Environmental determinants of food-chain length: a meta-analysis

Food-chain length is an important character of ecological communities that affects many of their functional aspects. Recently, an increasing number of studies have tested the effects of productivity, disturbance, or ecosystem size on food-chain length in a variety of natural systems. Here we conduct a formal meta-analysis to summarize findings from these empirical studies. We found significant positive mean effects of productivity and ecosystem size but no significant mean effect of disturbance on food-chain length. The strength of mean effect sizes was not significantly different between productivity and ecosystem size. These results lend general support to previous theories predicting the effect of productivity and ecosystem size, but fail to support the prediction that disturbance shortens food chains. In addition, our meta-analysis found that the effect sizes of primary studies were significantly heterogeneous for ecosystem size and disturbance, but not for productivity. This pattern might reflect that ecosystem size and disturbance can affect food-chain length through multiple different mechanisms, while productivity influences food-chain length in a simple manner through energy limitation.     

Hydrogeomorphology and river impoundment affect food-chain length of diverse Neotropical food webs

Food-chain length is a central characteristic of ecological communities that affects community structure and ecosystem function. What determines the length of food chains is not well resolved for most ecosystems. Herein, we examine environmental correlates of food-chain length based on the productivity hypothesis, compare food-chain lengths among aquatic ecosystem types and identify bi-directional effects of river impoundment on food-chain length in the Paraná River Basin of South America. Both temperature regime, a surrogate of productivity, and ecosystem type significantly affected food-chain length in independent analyses. However, when analyzed together, only ecosystem type explained significant variation in food-chain length. Food chains were longest in reservoirs, and shortest in high-gradient rivers. The proximate mechanism driving this pattern appears to be body-size ratios of primary consumers to apex predators, which differ among trophic pathways. Food chains based on phytoplankton production may have an additional size-structured link not present in food chains based on other basal sources such as detritus and algae. Hydrogeomorphology is the ultimate mechanism influencing food-chain length because it affects the relative importance of basal carbon sources supporting higher trophic levels, which through differences in the number of trophic links along the different size-structured pathways, appears to drive the observed patterns in food-chain length. We discuss a hypothesis of food-chain length that integrates energy flow and size-structure, facilitates inclusion of temporal dynamics and which is readily testable in both 'closed' and 'open' ecosystems.     

Proximate structural mechanisms for variation in food-chain length

Food-chain length is a central characteristic of ecological communities because of its strong influence on community structure and ecosystem function. While recent studies have started to better clarify the relationship between food-chain length and environmental gradients such as resource availability and ecosystem size, much less progress has been made in isolating the ultimate and proximate mechanisms that determine food-chain length. Progress has been slow, in part, because research has paid little attention to the proximate changes in food web structure that must link variation in food-chain length to the ultimate dynamic mechanism. Here we outline the structural mechanisms that determine variation in food-chain length. We explore the implications of these mechanisms for understanding how changes in food-web structure influence food-chain length using both an intraguild predation community model and data from natural ecosystems. The resulting framework provides the mechanisms for linking ultimate dynamic mechanisms to variation in food-chain length. It also suggests that simple linear food-chain models may make misleading predictions about patterns of variation in food-chain length because they are unable to incorporate important structural mechanisms that alter food-web dynamics and cause non-linear shifts in food-web structure. Intraguild predation models provide a more appropriate theoretical framework for understanding food-chain length in most natural ecosystems because they accommodate all of the proximate structural mechanisms identified here.     

Food Webs in Relation to Variation in the Environment and Species Assemblage: A Multivariate Approach

The abiotic environment has strong influences on the growth, survival, behavior, and ecology of aquatic organisms. Biotic interactions and species life histories interact with abiotic factors to structure the food web. One measure of food-web structure is food-chain length. Several hypotheses predict a linear relationship between one environmental variable (e.g., disturbance or ecosystem size) and food-chain length. However, many abiotic and biotic variables interact in diverse ways to structure a community, and may affect other measures of food web structure besides food-chain length. This study took a multivariate approach to test the influence of several important environmental variables on four food-web characteristics measured in nine ponds along a hydroperiod gradient over two years. This approach allowed for testing the ecosystem size and dynamic constraints hypotheses while in context of other possibly interacting environmental variables. The relationship between amphibian and invertebrate communities and pond habitat variables was assessed to understand the underlying food-web structure. Hydroperiod and pond area had a strong influence on amphibian and invertebrate communities, trophic diversity and δ¹⁵N range. The range in δ¹³C values responded strongly to dissolved oxygen. Food-chain length responded to multiple environmental variables. Invertebrate and amphibian communities were structured by pond hydroperiod which in turn influenced the trophic diversity of the food web. The results of this study suggest food-chain length is influenced by environmental variation and species assemblage and that a multivariate approach may allow us to better understand the dynamics within and across aquatic food webs.     

食物链长度远因与近因研究进展综述

食物链长度是生态系统的基本属性,其变化决定着群落结构和生态系统功能。稳定同位素分析技术的进步推进了生态系统中食物链长度决定因子相关研究的开展。尽管近期的研究证明了食物链长度与资源可利用性、生态系统大小、干扰等远因之间的关系,但是对于食物网内部结构变化这一近因对食物链长度的影响作用关注较少。综述了边界明确和开放类型淡水生态系统中食物链长度的相关研究进展;探讨了远因和近因机制在决定食物链长度中的作用;给出了判断不同层次和尺度上决定食物链长度机制的概念框架;为今后更好的开展不同生态系统间食物链长度的比较研究提出了建议。     

Modelling Size Structured Food Webs Using a Modified Niche Model with Two Predator Traits

The structure of food webs is frequently described using phenomenological stochastic models. A prominent example, the niche model, was found to produce artificial food webs resembling real food webs according to a range of summary statistics. However, the size structure of food webs generated by the niche model and real food webs has not yet been rigorously compared. To fill this void, I use a body mass based version of the niche model and compare prey-predator body mass allometry and predator-prey body mass ratios predicted by the model to empirical data. The results show that the model predicts weaker size structure than observed in many real food webs. I introduce a modified version of the niche model which allows to control the strength of size-dependence of predator-prey links. In this model, optimal prey body mass depends allometrically on predator body mass and on a second trait, such as foraging mode. These empirically motivated extensions of the model allow to represent size structure of real food webs realistically and can be used to generate artificial food webs varying in several aspects of size structure in a controlled way. Hence, by explicitly including the role of species traits, this model provides new opportunities for simulating the consequences of size structure for food web dynamics and stability.     

食物链长度理论研究进展

食物链长度（Food chain length,FCL）是生态系统中最重要的特点之一,它通过改变生物间的营养关系,影响着生物多样性,群落的结构以及稳定性;它是反映食物网物质转换与能量传递的综合指数,食物链及其动态特征是生态学许多重要理论的基础,食物链长度理论的研究进展,推动了人们对水域生态系统中生物和非生物相互作用的理解。回顾了食物链长度的3种度量方法及其详细的计算方法,在此基础上简述了各方法的特点。综述了食物链长度的决定因素的4种假说（资源可利用性假说、生产力空间假说、生态系统大小假说、动态稳定性假说）及其交互作用,重点总结了湖泊食物链长度的空间格局与决定因素的研究进展。最后,食物链长度研究展望,包括食物链长度决定因子研究存在的问题及发展方向的总结,以及在在水域生态学中的应用的研究进展,例如食物链长度在指示污染物的生物富集中的研究进展、食物链食物链长度在指导生物操作、以及在食物链长度在对气候变化响应方面的研究进展等等。     

Coevolution of species colonisation rates controls food-chain length in spatially structured food webs

How the complexity of food webs depends on environmental variables is a long-standing ecological question. It is unclear though how food-chain length should vary with adaptive evolution of the constitutive species. Here we model the evolution of species colonisation rates and its consequences on occupancies and food-chain length in metacommunities. When colonisation rates can evolve, longer food-chains can persist. Extinction, perturbation and habitat loss all affect evolutionarily stable colonisation rates, but the strength of the competition-colonisation trade-off has a major role: weaker trade-offs yield longer chains. Although such eco-evo dynamics partly alleviates the spatial constraint on food-chain length, it is no magic bullet: the highest, most vulnerable, trophic levels are also those that least benefit from evolution. We provide qualitative predictions regarding how trait evolution affects the response of communities to disturbance and habitat loss. This highlights the importance of eco-evolutionary dynamics at metacommunity level in determining food-chain length.     

Global stability and persistence of simple food chains

The main purpose of this paper is to develop criteria for which a simple food-chain model of intermediate type and of arbitrary length has a globally stable positive equilibrium and to develop criteria under which such a food chain exhibits uniform persistence. The same techniques are used to obtain conditions for a model of a predator-prey system with mutual interference of the predator to possess a globally stable positive equilibrium.     

The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.     

Effects of productivity, disturbance, and ecosystem size on food-chain length: insights from a metacommunity model of intraguild predation

Traditionally, productivity and disturbance have been hypothesized as important determinants of food-chain length. More recently, growing empirical evidence suggests a strong role of ecosystem size. To theoretically explore the effects of basal productivity, disturbance, and ecosystem size on food-chain length, we develop and analyze a metacommunity model of intraguild predation (IGP). The model finds that, when local IGP is weak, increasing basal productivity, weakening disturbance, and increasing ecosystem size will generally increase food-chain length. When local IGP is strong, by contrast, increasing basal productivity or weakening disturbance favors intraguild predators and hinders the coexistence of intraguild predators and intraguild prey, limiting food-chain length. In contrast, increasing ecosystem size can promote coexistence even when local IGP is strong, increasing food-chain length through inserting intraguild prey and changing the degree of omnivory by intraguild predators. Intraguild prey needs to be the superior colonizer to intraguild predators for this to occur. We discuss that these theoretical predictions appear consistent with empirical patterns.     

Stepping in Elton's footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.     

Effect of Sub-Polar Gyre, North Atlantic Oscillation and ambient temperature on size and abundance in the Icelandic Arctic fox

Animals are exposed to environmental factors that influence their life history and body size. Here we used the Arctic fox ( Vulpes lagopus ) as an indicator of the complex links between largescale environmental variables that influence both marine and tundra trophic dynamics to demonstrate how they affect the fox's body size and abundance. The Arctic fox inhabits throughout Iceland, where it preys mainly on birds. We studied the effects of the Sub-Polar Gyre (SPG), winter and summer North Atlantic Oscillation (NAO), mean annual winter and summer temperature, and geographic sector (eastern and western Iceland, which differ in their ecology) on variations in mandible size (6345 specimens) and body mass (2732 specimens) as well as abundance on the Arctic fox in Iceland. We found that (a) SPG index negatively affected male mandible length as well as body mass of both sexes. SPG was also negatively related to fox abundance. (b) Summer NAO had a negative effect on Arctic foxes, that is, cold summers were correlated with shorter mandibles and lower body mass. (c) Winter NAO had a significant negative effect (although weaker than that of summer NAO) on female mandible length, but not on body mass. (d) Summer temperature had a positive effect on female mandible length, but no effect on body mass. However, winter temperature had no effect on either the mandible or body mass. (e) Foxes in the eastern sector had shorter mandibles and were of lighter mass than those in the western sector. We suggest that climate conditions during the growth period of the young affected their final size both directly, by influencing energy metabolism for maintenance, but mainly through their effects on food availability. As far as we are aware, this is the first report that the SPG has an effect on vertebrates, let alone terrestrial ones.     

Disentangling the effects of spring anomalies in climate and net primary production on body size of temperate songbirds

Body size is implicated in individual fitness and population dynamics. Mounting interest is being given to the effects of environmental change on body size, but the underlying mechanisms are poorly understood. We tested whether body size and body condition are related to ambient temperature (heat maintenance hypothesis), or/and explained by variations in primary production (food availability hypothesis) during the period of body growth in songbirds. We also explored whether annual population‐level variations of mean body size are due to changes of juvenile growth and/or size‐dependent mortality during the first year. For 41 species, from 257 sites across France, we tested for relationships between wing length (n = 107 193) or body condition (n = 82 022) and local anomalies in temperature, precipitation and net primary production (NDVI) during the breeding period, for juveniles and adults separately. Juvenile body size was best explained by primary production: wings were longer in years with locally high NDVI, but not shorter in years with low NDVI. Temperature showed a slightly positive effect. Body condition and adult wing length did not covary with any of the other tested variables. We found no evidence of climate‐driven size‐dependent mortality for the breeding season. In our temperate system, local climatic anomalies explained little of the body size variation. A large part of wing length variance was site‐specific, suggesting that avian size was more dependent on local drivers than global ones. Net primary production influenced juvenile size the most through effects on body growth. We suggest that, during the breeding season in temperate systems, thermoregulatory mechanisms are less involved in juvenile growth than food assimilation.     

Food-chain length and adaptive foraging

Food-chain length, the number of feeding links from the basal species to the top predator, is a key characteristic of biological communities. However, the determinants of food-chain length still remain controversial. While classical theory predicts that food-chain length should increase with increasing resource availability, empirical supports of this prediction are limited to those from simple, artificial microcosms. A positive resource availability–chain length relationship has seldom been observed in natural ecosystems. Here, using a theoretical model, we show that those correlations, or no relationships, may be explained by considering the dynamic food-web reconstruction induced by predator''s adaptive foraging. More specifically, with foraging adaptation, the food-chain length becomes relatively invariant, or even decreases with increasing resource availability, in contrast to a non-adaptive counterpart where chain length increases with increasing resource availability; and that maximum chain length more sharply decreases with resource availability either when species richness is higher or potential link number is larger. The interactive effects of resource availability, adaptability and community complexity may explain the contradictory effects of resource availability in simple microcosms and larger ecosystems. The model also explains the recently reported positive effect of habitat size on food-chain length as a result of increased species richness and/or decreased connectance owing to interspecific spatial segregation.     

Food availability modulates temperature‐dependent effects on growth,reproduction, and survival in Daphnia magna

Reduced body size and accelerated life cycle due to warming are considered major ecological responses to climate change with fitness costs at the individual level. Surprisingly, we know little about how relevant ecological factors can alter these life history trade‐offs and their consequences for individual fitness. Here, we show that food modulates temperature‐dependent effects on body size in the water flea Daphnia magna and interacts with temperature to affect life history parameters. We exposed 412 individuals to a factorial manipulation of food abundance and temperature, tracked each reproductive event, and took daily measurements of body size from each individual. High temperature caused a reduction in maximum body size in both food treatments, but this effect was mediated by food abundance, such that low food conditions resulted in a reduction of 20% in maximum body size, compared with a reduction of 4% under high food conditions. High temperature resulted in an accelerated life cycle, with pronounced fitness cost at low levels of food where only a few individuals produced a clutch. These results suggest that the mechanisms affecting the trade‐off between fast growth and final body size are food‐dependent, and that the combination of low levels of food and high temperature could potentially threaten viability of ectotherms.     

The phylogenetic component of food web structure and intervality

Despite the exceptional complexity formed by species and their interactions in ecological networks, such as food webs, regularities in the network structures are repeatedly demonstrated. The interactions are determined by the characteristics of a species. The characteristics are in turn determined by the species’ phylogenetic relationships, but also by factors not related to evolutionary history. Here, we test whether species’ phylogenetic relationships provides a significant proxy for food web intervality. We thereafter quantify the degree to which different species traits remain valuable predictors of food web structure after the baseline effect of species’ relatedness has been removed. We find that the phylogenetic relationships provide a significant background from which to estimate food web intervality and thereby structure. However, we also find that there is an important, non-negligible part of some traits, e.g., body size, in food webs that is not accounted for by the phylogenetic relationships. Additionally, both these relationships differ depending if a predator or a prey perspective is adopted. Clearly, species’ evolutionary history as well as traits not determined by phylogenetic relationships shapes predator-prey interactions in food webs, and the underlying evolutionary processes take place on slightly different time scales depending on the direction of predator-prey adaptations.     

Ecological correlates and determinants in the geographical variation of deer morphology

Previous studies on patterns in ungulate size variations have emphasized the effect of a particular environmental factor such as Bergmann's rule and the island rule. However, although multiple environmental factors may influence the body size, these studies focused on a single factor, and various measurements that may be influenced by different environmental factors (at least partly) were used as indices of body size. In this study, we used several skull and limb measurements to examine size variations among island populations of sika deer (Cervus nippon) in southern Japan considering the effects of multiple environmental factors. We found that all measurements differed markedly between populations. We focused on the skull and limb condylobasal length (CBL) and metacarpal length because they had the most important variations among the populations and the largest sample sizes. The common environmental factors influencing CBL and metacarpal length were island area and precipitation. Since these environmental factors reflect the availability of food resources, the causal factor of body size variation may be food resources. Interpopulation variation in metacarpal length was greater than that of CBL, indicating that metacarpal length may be affected by additional factors besides the common factors shared with CBL. Specific environmental factors influencing relative (CBL adjusted) metacarpal length were precipitation and slope. A common direct cause of those environmental factors was discussed in relation to topography. Analyses of phenotypic variation using multiple measurements with multiple environmental factors are useful to gain insight into underlying causes and can lead to identification of a measurement-specific variation with a specific driving force.     

Biotic and abiotic effects on cohort size distributions in fish

Intraspecific variation in body size is common in animals and plants. Body size affects trophic interactions like foraging ability and vulnerability to predation, which in turn affect individual fitness as well as population stability and extinction risk. Experimental and theoretical work has shown that the size distribution of individuals within cohorts is strongly influenced by intraspecific competition for resources, often leading to skewed frequency distributions. However, little is known about the effects of environmental factors such as climate and eutrophication on the cohort size‐structure of natural populations. We use a long‐term time series of scientific monitoring of a freshwater fish (European perch Perca fluviatilis) to investigate the effects of density dependence, predation, nutrient availability, climate and the timing of spawning on the cohort size distributions. We find that the mean length of the fish is best predicted by the extrinsic factors phosphorus concentration and summer temperature, and the densities of the different age‐classes, whereas the skewness of the length distribution is best predicted by phosphorus concentration, summer temperature, abundance of small fish, and the timing of spawning. Higher nutrient levels, temperatures and densities of small fish increase food availability and thus reduce competition, which is reflected in increased mean length and decreased skewness. The timing of spawning affects skewness presumably through changes in the initial size variation of the cohort and the length of the first growth season. Our results indicate that higher temperatures increase the mean length and decrease skewness due to the concurrent eutrophication of the lake. The study thereby highlights the potential impact of human‐induced environmental change on the size structure of fish populations. More studies are needed to understand better the complex mechanisms through which these factors alter the intensity of intraspecific competition in fish communities.     

Growth pattern and survival in populations of Poecilia vivipara (Teleostei; Poeciliidae) inhabiting an environmental gradient: a common garden study

We performed a common garden experiment to assess the existence of genetic differences on growth and body size between two populations of Poecilia vivipara inhabiting extremes of an environmental gradient caused by water salinity in lagoons of Northern Rio de Janeiro State, Brazil: the Campelo lagoon (freshwater) and Açu lagoon (brackish/saltwater). The two populations show extreme differences in average phenotypes for body size, shape and life history (freshwater populations with smaller body size, lower fecundity and larger reproductive allotment). Pregnant females were brought to the lab and the offspring from both groups were kept in a common recirculating system with freshwater. Standard length and survival were measured weekly over a period of 200 days and growth models were fitted and selected with information criteria. The offspring originally from the brackish water lagoon presented larger asymptotic length, higher maximum growth rate but lower survival than the offspring originally from the freshwater lagoon. Potential confounding variables such as density differences due to mortality and maternal effects (offspring size) were included as covariates in comparisons of growth rates between groups. The results are consistent with phenotypic differences among populations having some genetic basis, and with the existence of a trade-off between growth and maintenance due to the high growth/low survival observed in the group that changed from salt to freshwater. Comparisons of captive and natural populations suggest that the influence of environmental factors, such as salinity, food availability, fish density and predation should also be considered relevant to explain phenotypic variation in this system.