Conditions under which plants help herbivores and benefit from predators through apparent competition

Leaf domatia are tiny structures in leaf vein axils that are typically inhabited by predatory and fungivorous mites. A recent article reported plant domatia specifically suited for herbivorous mites, which seems paradoxical, since the plant is thus supporting a natural enemy that can harm itself. The authors claimed that domatia are created to promote herbivorous mites as "fodder" for predatory mites that attack another herbivorous mite damaging the plant, and that the relationship among the plant, the fodder mite, and the predatory mite constitute a multiway mutualism because all three species benefit from the interaction. I formulate this system using two simple mathematical models of apparent competition, which differ in how domatia are modeled, and then assess when it is advantageous for the plant to create such space for a natural enemy. As a necessary condition for mutualism, the product of reproductive efficiency and nutritious value of the fodder prey should exceed that of the pest prey. This condition is also sufficient, if the direct costs for making the structure of domatia are negligible. If there are significant costs, however, the condition is broader for predators with lower reproductive efficiency and higher mortality, and for non-fodder prey with high consumption rate and low predation rate. I suggest that creating domatia is more effective when predators are less prolific and non-fodder prey are more severe as pests. Finally, I discuss how this mathematical model can apply to a wider range of tritrophic mutualistic relationships such as those among plants, aphids, and ants.     

Exploitative competition between invasive herbivores benefits a native host plant

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).     

Host-plant-mediated competition via induced resistance: interactions between pest herbivores on potatoes.

Plant-mediated competition among insect herbivores occurs when one species induces changes in plant chemistry, nutrition, or morphology that render plants resistant to attack by others. We explored plant-mediated interspecific interactions between the potato leafhopper (Empoasca fabae) and the Colorado potato beetle (Leptinotarsa decemlineata), two important pests on potatoes. Leafhoppers colonize fields in advance of beetles, and thus the possibility exists that previous feeding by leafhoppers induces changes in potato plants that have adverse consequences for beetles. The consequences of leafhopper-induced resistance for beetle performance were studied in the greenhouse, field cages, and in large open-field plots. Potato plants were exposed to four densities of leafhoppers (none, low, moderate, and high), and visible feeding symptoms were measured as percentage leaf curling, chlorosis, and necrosis. The oviposition preference, performance, and survivorship of Colorado potato beetles were then measured on the four categories of induced plants in field-cage and greenhouse settings. In open field plots, survival on the four categories of induced plants was determined by placing cohorts of beetle adults onto plants and measuring the densities of resulting eggs, larvae, and emerging Fl adults. Leafhopper-induced symptoms on potato plants were density dependent, with the percentage of curled, chlorotic, and necrotic leaves increasing with leafhopper density. Previous feeding by leafhoppers adversely affected oviposition and larval performance of beetles. Fewer egg masses were deposited on plants that incurred high levels of leafhopper feeding. Similarly, larval development was delayed and emerging adult beetles weighed less when fed induced foliage from the high leafhopper-density treatment. Beetles survived less well in the field on plants experiencing moderate and high levels of leafhopper feeding as evidenced by lower densities of eggs, larvae, and emerging F1 adults. Overall, leafhoppers and beetles competed through feeding-induced changes in plant quality. Notably, the asymmetric interaction took place at a large spatial scale in open field plots and had negative consequences that persisted to the next beetle generation. Ultimately, to establish an effective management strategy for crop pests such as leafhoppers, it is essential to consider the positive indirect effects of induced resistance along with the negative direct effects on crop yield.     

Behavioral response to predators reverses the outcome of competition between prey species

Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.     

The function of hatching asynchrony in the blue-footed booby

The blue-footed booby (Sula nebouxii) commonly hatches two eggs 4 days apart; then the senior (first-hatched) chick aggressively dominates the other and sometimes kills it. Two hypotheses explaining the function of the hatching interval were tested by creating broods with synchronous hatching: the facultative brood reduction hypothesis of Lack (1954) and the sibling rivalry reduction hypothesis of Hahn (1981). The results contradicted most predictions of both hypotheses: synchronous broods formed an aggressive hierarchy similarly to asynchronous broods (controls), and subordinate chicks grew poorly (Fig. 1) and died frequently, similarly to junior chicks in control broods. However, compared with synchronous broods, asynchronous broods showed less aggression (Fig. 2), diminished food allocation to subordinate chicks (Fig. 3) and less total food consumption (30% fewer feeds at age 0–10 days). These behavioral comparisons took into account the different ages of chicks in different treatments. The results suggest that natural asynchrony makes brood reduction more efficient and decreases the costs of sibling aggression to parents, in terms of their future survival or fecundity, as proposed by Mock and Ploger (1987). Further, in exaggeratedly asynchronous broods (8-day hatch interval) junior chicks suffered more aggression (Fig. 4) and grew more slowly than junior chicks in control broods. This result supports the hypothesis of optimal hatch asynchrony of Mock and Ploger (1987).     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.     

Specificity between Neotropical tree seedlings and their fungal mutualists leads to plant-soil feedback

A growing body of evidence obtained largely from temperate grassland studies suggests that feedbacks occurring between plants and their associated soil biota are important to plant community assemblage. However, few studies have examined the importance of soil organisms in driving plant-soil feedbacks in forested systems. In a tropical forest in central Panama, we examined whether interactions between tree seedlings and their associated arbuscular mycorrhizal fungi (AMF) lead to plant-soil feedback. Specifically, do tropical seedlings modify their own AMF communities in a manner that either favors or inhibits the next cohort of conspecific seedlings (i.e., positive or negative feedback, respectively)? Seedlings of two shade-tolerant tree species (Eugenia nesiotica, Virola surinamensis) and two pioneer tree species (Luehea seemannii, Apeiba aspera) were grown in pots containing identical AMF communities composed of equal amounts of inoculum of six co-occurring AMF species. The different AMF-host combinations were all exposed to two light levels. Under low light (2% PAR), only two of the six AMF species sporulated, and we found that host identity did not influence composition of AMF spore communities. However, relative abundances of three of the four AMF species that produced spores were influenced by host identity when grown under high light (20% PAR). Furthermore, spores of one of the AMF species, Glomus geosporum, were common in soils of Luehea and Eugenia but absent in soils of Apeiba and Virola. We then conducted a reciprocal experiment to test whether AMF communities previously modified by Luehea and Apeiba differentially affected the growth of conspecific and heterospecific seedlings. Luehea seedling growth did not differ between soils containing AMF communities modified by Luehea and Apeiba. However, Apeiba seedlings were significantly larger when grown with Apeiba-modified AMF communities, as compared to Apeiba seedlings grown with Luehea-modifed AMF communities. Our experiments suggest that interactions between tropical trees and their associated AMF are species-specific and that these interactions may shape both tree and AMF communities through plant-soil feedback.     

Sperm competition as a mechanism of female choice in the field cricket,Gryllus bimaculatus

Summary While traditionally viewed as an extension of intermale competition, mechanisms of sperm competition may be used by multiply mating females for mate choice. In the field cricket G. bimaculatus sperm were shown to mix in the spermatheca. The proportion of offspring sired by the second male increased with spermatophore attachment duration and, therefore, the number of sperm transferred. There was no second male advantage for single matings after an initial double mating. However, the proportion ofoffspring sired by the second male increased in proportion to the number of times he mated such that second males mating three times after an initial double mating had the advantage at fertilization. The data suggested that sperm were utilized in proportion to their numerical representation in the spermatheca. The mechanism of sperm precedence may, therefore, be one of sperm dilution. Female G. bimaculatus may control the degree of sperm competition as a mechanism of mate choice. By accepting large quantities of sperm from chosen males they may determine the paternity of their offspring by diluting out the sperm stored from previous matings.     

Invasional interference due to similar inter- and intraspecific competition between invaders may affect management

As the number of biological invasions increases, the potential for invader-invader interactions also rises. The effect of multiple invaders can be superadditive (invasional meltdown), additive, or subadditive (invasional interference); which of these situations occurs has critical implications for prioritization of management efforts. Carduus nutans and C. acanthoides, two congeneric invasive weeds, have a striking, segregated distribution in central Pennsylvania, U.S.A. Possible hypotheses for this pattern include invasion history and chance, direct competition, or negative interactions mediated by other species, such as shared pollinators. To explore the role of resource competition in generating this pattern, we conducted three related experiments using a response-surface design throughout the life cycles of two cohorts. Although these species have similar niche requirements, we found no differential response to competition between conspecifics vs. congeners. The response to combined density was relatively weak for both species. While direct competitive interactions do not explain the segregated distributional patterns of these two species, we predict that invasions of either species singly, or both species together, would have similar impacts. When prioritizing which areas to target to prevent the spread of one of the species, it is better to focus on areas as yet unaffected by its congener; where the congener is already present, invasional interference makes it unlikely that the net effect will change.     

Lack of phlorotannin induction in the kelp Laminaria hyperborea in response to grazing by two gastropod herbivores

Phlorotannins (polyphenolic compounds) are found exclusively in brown algae and are commonly regarded as an important component of algal defense against herbivores. The variation in phlorotannin concentration within and among individual seaweed specimens of the same species can be substantial, and seaweeds may respond to herbivore damage by an increased production of phlorotannins in order to prevent further herbivory. In the present study, we investigated the natural variation of phlorotannins in different parts of the kelp Laminaria hyperborea in an observational field survey. We also performed a manipulative induction experiment in which we studied the effect of natural grazing by the patellid limpet Ansates (=Helcion) pellucida and the littorinid snail Lacuna vincta, as well as artificial damage on the production of phlorotannins in L. hyperborea. There was a large variation in phlorotannin content both among and within L. hyperborea individuals and populations. The control plants in the induction experiment had a higher phlorotannin content compared to plants in natural populations, but grazing by A. pellucida or L. vincta, or artificial damage did not induce a higher phlorotannin concentration in the kelps. Instead, the phlorotannin content in wounded kelps was significantly lower than in the control plants, possibly due to removal of phlorotannin-rich outer layers of the kelp thallus, or to an induced compensatory growth response in the L. hyperborea plants. These results imply that L. hyperborea phlorotannins probably do not function as an inducible chemical defense against A. pellucida and L. vincta, instead grazing by these herbivores decreases the phlorotannin content of the kelps.     

Intraspecific competition: the role of lags between attack and death in host-parasitoid interactions

Many natural enemies do not immediately kill their host, and the lag this creates between attack and host death results in mixed populations of uninfected and infected hosts. Both competition and parasitism are known to be major structuring forces in ecological communities; however, surprisingly little is known about how the competitive nature of infected hosts could affect the survival and dynamics of remaining uninfected host populations. Using a laboratory system comprising the Indian meal moth, Plodia interpunctella, and a solitary koinobiont parasitoid, Venturia canescens, we address this question by conducting replicated competition experiments between the unparasitized and parasitized classes of host larvae. For varying proportions of parasitized host larvae and competitor densities, we consider the effects of competition within (intraclass) and between (interclass) unparasitized and parasitized larvae on the survival, development time, and size of adult moths and parasitoid wasps. The greatest effects were on survival: increased competitor densities reduced survival of both parasitized and unparasitized larvae. However, unparasitized larvae survival, but not parasitized larvae survival, was reduced by increasing interclass competition. To our knowledge, this is the first experimental demonstration of the competitive superiority of parasitized over unparasitized hosts for limiting resources. We discuss possible mechanisms for this phenomenon, why it may have evolved, and its possible influence on the stability of host-parasite dynamics.     

The importance of competitor identity,morphology and ranking methodology to outcomes in interference competition between sponges

The nature or structure of competition within communities has been a dynamic area of practical and theoretical research for a number of decades. Certain components of some communities have proved hard to incorporate within such studies; in marine benthic work this has included the sponges. In this study, intra- and inter-phyletic interactions involving sponges (7,817 in total) were studied on 125 rocks (ranging in surface area from 10 to 2,438 cm ²) at a site experiencing low disturbance levels at Lough Hyne Marine Nature Reserve, Co. Cork, Ireland. The outcomes of sponge-sponge (intra-phyletic) interactions were significantly different in both outcome and structure to those involving sponges and representatives of other phyla. Typically sponge-sponge competition resulted in a much higher number of tied outcomes than did inter-phyletic encounters. Sponges over-grew all groups of organisms with the exception of some cnidarians and ascidians. The morphology of any sponge species (thickness in most cases) was important in determining the outcome of intra-phyletic, but not inter-phyletic, interactions. Sponges which exhibited thick (>2 mm) crusts were, in the majority of interactions, superior competitors compared to the thin (<1 mm) crusts. However, the method used to rank species (i.e. wins:loss or wins:total ratio) made a significant difference to the ranking of sponge competitors. The transitivity of the sponge assemblage investigated was calculated using the index formulated by Tanaka and Nandakumar (1994) as 0.23, indicating the assemblage to be organised as a network rather than a hierarchical system. We suggest this network of principally tied outcomes (due to standoffs) may be maintained in part by chemical interaction. However, in the absence of disturbance, the network is more likely to be mediated through processes of growth and regression of tissues during summer and winter months respectively.     

Species-specific responses of herbivores to within-plant and environmentally mediated between-plant variability in plant chemistry

Allocation of resources to growth and defense against herbivores crucially affects plant competitiveness and survival, resulting in a specific distribution of assimilates and defense compounds within plant individuals. Additionally, plants rarely experience stable environmental conditions, and adaptations to abiotic and biotic stresses may involve shifts in resistance to herbivores. We studied the allocation of phytochemicals in Brassica oleracea (Brussels sprouts) due to leaf age, drought stress and herbivore damage and assessed effects on two lepidopteran herbivores differing in diet breadth: the generalist Spodoptera littoralis and the specialist Pieris brassicae. Glucosinolates as secondary defense compounds and total nitrogen and carbon were quantified and linked to plant palatability, i.e., herbivore feeding preference. Herbivore responses were highly species-specific and partially related to changes in phytochemicals. Spodoptera littoralis preferred middle-aged leaves with intermediate levels of glucosinolates and nitrogen over young, glucosinolate and nitrogen rich leaves, as well as over old leaves, poor in glucosinolates and nitrogen. In contrast, P. brassicae preferred young leaves. Both species preferred severely drought-stressed plants to the well-watered control, although analyzed glucosinolate concentrations did not differ. Both S. littoralis and P. brassicae feeding induced an increase of indole glucosinolate levels, which may explain a reduced consumption of damaged plants detected for S. littoralis but not for P. brassicae. By revealing distinct, sometimes contrasting responses of two insect herbivores to within-plant and stress-mediated intraspecific variation in phytochemistry of B. oleracea, this study emphasizes the need to consider specific herbivore responses to understand and predict the interactions between herbivores and variable plants.     

溶藻放线菌对铜绿微囊藻和小球藻竞争生长的影响

从土壤中分离获得l株对铜绿微囊藻有明显抑制作用的橄榄网状链霉菌SG-001（Streptomyces olivoreticuli SG-001）,研究其对铜绿微囊藻（Microcystis aeruginosa）和小球藻（Chlorefla pyrenoidosa）竞争生长的影响。结果表明：SG-001菌株的活性物质主要存在于无菌滤液中,能够强烈抑制铜绿微囊藻的生长,但对小球藻的生长具有明显的促进作用。当混藻中两种藻细胞初始接种浓度均为4.0×106 mL-1时,在BG11纯培养条件下,添加SG-001无菌滤液有利于小球藻生长,但对铜绿微囊藻抑制作用不明显;而SG-001无菌滤液对天然加富水样中的铜绿微囊藻具有明显抑制作用,在同时接种有铜绿微囊藻和小球藻的混合藻液中,添加SG-001无菌滤液能够明显提高小球藻的生长竞争能力,且水体中氨氮和可溶性总磷的去除率可分别达到85%和93.33%,而铜绿微囊藻在第8天时生长基本被小球藻抑制。     

Local consumers induce resistance differentially between Spartina populations in the field

Intraspecific variation in the strength of inducible plant defenses plays a central role in the interactions between plants and herbivores. Studies of this variation are typically conducted in the greenhouse or laboratory rather than the field. We simultaneously manipulated densities of local consumers in the field within Maine and South Carolina populations of the smooth cordgrass Spartina alterniflora. South Carolina, but not Maine, plants induced resistance when grazed by local consumers. South Carolina populations of Littoraria snails and planthoppers colonized control more than previously grazed South Carolina plants, and Littoraria snails consumed more control than previously grazed plants. The inducible feeding deterrents in South Carolina plants appear to be water soluble, but not phenolic based. In contrast, grazed and control plants from Maine populations did not differ in attractiveness or palatability to Maine consumers. Thus, inducible plant responses by South Carolina plants had a strong effect on the South Carolina consumer community, but no analogous effect occurred in Maine. Field experiments are a powerful approach to detecting the strength of inducible plant resistance and its impacts on local consumers, which in this case were shown to vary with location.     

Insect herbivory may cause changes in the visual properties of leaves and affect the camouflage of herbivores to avian predators

Behavioral Ecology and Sociobiology - An individual’s metabolic competence is important when escalating to costly behaviours in agonistic encounters. The use of broken shells in the wild...     

Population-level variation in plant secondary chemistry,and the population biology of herbivores

Summary Modern population ecology is becoming increasingly spatially-explicit. For insect hervibores, spatial variation in plant quality is a critical component of distribution and abundance. We argue that populationlevel measurements of phytochemical variation among individual plants has not kept pace with recent developments in population ecology. With examples from our own work, we demonstrate the importance of spatial variation in plant chemistry for insect herbivores, and suggest that phytochemistry should play a central role in the development of any spatially-based ecological theory.     

Assessing competition between Steller sea lions and the commercial groundfishery in Alaska: A bioenergetics modelling approach

In the last three decades the western stock of the Steller sea lion (Eumetopias jubatus) has declined by more than 85%. Nutritional stress resulting in increased juvenile mortality is one of the leading hypotheses to account for this decline. Competition between Steller sea lions and the commercial groundfishery for walleye pollock (Theragra chalcogramma) has been proposed as a mechanism underlying the nutritional stress. In order to examine the competition component of the nutritional stress hypothesis, we developed a bioenergetics-based model to project the population trends of Steller sea lions under various scenarios of continued groundfish harvest. Annual energy budgets were calculated for the Gulf of Alaska population of Steller sea lions, and compared with projected available energy from walleye pollock under a variety of harvest scenarios. Model simulations produced 50-year Steller sea lion population projections consistent with current trends, as well as with published projections for stable and increasing populations from stable age distribution life table models. Model simulations were unable to produce energy deficits sufficient to account for the decline in Steller sea lions, but do suggest areas where existing data need supplementing.     

Chemically mediated competition between microbes and animals: microbes as consumers in food webs

Microbes are known to affect ecosystems and communities as decomposers, pathogens, and mutualists. However, they also may function as classic consumers and competitors with animals if they chemically deter larger consumers from using rich food-falls such as carrion, fruits, and seeds that can represent critical windfalls to both microbes and animals. Microbes often use chemicals (i.e., antibiotics) to compete against other microbes. Thus using chemicals against larger competitors might be expected and could redirect significant energy subsidies from upper trophic levels to the detrital pathway. When we baited traps in a coastal marine ecosystem with fresh vs. microbe-laden fish carrion, fresh carrion attracted 2.6 times as many animals per trap as microbe-laden carrion. This resulted from fresh carrion being found more frequently and from attracting more animals when found. Microbe-laden carrion was four times more likely to be uncolonized by large consumers than was fresh carrion. In the lab, the most common animal found in our traps (the stone crab Menippe mercenaria) ate fresh carrion 2.4 times more frequently than microbe-laden carrion. Bacteria-removal experiments and feeding bioassays using organic extracts of microbe-laden carrion showed that bacteria produced noxious chemicals that deterred animal consumers. Thus bacteria compete with large animal scavengers by rendering carcasses chemically repugnant. Because food-fall resources such as carrion are major food subsidies in many ecosystems, chemically mediated competition between microbes and animals could be an important, common, but underappreciated interaction within many communities.     

Sperm competition in the mite Caloglyphus berlesei

Summary The sperm utilisation pattern of C. berlesei was investigated using the sterile male method. The last male to mate with a female fertilised most of her eggs. In experiments where females mated with two virgin males, second males fertilised on average 86% of eggs when second matings took place immediately after the first. Last male sperm priority was also confirmed in experiments where females mated sequentially with three males. The proportion of eggs fertilised by the second male decreased to 71% and 56% when the interval between matings was 2 and 6 h, respectively. Males that had opportunities to mate prior to experiments copulated for a shorter time than virgin males, and second males fertilised on average 74% eggs.