Intraspecific genetic variation and competition interact to influence niche expansion

Theory and empirical evidence show that intraspecific competition can drive selection favouring the use of novel resources (i.e. niche expansion). The evolutionary response to such selection depends on genetic variation for resource use. However, while genetic variation might facilitate niche expansion, genetically diverse groups may also experience weaker competition, reducing density-dependent selection on resource use. Therefore, genetic variation for fitness on different resources could directly facilitate, or indirectly retard, niche expansion. To test these alternatives, we factorially manipulated both the degree of genetic variation and population density in flour beetles (Tribolium castaneum) exposed to both novel and familiar food resources. Using stable carbon isotope analysis, we measured temporal change and individual variation in beetle diet across eight generations. Intraspecific competition and genetic variation acted on different components of niche evolution: competition facilitated niche expansion, while genetic variation increased individual variation in niche use. In addition, genetic variation and competition together facilitated niche expansion, but all these impacts were temporally variable. Thus, we show that the interaction between genetic variation and competition can also determine niche evolution at different time scales.     

High intraguild predator density induces thinning effects on and increases temporal overlap with prey populations

Intraguild (IG) predator density can alter its effects on intraguild prey populations through several mechanisms, including density-dependent processes that affect IG predator traits such as size or growth that enhance or limit its predatory abilities. We examined whether intraspecific density-dependence altered IG predator traits, as well as the subsequent interspecific effects among its intraguild prey within a larval salamander guild. Four densities of ringed salamanders (Ambystoma annulatum), the IG predator, were combined with the presence/absence of spotted salamanders (A. maculatum), the IG prey, within experimental mesocosms. We modeled the effects of A. annulatum density on both conspecific and heterospecific responses that would be indicative of density-dependent competition and predation, respectively. We also modeled the reciprocal interspecific effects of A. maculatum on A. annulatum. We found that increasing intraspecific density negatively affected morphological traits but not survival of A. annulatum. No interspecific effects of A. maculatum on A. annulatum were observed. Alternatively, traits of A. maculatum showed nonlinear relationships with increasing A. annulatum density. Thinning effects of A. annulatum on A. maculatum were observed, as survival was positively and size negatively related for A. maculatum with IG predator density. The temporal overlap of the IG predator and prey also increased nonlinearly with IG predator density, intensifying the potential encounter rate of the two species. Overall, this study shows that density-dependent processes in IG predators can significantly affect traits of both themselves, as well as IG prey, which could ultimately change whether competition or predation occurs between the two groups.     

Soil microbial biomass C and symbiotic processes associated with soybean after sulfentrazone herbicide application

Resource competition and chemical interference are mechanisms of interaction among plants that may occur simultaneously. However, both mechanisms are rarely considered together when modelling plant growth. We propose a new empirical model that estimates biologically significant parameters on both plant competition and chemical interference. The model is tested with data sets from different density-dependent experiments done with two species (the grass Lolium rigidum Gaud. and the legume Glycine max soya L.) subjected to a noxious chemical environment when growing (allelochemicals and herbicides, respectively). Hypotheses on the effect of allelochemicals and its interaction with density are tested using maximum likelihood ratio tests in order to ask, for these species, whether chemical interference is playing a significant role in the interactions among plants or on the contrary, whether interactions among plants are sufficiently explained by the resource competition. In all cases a significant interaction between chemicals and density is observed. This interaction is inconsistent with the hypothesis of only resource competition having an influence of plant biomass and suggests a significant density-dependent effect of chemicals on plant growth.     

Intraspecific Scaling of the Resting and Maximum Metabolic Rates of the Crucian Carp (Carassius auratus)

The question of how the scaling of metabolic rate with body mass (M) is achieved in animals is unresolved. Here, we tested the cell metabolism hypothesis and the organ size hypothesis by assessing the mass scaling of the resting metabolic rate (RMR), maximum metabolic rate (MMR), erythrocyte size, and the masses of metabolically active organs in the crucian carp (Carassius auratus). The M of the crucian carp ranged from 4.5 to 323.9 g, representing an approximately 72-fold difference. The RMR and MMR increased with M according to the allometric equations RMR = 0.212M ^0.776 and MMR = 0.753M ^0.785. The scaling exponents for RMR (b _r) and MMR (b _m) obtained in crucian carp were close to each other. Thus, the factorial aerobic scope remained almost constant with increasing M. Although erythrocyte size was negatively correlated with both mass-specific RMR and absolute RMR adjusted to M, it and all other hematological parameters showed no significant relationship with M. These data demonstrate that the cell metabolism hypothesis does not describe metabolic scaling in the crucian carp, suggesting that erythrocyte size may not represent the general size of other cell types in this fish and the metabolic activity of cells may decrease as fish grows. The mass scaling exponents of active organs was lower than 1 while that of inactive organs was greater than 1, which suggests that the mass scaling of the RMR can be partly due to variance in the proportion of active/inactive organs in crucian carp. Furthermore, our results provide additional evidence supporting the correlation between locomotor capacity and metabolic scaling.     

Interactions between two closely related phytal harpacticoid copepods, asymmetric positive and negative effects

Competition for food is generally thought to exert a strong evolutionary pressure, driving trophic niche separation, either by specialization and/or by widening the choice of potential food resources. Harpacticoid copepods are common inhabitants of phytal assemblages, where several closely related species of the so-called phytal dwelling families often co-occur. However, direct competition among phytal harpacticoids has been thought to be unlikely, due to the abundant and continuously available food supplies. We conducted a series of field and laboratory studies to assess the role of competition in the abundance distribution of two closely related harpacticoid species, Mesochra rapiens and M. aestuari. We found that the abundance of both species co-varied on several seaweed species in the northern Baltic Sea, during a 3-month period. Stable isotope ratios in the green alga Cladophora glomerata field samples indicated different resource utilization of the two species, both in fresh and deteriorated C. glomerata, and in drifting algae. We tested in the laboratory if resource utilization was different between the species in sympatry and allopatry. We used enriched stable carbon isotope ratios (¹³C/¹²C) of the diatom Phaeodactylum tricornutum to trace the uptake in both species. Results from these experiments showed a much higher assimilation by M. aestuari in sympatry with M. rapiens, while the latter species showed a higher assimilation in allopatry. Our results show that while there is no apparent competition for resources between these two species in the field, there seems to be an asymmetric reaction when in sympatry and provided one single resource in the laboratory. We suggest that M. rapiens may facilitate assimilation by M. aestuari and discuss the mechanisms by which this may take place.     

Resource Elasticity of Offspring Survival and the Optimal Evolution of Sex Ratios

The fitness of any organisms includes the survival and reproductive rate of adults and the survival of their offspring. Environmental selection pressures might not affect these two aspects of an organism equally. Assuming that an organism first allocates its limited resources to maintain its survival under environmental selection pressure, our model, based on the evolutionarily stable strategy theory, surprisingly shows that the sex ratio is greatly affected by the environmental pressure intensity and by the reproductive resource elasticity of offspring survival. Moreover, the concept of the resource elasticity of offspring survival intrinsically integrates the ecological concepts of K selection and r selection. The model shows that in a species with reproductive strategy K, increased environmental selection pressure will reduce resource allocation to the male function. By contrast, in a species with reproductive strategy r, harsher environmental selection pressure will increase allocation to the male function. The elasticity of offspring survival might vary not only across species, but also across many other factors affecting the same species (e.g., age structure, spatial heterogeneity), which explains sex ratio differences across species or age structures and spatial heterogeneity in the same species.     

Analysis of changes of insect-plant ratio-improvement of key-factor analysis for evaluating bottom-up effects in insect population dynamics

A revised key-factor analysis was presented for analyzing the temporal changes in the ratio of insect absolute number to plant resource. Ten data sets for 5 insect species were then analyzed. In this key-factor analysis, the key factor is defined as the factor contributing highly to between-year variation inR _r , the log rate of the inter-year change of the insect-plant ratio. The yearly change of plant resource was handled as a separate factor, expressed byr _pl , log ratio of plant resource in yearn to plant resource in yearn+1. The following was revealed: 1) In 7 of the 10 data sets examined,r _pl influenced variations ofR _r ; in particular in 3 casesr _pl was the main key factor. 2) Generation-to-generation fluctuations of absolute insect densities showed density dependence in 4 cases, while those of insect-plant ratios, in 8 cases. 3) The Royama model or a linear model, explained well the relationship between log insect-plant ratio (X _r ) andR _r and the relationship betweenX _r and log yearly change rate of absolute insect density (R _abs ). However, in the 7 cases in whichr _pl was a critical factor for variations ofR _r , with, increase ofX _r ,R _r showed a steeper, decrease around the equilibrium point (the point for whichR _r is 0) thanR _abs . This occurred becauser _pl tended to be negatively correlated withX _r . Consequently, in two casesX _r fluctuated cyclicly or chaotically although without the changes in plant resource, fluctuations ofX _r would be damped oscillations approaching equilibrium.     

Size-dependent foraging capacities and intercohort competition in an ontogenetic omnivore (Arctic char)

Intraspecific competition for resources is strongly influenced by the size of competitors. In this study, we estimated the size‐scaling of the foraging capacities on zooplankton and benthic macroinvertebrates in Arctic char (Salvelinus alpinus) to link size‐dependent performance to effects from competition. The competitive interactions between two size‐classes (YOY and 1‐y) of char were then studied in a large‐scale pond experiment and in two small subarctic lakes. The attack rate function on zooplankton was hump‐shaped whereas the attack rate on benthic chironomids increased monotonically with size. The size‐scaling exponent's for zooplankton and chironomids were 0.65 and 0.30, respectively, leading to that critical resource density (CRD) and maximum growth resource density (G_MRD) increases with size, suggesting an exploitative competitive advantage of small individuals over large. Correspondingly, large (1‐y) char growth was negatively affected by cohort competition whereas small (YOY) char growth was not. Diets of both size classes were dominated by macroinvertebrates with large overlap in prey size suggesting only small gape size advantages for large char. Small char fed to a larger extend on cladocerans which, due to the hump‐shaped foraging efficiency function on zooplankton, were a relatively more profitable resource for small than large char. Estimates of CRD and G_mRD were in correspondence with observed growth responses and resource estimates for zooplankton, whereas for macroinvertebrates only qualitatively correspondence with foraging estimates and char performance was found. Although we were able to explain our results with exploitative competition only, we suggest a general need for size‐dependent foraging estimates on prey in more complex habitats in order to quantitatively link performance and resource abundances. Interference and size‐dependent resource use as mechanisms for observed stable population dynamics in char was not supported by this study and instead a low per capita fecundity and early cannibalism on recruits are suggested to be potential mechanisms that may stabilize char dynamics.     

Intraspecies competition in a field population ofGregopimpla himalayensis (Hymenoptera: Ichneumonidae) parasitic onMalacosoma neustria testacea (Lepidoptera: Lasiocampidae)

Intraspecies competition in a field population of Gregopimpla himalayensis (Hym.: Ichneumonidae) parasitic on the prepupae of Malacosoma neustria testacea (Lep.: Lasiocampidae) was investigated. The parasite oviposits the sufficient number of progeny (5 individuals/0.1 g dry weight of host) to exhaust a single host in a single attack. However, at the intensity less than 22–26 individuals/0.1 g d.w. of host, all individuals can emerge, i.e. density-dependent mortality does not occur. Within this range of intensity, survival of parasite larvae is guaranteed by diminution in body size and decreasing sex ratio. In contrast, total biomass of parasites showed a peak at 5 individuals/0.1 g d.w. of host at which a single host is exhausted. Above the intensity of 22–26, extraordinary minute individuals appeared and they died before maturation. If intraspecies competition play a role in regulation of G. himalayensis population in the field, the process is usually not through density-dependent mortality but through decreasing reproductive rate caused by decrease in the sex ratio, adult longevity and fecundity.     

Anthocyanin production in the hyperaccumulator plant Noccaea caerulescens in response to herbivory and zinc stress

Stomatal behavior in response to drought has been the focus of intensive research, but less attention has been paid to stomatal density. In this study, 5-week-old maize seedlings were exposed to different soil water contents. Stomatal density and size as well as leaf gas exchange were investigated after 2-, 4- and 6-week of treatment, which corresponded to the jointing, trumpeting, and filling stages of maize development. Results showed that new stomata were generated continually during leaf growth. Reduced soil water content significantly stimulated stomatal generation, resulting in a significant increase in stomatal density but a decrease in stomatal size and aperture. Independent of soil water conditions, stomatal density and length in the trumpeting and filling stages were greater than in the jointing stage. Irrespective of growth stage, severe water deficit significantly reduced stomatal conductance (G _s), decreasing the leaf transpiration rate (T _r) and net photosynthetic rate (P _n). Stomatal density was significantly negatively correlated with both P _n and T _r but more strongly with T _r, so the leaf instantaneous water use efficiency (WUE _i ) correlated positively with stomatal density. In conclusion, drought led to a significant increase in stomatal density and a reduction in stomatal size and aperture, resulting in decreased P _n and T _r. Because the negative correlation of stomatal density to T _r was stronger than that to P _n, leaf WUE _i tended to increase.     

Animal response to nested self-similar patches: a test with woolly bears

To gain insight into how animals respond to resource patchiness at different spatial scales, we envision their responses in environments comprised of nested, self-similar patches. In these environments, all resources reside within the smallest patches, and resource density declines as a constant exponent of patch size. Accordingly, we use simple mathematical formulations to describe a self-similar environment and a null model of how animals should respond to this environment if they do not perceive resource distribution. We then argue that animals that can perceive resource distribution should partition space by reducing the relative time searching between patches as patch size increases. On an experimental landscape, we found that woolly bear caterpillars Grammia geneura could partition space in this manner, but the range of patch sizes over which they did so tended to increase with resource aggregation. Nevertheless, scaling efficiency (i.e. the scaling of search time versus the scaling or resource density) was similar in all distributions when averaged over all patch sizes. These disparate patterns with similar outcomes resulted from differences in caterpillars' abilities to discriminate spatially among patches of different sizes via their movement pathways, and differences in their use of speed to detect resource items. Our work is relevant to the characterization of resource availability from an animal's perspective, and to the linking of optimal foraging theory to the modeling of search behavior.     

Trophic Strategies of a Non-Native and a Native Amphibian Species in Shared Ponds

One of the critical factors for understanding the establishment, success and potential impact on native species of an introduced species is a thorough knowledge of how these species manage trophic resources. Two main trophic strategies for resource acquisition have been described: competition and opportunism. In the present study our objective was to identify the main trophic strategies of the non-native amphibian Discoglossus pictus and its potential trophic impact on the native amphibian Bufo calamita. We determine whether D. pictus exploits similar trophic resources to those exploited by the native B. calamita (competition hypothesis) or alternative resources (opportunistic hypothesis). To this end, we analyzed the stable isotope values of nitrogen and carbon in larvae of both species, in natural ponds and in controlled laboratory conditions. The similarity of the δ¹⁵N and δ¹³C values in the two species coupled with isotopic signal variation according to pond conditions and niche partitioning when they co-occurred indicated dietary competition. Additionally, the non-native species was located at higher levels of trophic niches than the native species and B. calamita suffered an increase in its standard ellipse area when it shared ponds with D. pictus. These results suggest niche displacement of B. calamita to non-preferred resources and greater competitive capacity of D. pictus in field conditions. Moreover, D. pictus showed a broader niche than the native species in all conditions, indicating increased capacity to exploit the diversity of resources; this may indirectly favor its invasiveness. Despite the limitations of this study (derived from potential variability in pond isotopic signals), the results support previous experimental studies. All the studies indicate that D. pictus competes with B. calamita for trophic resources with potential negative effects on the fitness of the latter.     

Evolution of density-dependent dispersal in a structured metapopulation

We study the evolution of density-dependent dispersal in a structured metapopulation subject to local catastrophes that eradicate local populations. To this end we use the theory of structured metapopulation dynamics and the theory of adaptive dynamics.The set of evolutionarily possible dispersal functions (i.e., emigration rates as a function of the local population density) is derived mechanistically from an underlying resource-consumer model. The local resource dynamics is of a flow-culture type and consumers leave a local population with a constant probability per unit of time κ when searching for resources but not when handling resources (i.e., eating and digesting). The time an individual spends searching (as opposed to handling) depends on the local resource density, which in turn depends on the local consumer density, and so the average per capita emigration rate depends on the local consumer density as well.The derived emigration rates are sigmoid functions of local consumer population density. The parameters of the local resource-consumer dynamics are subject to evolution. In particular, we find that there exists a unique evolutionarily stable and attracting dispersal rate κ^∗ for searching consumers. The κ^∗ increases with local resource productivity and decreases with resource decay rate. The κ^∗ also increases with the survival probability during dispersal, but as a function of the catastrophe rate it reaches a maximum before dropping off to zero again.     

The functional responses of adaptive consumers of two resources

This article investigates some aspects of the shape of the functional responses of consumers that utilize two resources. Adaptive variation in consumption behavior is shown to have a major effect on the relationship between amount of resource available and its rate of consumption by an average consumer individual. The effects of adaptive variation are dependent on the nutritional status of the two resources. If the resources are linearly substitutable, increases in the density of resource i will usually increase the quantity, functional response on i divided by density of i, and increases in the density of resource j will decrease this quantity. The result is that the functional response to resource i will generally decrease with the density of resource j, and will increase faster than it would otherwise have increased with the density of resource i. If resources are nonsubstitutable, an adaptive functional response to resource i will increase with the density of resource j, and it will increase more slowly with the density of resource i than it would have without adaptive change. If resources are both complementary and substitutable, the functional response will exhibit ranges of smooth change separated by rapid jumps between values, and different ranges of resource densities will result in a functional response with the characteristics of linearly substitutable or of non-substitutable resources. Adaptive functional response shape is dependent upon the tradeoff involved in raising each functional response. These results have implications for the types of indirect interactions that occur between resources as the result of a common consumer's functional response. They also suggest that the adaptive response of competing consumers to each other will differ depending on the nutritional status of the resources for which they are competing. Implications of these findings for consumer growth isocline shape and several other issues are explored.     

Density dependence,territoriality, and divisibility of resources: from optimality models to population processes

Species differ enormously in their territorial systems. Some species defend only small areas surrounded by undefended space, while others defend large contiguous territories. Using an optimization approach, we show that this variation can be explained from the density of two types of resources: divisible and nondivisible. We assume that benefits of territories are monotonously related to the defended amount of divisible resources (hereafter called food). In contrast, no benefits are obtained without a nondivisible resource (hereafter called nest site) in the territory, while more than one nest site does not further increase the benefits. The optimal territory size depends on the relative abundance of these resources. With a low density of nest sites, the optimal territory size is small and includes only the nest site. If the density of nest sites is relatively large, the optimal territory size is high, and territories are contiguous. Competition for these different resources yields contrasting patterns of how populations are regulated. If there is mainly competition for nest sites, we expect density-dependent exclusion through territoriality and no density-dependent reproduction. When competition is mainly for food, we expect density-dependent reproduction because optimal territory size will be compressed at higher densities, resulting in lower reproductive success. These predicted patterns indeed are observed in some well-studied passerine species for which both the territorial system and the occurrence of density dependence is known.     

Emotion Based Attentional Priority for Storage in Visual Short-Term Memory

A plethora of research demonstrates that the processing of emotional faces is prioritised over non-emotive stimuli when cognitive resources are limited (this is known as ‘emotional superiority’). However, there is debate as to whether competition for processing resources results in emotional superiority per se, or more specifically, threat superiority. Therefore, to investigate prioritisation of emotional stimuli for storage in visual short-term memory (VSTM), we devised an original VSTM report procedure using schematic (angry, happy, neutral) faces in which processing competition was manipulated. In Experiment 1, display exposure time was manipulated to create competition between stimuli. Participants (n = 20) had to recall a probed stimulus from a set size of four under high (150 ms array exposure duration) and low (400 ms array exposure duration) perceptual processing competition. For the high competition condition (i.e. 150 ms exposure), results revealed an emotional superiority effect per se. In Experiment 2 (n = 20), we increased competition by manipulating set size (three versus five stimuli), whilst maintaining a constrained array exposure duration of 150 ms. Here, for the five-stimulus set size (i.e. maximal competition) only threat superiority emerged. These findings demonstrate attentional prioritisation for storage in VSTM for emotional faces. We argue that task demands modulated the availability of processing resources and consequently the relative magnitude of the emotional/threat superiority effect, with only threatening stimuli prioritised for storage in VSTM under more demanding processing conditions. Our results are discussed in light of models and theories of visual selection, and not only combine the two strands of research (i.e. visual selection and emotion), but highlight a critical factor in the processing of emotional stimuli is availability of processing resources, which is further constrained by task demands.     

Scaling in Free-Swimming Fish and Implications for Measuring Size-at-Time in the Wild

This study was motivated by the need to measure size-at-age, and thus growth rate, in fish in the wild. We postulated that this could be achieved using accelerometer tags based first on early isometric scaling models that hypothesize that similar animals should move at the same speed with a stroke frequency that scales with length^-1, and second on observations that the speed of primarily air-breathing free-swimming animals, presumably swimming ‘efficiently’, is independent of size, confirming that stroke frequency scales as length^-1. However, such scaling relations between size and swimming parameters for fish remain mostly theoretical. Based on free-swimming saithe and sturgeon tagged with accelerometers, we introduce a species-specific scaling relationship between dominant tail beat frequency (TBF) and fork length. Dominant TBF was proportional to length^-1 (r² = 0.73, n = 40), and estimated swimming speed within species was independent of length. Similar scaling relations accrued in relation to body mass^-0.29. We demonstrate that the dominant TBF can be used to estimate size-at-time and that accelerometer tags with onboard processing may be able to provide size-at-time estimates among free-swimming fish and thus the estimation of growth rate (change in size-at-time) in the wild.     

Interspecific competition between the red imported fire ant,Solenopsis invicta Buren and ghost ant,Tapinoma melanocephalum Fabricius for honeydew resources produced by an invasive mealybug,Phenacoccus solenopsis Tinsiley

In natural as in agricultural ecosystems, interactions between ants and honeydew-producing hemipterans are commonly observed. Mutualisms between invasive ants and hemipterans have been extensively studied in recent years. However, native ant species can equally exploit the honeydew excreted by hemipterans, and establish close relationships with them. Up till present, little is known about the competition between exotic ants (such as Solenopsis invicta) and its co-occurring species (e.g., Tapinoma melanocephalum) for this food resource. In this study, we compared the competitive ability of the invasive ant S. invicta and its co-occurring species T. melanocephalum in the laboratory. We also determined whether the two ant species could coexist and share honeydew resource. Our results indicate that the foraging activity of T. melanocephalum was restrained by S. invicta. Mortality of S. invicta and T. melanocephalum was significantly higher in T. melanocephalum colony case than that in other cases. The invasive ability between the two ant species was significantly different. These results suggest that S. invicta suppresses exploitation of honeydew-producing hemipterans by native ants and occupies most of honeydew resource. S. invicta could not completely drive T. melanocephalum out of honeydew competition, with small numbers of T. melanocephalum workers coexisting and sharing the honeydew with S. invicta. This finding permits a better understanding of the invasion success of S. invicta, and its ability to occupy new habitats.