutrients, fibre components and old leaf phenophases. No consistently significant correlation was found between acceptability and any single chemical factor. Based on an a priori palatability classification, discriminant function analysis separated relatively palatable species from unpalatable species in terms of a linear combination of protein and condensed tannin concentrations. The high acceptability of certain otherwise unpalatable species during the new leaf phenophase was related to elevation of protein levels relative to condensed tannin contents. Species were added to the diet during the dry season approximately in the order of their relative protein-condensed tannin difference.

Inter-specific interactions linking predation and scavenging in terrestrial vertebrate assemblages.

Predation and scavenging have been classically understood as independent processes, with predator-prey interactions and scavenger-carrion relationships occurring separately. However, the mere recognition that most predators also scavenge at variable rates, which has been traditionally ignored in food-web and community ecology, leads to a number of emergent interaction routes linking predation and scavenging. The general goal of this review is to draw attention to the main inter-specific interactions connecting predators (particularly, large mammalian carnivores), their live prey (mainly ungulates), vultures and carrion production in terrestrial assemblages of vertebrates. Overall, we report an intricate network of both direct (competition, facilitation) and indirect (hyperpredation, hypopredation) processes, and provide a conceptual framework for the future development of this promising topic in ecological, evolutionary and biodiversity conservation research. The classic view that scavenging does not affect the population dynamics of consumed organisms is questioned, as multiple indirect top-down effects emerge when considering carrion and its facultative consumption by predators as fundamental and dynamic components of food webs. Stimulating although challenging research opportunities arise from the study of the interactions among living and detrital or non-living resource pools in food webs.

Consumer-resource dynamics: quantity, quality, and allocation.

BACKGROUND: The dominant paradigm for modeling the complexities of interacting populations and food webs is a system of coupled ordinary differential equations in which the state of each species, population, or functional trophic group is represented by an aggregated numbers-density or biomass-density variable. Here, using the metaphysiological approach to model consumer-resource interactions, we formulate a two-state paradigm that represents each population or group in a food web in terms of both its quantity and quality. METHODOLOGY AND PRINCIPAL FINDINGS: The formulation includes an allocation function controlling the relative proportion of extracted resources to increasing quantity versus elevating quality. Since lower quality individuals senescence more rapidly than higher quality individuals, an optimal allocation proportion exists and we derive an expression for how this proportion depends on population parameters that determine the senescence rate, the per-capita mortality rate, and the effects of these rates on the dynamics of the quality variable. We demonstrate that oscillations do not arise in our model from quantity-quality interactions alone, but require consumer-resource interactions across trophic levels that can be stabilized through judicious resource allocation strategies. Analysis and simulations provide compelling arguments for the necessity of populations to evolve quality-related dynamics in the form of maternal effects, storage or other appropriate structures. They also indicate that resource allocation switching between investments in abundance versus quality provide a powerful mechanism for promoting the stability of consumer-resource interactions in seasonally forcing environments. CONCLUSIONS/SIGNIFICANCE: Our simulations show that physiological inefficiencies associated with this switching can be favored by selection due to the diminished exposure of inefficient consumers to strong oscillations associated with the well-known paradox of enrichment. Also our results demonstrate how allocation switching can explain observed growth patterns in experimental microbial cultures and discuss how our formulation can address questions that cannot be answered using the quantity-only paradigms that currently predominate.