On the Measurement of Ecological Novelty: Scale-Eating Pupfish Are Separated by 168 my from Other Scale-Eating Fishes

The colonization of new adaptive zones is widely recognized as one of the hallmarks of adaptive radiation. However, the adoption of novel resources during this process is rarely distinguished from phenotypic change because morphology is a common proxy for ecology. How can we quantify ecological novelty independent of phenotype? Our study is split into two parts: we first document a remarkable example of ecological novelty, scale-eating (lepidophagy), within a rapidly-evolving adaptive radiation of Cyprinodon pupfishes on San Salvador Island, Bahamas. This specialized predatory niche is known in several other fish groups, but is not found elsewhere among the 1,500 species of atherinomorphs. Second, we quantify this ecological novelty by measuring the time-calibrated phylogenetic distance in years to the most closely-related species with convergent ecology. We find that scale-eating pupfish are separated by 168 million years of evolution from the nearest scale-eating fish. We apply this approach to a variety of examples and highlight the frequent decoupling of ecological novelty from phenotypic divergence. We observe that novel ecology is not always tightly correlated with rates of phenotypic or species diversification, particularly within recent adaptive radiations, necessitating the use of additional measures of ecological novelty independent of phenotype.     

Niche width and niche overlap: a method based on type-2 fuzzy sets

Complicated ecosystems and the high non-linearity of evolution has made biology more adaptable to a variety of environments. The relationship between life and environment demands a dynamic definition of niche and its measurement. In this paper we propose a model of niche with dynamic character based on the “broad band” effect in type-2 fuzzy sets. The niche in this definition is an interval in each ecological dimension which is dynamic in character and depends on the actual environment. We also give formulas for niche width and niche overlap. We compute the niche width and overlap for plants and animals and then compare these results with previous results. The results for niche width in this paper reflect the diversity of resources used by species or communities. The results for niche overlap demonstrate overlap under different environmental conditions. The results are, moreover, intervals, which could provide more information. The model in this paper could therefore be used to describe the state of every resource comprehensively, reflecting the interaction between species and environment.     

Temperature as an Ecological Resource

Ectothermic vertebrates respond to the temperature of theirhabitat in a manner that is remarkably similar to their responseto more traditional ecological resources such as food. We reviewthe response to temperature primarily from literature on fishesin terms of ecological concepts related to niche theory andcompetition. The width of the fundamental thermal niche is about4°C when measured by a mean plus and minus one standarddeviation of the distribution of temperature occupied in a laboratorygradient. Fish of temperate freshwater appear to fall into threethermal guilds along the temperature resource axis —cold,cool, and warm water fishes. Realized thermal niches are similarin central tendency to fundamental niches, but niche width appearsto be more narrow for the realized niche in limited sample data.The success of interference competition for space with preferredtemperature is tied to social dominance in a manner analogousto food competition. Thermal niche shifts in the face of interspecificcompetition for preferred temperature appear supported by onelaboratory study. Exploitation competition in respect to temperatureseems nebulous. If animals successfully compete for their thermalniche, growth and perhaps other measures of fitness are maximized.Cost/benefit models for thermal resources and food resourceslead to similar predictions about resource use. We suggest thatviewing temperature and other niche axes in the way ecologistshave viewed food resources would be useful.     

Beyond ecological opportunity: Prey diversity rather than abundance shapes predator niche variation

1. Ecological opportunity (i.e. the diversity of available resources) has a pivotal role in shaping niche variation and trophic specialisation of animals. However, ecological opportunity can be described with regard to both diversity and abundance of resources. The degree to which these two components contribute to niche variation remains unexplored.
2. To address this, we used an extensive dataset on fish diet and benthic invertebrate diversity and density from 73 sampling events in three Norwegian rivers in order to explore realised trophic niches and the response of dietary niche variation along gradients of resource diversity (potential trophic niches), resource density (as a proxy of resource abundance) and fish density (as a proxy of inter‐ and intra‐specific competition) in a freshwater top predator (the brown trout, Salmo trutta L.).
3. Linear models indicated that individual and population niche variation increased with increasing ecological opportunity in terms of prey diversity. However, no simple cause‐and‐effect associations between niche indices and prey abundance were found. Our multiple regression analyses indicated that the abundance of certain resources (e.g. Chironomidae) can interact with prey diversity to determine individual and population realised trophic niches. Niche variation (within‐individual component and inter‐individual diet variation) decreased with increasing inter‐ and intra‐specific competition.
4. This study extends prevailing trophic ecology theory by identifying diversity, rather than density, of available prey resources as a primary driver of niche variation in fish of temperate riverine systems with no extensive resource limitation. The study also shows that ecological opportunity may mask the direction of the effect (compression or expansion) of competition on niche variation when food resources are diverse.
5. Our study provides novel empirical insight to the driving forces behind niche variation and reveals that diversity, rather than density, of available prey resources may be a primary driver of niche variation in freshwater fish. Our study supports the view that a broader potential trophic niche promotes broader realised trophic niche variation by individuals, which leads to individual niche diversification by opening access to alternatives resources, resulting in a concomitant rise in the realised trophic niche width of the population.

     

Coevolution in resource-limited competition communities

A coevolutionary model of species packing is developed which allows evolutionary adjustment in both niche position and within-phenotype niche width of from one of three competing species. The environment is specified as a single resource dimension x and availability of resources along x is given by a Gaussian curve with parameters ($\text{x}\text{?}{}_{\mathrm{R}}\text{and}\text{σ}{}_{\mathrm{R}}$). The model predicts that for S species the ratio of optimal niche width (?) to σ_R is roughly independent of σ_R and can be approximated by $\text{1}\text{S}$ when the competitors are completely resource limited. Niche separation ($\text{d}\text{?}\text{w}\text{?}$) increases only moderately with increases in resource diversity σ_R and is greater for two than for three competing species. To the extent that the competitors do not utilize all resources, both optimal niche separation and niche width decrease. Many of the general trends in niche width and niche separation predicted by this coevolutionary model parallel those from optimal foraging theory and limiting similarity models of community structure. The coevolutionary model stands out, however, in the singularly high values predicted for niche separation, making coevolved communities highly invadeable. Hence, the theory suggests, as some empirical evidence indicates, that coevolved competition communities can only exist as such on remote islands or in other habitats which might be free from invasion by outside species. For such communities, resource diversity (σ_R) should have much less effect on species packing than either species number itself or the extent that these competitors are resource limited.     

Trait plasticity in species interactions: a driving force of community dynamics

Evolutionary community ecology is an emerging field of study that includes evolutionary principles such as individual trait variation and plasticity of traits to provide a more mechanistic insight as to how species diversity is maintained and community processes are shaped across time and space. In this review we explore phenotypic plasticity in functional traits and its consequences at the community level. We argue that resource requirement and resource uptake are plastic traits that can alter fundamental and realised niches of species in the community if environmental conditions change. We conceptually add to niche models by including phenotypic plasticity in traits involved in resource allocation under stress. Two qualitative predictions that we derive are: (1) plasticity in resource requirement induced by availability of resources enlarges the fundamental niche of species and causes a reduction of vacant niches for other species and (2) plasticity in the proportional resource uptake results in expansion of the realized niche, causing a reduction in the possibility for coexistence with other species. We illustrate these predictions with data on the competitive impact of invasive species. Furthermore, we review the quickly increasing number of empirical studies on evolutionary community ecology and demonstrate the impact of phenotypic plasticity on community composition. Among others, we give examples that show that differences in the level of phenotypic plasticity can disrupt species interactions when environmental conditions change, due to effects on realized niches. Finally, we indicate several promising directions for future phenotypic plasticity research in a community context. We need an integrative, trait-based approach that has its roots in community and evolutionary ecology in order to face fast changing environmental conditions such as global warming and urbanization that pose ecological as well as evolutionary challenges.     

Trophic niche differences between coexisting omnivores silver carp and bighead carp in a pelagic food web

Understanding how omnivorous consumers are affected by their resources and how this is expressed through the food chain is a fundamental issue in ecology. We used stable isotope analysis of archived scales of two pelagic single-chain omnivorous fish species, bighead carp (Hypophthalmichthys nobilis) and silver carp (H. molitrix), to reconstruct historical trophic interactions patterns along a gradient of resources. We found that, although bighead carp and silver carp utilize the similar resources from the pelagic food chain, they can coexist and persist not only by regulating their trophic position and trophic dissimilarity, but also by regulating trophic niche width. Omnivorous fish often exhibit flexible foraging strategies, which is closely related to the availability of ecologic context. We found a positive relationship between trophic dissimilarity and zooplankton density, which may indicate that the competitive interactions induce strong top-down effects on zooplankton, and/or that high zooplankton availability release the between-population trophic interaction through bottom-up effect. The trophic niche width of bighead carp was positively related with zooplankton availability, probably reflecting that the niche of an omnivore at a higher trophic position is more sensitive to high quality resources. Our results indicate how different aspects of the trophic partitioning of coexisting omnivores may be regulated by different ecological contexts. These alternatives are not mutually exclusive and further theoretical work should include both these mechanisms to re-evaluate the effects of omnivory on food web properties.     

The effects of morphology,phylogeny and prey availability on trophic resource partitioning in an anuran community

Several factors influence the partitioning of trophic resources in ecological communities, such as morphology, evolutionary history, and resource availability. Although the effects of morphology, phylogeny, and resource availability on trophic ecology have long been explored by theoretical studies, little has been done to empirically test these relationships. Here, we tested whether phylogenetic and morphological distances correlate with trophic niche overlap using a path analysis of multiple partial regression of distance matrices. Also, we tested whether niche breadth is influenced by body size using Phylogenetic Generalized Least Squares analysis. Trophic niche overlap was better explained by morphology per se than by the phylogenetic distance. We also found that predator's body size influences niche breadth calculated considering prey traits and availability, but not when we do not include these availability data. Additionally, trophic niche breadth was usually smaller when we considered prey traits and availability, differently from niche overlap, whose values increased when we did not consider these data. Our findings show that the interpretation of trophic niche in communities changes if we consider availability data, affecting inferences about coexistence and trophic specialization. Our study contributes to understanding trophic specialization and emphasizes the importance of incorporating prey availability and their traits into diet analysis.     

ON THE COEXISTENCE AND COEVOLUTION OF ASEXUAL AND SEXUAL COMPETITORS

The coexistence and coevolution of sexual and asexual species under resource competition are explored with three models: a nongenetic ecological model, a model including single locus genetics, and a quantitative-genetic model. The basic assumption underlying all three models is that genetic differences are translated into ecological differences. Hence if sexual species are genetically more variable, they will be ecologically more variable. Under classical competition theory, this increased ecological variability can, in many cases, be an advantage to individual sexual genotypes and to the sexual species as a whole. The purpose of this paper is to determine the conditions when this advantage will outway three disadvantages of sexuality: the costs of males, of segregation, and of the additive component of recombination. All three models reach similar conclusions. Although asexuality confers an advantage, it is much less than a two-fold advantage because minor increases in the overall species niche width of the sexual species will offset the reproductive advantage of the asexual species. This occurs for two reasons. First, an increase in species niche width increases the resource base of the sexual species. Second, to the extent that the increase in niche width is due to increased differences between individuals, a reduction in intraspecific competition will result. This is not to imply that the sexual species will always win. The prime conditions that enable sexual species to stably coexist with or even supplant an asexual sister species are:

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Making a living in the tropical forest: Yuquí foragers in the Bolivian Amazon

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Quantifying Inter- and Intra-Population Niche Variability Using Hierarchical Bayesian Stable Isotope Mixing Models

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Reconsidering the Specialist-Generalist Paradigm in Niche Breadth Dynamics: Resource Gradient Selection by Canada Lynx and Bobcat

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