Mechanisms and implications of bacterial-fungal competition for soil resources.

Elucidating complex interactions between bacteria and fungi that determine microbial community structure, composition, and functions in soil, as well as regulate carbon (C) and nutrient fluxes, is crucial to understand biogeochemical cycles. Among the various interactions, competition for resources is the main factor determining the adaptation and niche differentiation between these two big microbial groups in soil. This is because C and energy limitations for microbial growth are a rule rather than an exception. Here, we review the C and energy demands of bacteria and fungi-the two major kingdoms in soil-the mechanisms of their competition for these and other resources, leading to niche differentiation, and the global change impacts on this competition. The normalized microbial utilization preference showed that bacteria are 1.4-5 times more efficient in the uptake of simple organic compounds as substrates, whereas fungi are 1.1-4.1 times more effective in utilizing complex compounds. Accordingly, bacteria strongly outcompete fungi for simple substrates, while fungi take advantage of complex compounds. Bacteria also compete with fungi for the products released during the degradation of complex substrates. Based on these specifics, we differentiated spatial, temporal, and chemical niches for these two groups in soil. The competition will increase under the main five global changes including elevated CO2, N deposition, soil acidification, global warming, and drought. Elevated CO2, N deposition, and warming increase bacterial dominance, whereas soil acidification and drought increase fungal competitiveness.

Competitive interactions and coexistence of sympatric flagship carnivores in Asia.

Understanding the competition and coexistence of flagship carnivores is key to creating strategies for their conservation in the face of global carnivore declines. Although studies exploring the dynamics and competition between tigers (Panthera tigris) and leopards (P. pardus) span decades, there is a lack of understanding regarding the factors that influence their coexistence mechanisms on a broad scale, as well as the drivers determining their exploitative and interference competition. We gathered a comprehensive list of research papers among which 36 papers explored the interspecific interactions between tigers and leopards and tested the influence of biotic and abiotic factors on the coexistence mechanisms along three dimensions using multiple response variables regression models; we also tested the influence of ecological drivers determining the exploitative or interference competition between tigers and leopards. Elevation and ungulate density were the most important predictors in regulating the coexistence mechanisms. Tigers and leopards exhibited more positive relations/higher overlaps as elevation increased in the spatial niche. In addition, they showed a higher dietary overlap in the prey-rich regions. We determined that interference competition between tigers and leopards was less frequently observed in habitats with dense tree cover and homogeneous vegetation structures. Meanwhile, studies with multiple metrics would promote the detection of interference competition. Our study provides new insight into the competitive interactions and coexistence mechanisms of tigers and leopards on a broad scale. Policy-makers and managers should pay more attention to the factors of elevation, prey abundance, and habitat structures for the conservation of tigers and leopards.

Disentangling Exploitative and Interference Competition on Forest Dwelling Salamanders.

Exploitative competition and interference competition differ in the way access to resources is modulated by a competitor. Exploitative competition implies resource depletion and usually produces spatial segregation, while interference competition is independent from resource availability and can result in temporal niche partitioning. Our aim is to infer the presence of spatial or temporal niche partitioning on a two-species system of terrestrial salamanders in Northern Italy: Speleomantes strinatii and Salamandrina perspicillata. We conducted 3 repeated surveys on 26 plots in spring 2018, on a sampling site where both species are present. We modelled count data with N-mixture models accounting for directional interactions on both abundance and detection process. In this way we were able to disentangle the effect of competitive interaction on the spatial scale, i.e., local abundance, and from the temporal scale, i.e., surface activity. We found strong evidence supporting the presence of temporal niche partitioning, consistent with interference competition. At the same time, no evidence of spatial segregation has been observed.

Decline in wild bee species richness associated with honey bee (Apis mellifera L.) abundance in an urban ecosystem.

The spatial heterogeneity of urban landscapes, relatively low agrochemical use, and species-rich floral communities often support a surprising diversity of wild pollinators in cities. However, the management of Western honey bees (Apis mellifera L.) in urban areas may represent a new threat to wild bee communities. Urban beekeeping is commonly perceived as an environmentally friendly practice or a way to combat pollinator declines, when high-density beekeeping operations may actually have a negative influence on native and wild bee populations through floral resource competition and pathogen transmission. On the Island of Montréal, Canada there has been a particularly large increase in beekeeping across the city. Over the years following a large bee diversity survey ending in 2013, there was an influx of almost three thousand honey bee colonies to the city. In this study, we examined the wild bee communities and floral resources across a gradient of honey bee abundances in urban greenspaces in 2020, and compared the bee communities at the same sites before and after the large influx of honey bees. Overall, we found a negative relationship between urban beekeeping, pollen availability, and wild bee species richness. We also found that honey bee abundance had the strongest negative effect on small (inter-tegular span <2.25 mm) wild bee species richness. Small bee species may be at higher risk in areas with abundant honey bee populations as their limited foraging range may reduce their access to floral resources in times of increased competition. Further research on the influence of urban beekeeping on native and wild pollinators, coupled with evidence-based beekeeping regulations, is essential to ensure cities contain sufficient resources to support wild bee diversity alongside managed honey bees.

Extending the gleaner-opportunist trade-off.

Species exhibit various trade-offs that can result in stable coexistence of competitors. The gleaner-opportunist trade-off to fluctuations in resource abundance is one of the most intuitive, yet also misunderstood, coexistence-promoting trade-offs. Here, we review its history as an ecological concept, discuss extensions to the classical theory and outline opportunities to advance its understanding. The mechanism of coexistence between species that grow relatively faster than their competitors in a low-resource environment (i.e. a gleaner) versus a high-resource environment (i.e. an opportunist) was first proposed in the 1970s. Stable coexistence could emerge between gleaners and opportunists if the opportunist species (dominant in unstable environments) dampens resource fluctuations via relatively convex functional responses, while the gleaner species (dominant in stable environments) promotes fluctuations, or diminishes them less than the opportunist does, via relatively saturating functional responses. This fluctuation-dependent coexistence mechanism has since been referred to by various names, including the Armstrong-McGehee mechanism and relative nonlinearity of competition. Several researchers have argued this mechanism likely plays a relatively minor role in species coexistence owing in part to the restricted range of conditions that allow it to operate. More recent theoretical research, however, suggests that relative nonlinearity can operate over wider conditions than previously thought. Here, we identify several novel, or little explored, extensions to the gleaner-opportunist trade-off that can yield species coexistence under phenomena as diverse as fluctuations in predation/pathogen pressure, multiple resources, phenotypic plasticity and rapid evolution, amongst other phenomena. While the original definition of the gleaner-opportunist trade-off may be imperfect as a collective for these extensions, we argue that a subtle reframing of the trade-off focusing on species' performance in equilibrium versus fluctuating conditions (irrespective of preferences for high or low resources, predation pressure or other competitive factors) reveals their fundamental commonality in stable coexistence via relative nonlinearity. An extended framing shines a light on the potential ubiquity of this canonical trade-off in nature and on the breadth of theoretical and empirical terrain that remains to be trodden.

Statistical evidence that honeybees competitively reduced wild bee abundance in the Munich Botanic Garden in 2020 compared to 2019.

In a commentary on our paper (Renner et al., Oecologia 195:825-831, 2021), Harder and Miksha lay out why they think that our finding of higher honeybee abundances reducing wild bee abundances in an urban botanical garden is not statistically supported. Here, we explain the statistical test provided in our paper, which took advantage of a natural experiment offered by 2019 being a poorer year for bee keeping than 2020.

Extrinsic Inter- and Intraspecific Competition in Parasitoid Wasps.

The diverse ecology of parasitoids is shaped by extrinsic competition, i.e., exploitative or interference competition among adult females and males for hosts and mates. Adult females use an array of morphological, chemical, and behavioral mechanisms to engage in competition that may be either intra- or interspecific. Weaker competitors are often excluded or, if they persist, use alternate host habitats, host developmental stages, or host species. Competition among adult males for mates is almost exclusively intraspecific and involves visual displays, chemical signals, and even physical combat. Extrinsic competition influences community structure through its role in competitive displacement and apparent competition. Finally, anthropogenic changes such as habitat loss and fragmentation, invasive species, pollutants, and climate change result in phenological mismatches and range expansions within host-parasitoid communities with consequent changes to the strength of competitive interactions. Such changes have important ramifications not only for the success of managed agroecosystems, but also for natural ecosystem functioning.

Competition between obligate and facultative scavengers and infection: vulture-jackal-anthrax dynamics in Etosha National Park.

Different species of scavengers may compete for the same food in an ecosystem. This case study considers the competition between jackals and vultures in Etosha National Park in Namibia. While jackals are facultative scavengers, able to hunt for food if necessary, vultures are obligate scavengers wholly dependent on carcasses of animals like zebras for persistence. This competition may be further affected by outbreaks of infections such as anthrax, which temporarily increase the number of carcasses but lower the zebra population, acting in some ways as a third competitor. We use a dynamical system to model the interplay between competition dynamics and infection dynamics, and how it is affected by the nature of the competition: indirect (exploitative) or direct (interference). A bifurcation analysis using reproduction numbers shows how vultures' survival may depend on their direct competitive edge in reaching carcasses faster than jackals, and how the infection and the scavengers complicate each other's persistence. Vultures' interference causes a backward bifurcation which enables them to persist. One possible outcome is a "strange bedfellows" bistability in which anthrax and vultures persist only together, not apart, despite being competitors.

Contrasting population manipulations reveal resource competition between two large marsupials: bare-nosed wombats and eastern grey kangaroos.

Resource competition is an important interaction that can structure ecological communities, but is difficult to demonstrate in nature, and rarely demonstrated for large mammals including marsupials. We analysed 10 years of population survey data to investigate resource competition between bare-nosed wombats (Vombatus ursinus) and eastern grey kangaroos (Macropus giganteus) at two sites to assess whether resource competition is occurring. At one site, wombat abundance was reduced by increased mortality from mange disease, whereas at the other site, kangaroo abundance was reduced primarily by culling. We used the modified Lotka-Volterra competition (LVC) models to describe the mechanism of resource competition and fitted those models to the empirical data by maximum likelihood estimation. We found strong negative relationships between the abundance of wombats and kangaroos at each site, and resource competition was also mechanistically supported by the modified LVC models. The estimated competition coefficients indicate that bare-nosed wombats are a slightly superior competitor of eastern grey kangaroos than vice versa, and that intraspecific competition is almost twice as strong as interspecific competition. In addition, this study facilitated the calculation of the transmission rate associated with mange disease at one site (0.011), and the removal rate owing to culling, the introduction of a predator species, and drought at the other site (0.0006). Collectively, this research represents a rare empirical demonstration of resource competition between large mammals and contributes new insight into the ecology of two of Australia's largest grazing marsupials.

Community science data suggests that urbanization and forest habitat loss threaten aphidophagous native lady beetles.

Community scientists have illustrated rapid declines of several aphidophagous lady beetle (Coccinellidae) species. These declines coincide with the establishment of alien coccinellids. We established the Buckeye Lady Beetle Blitz program to measure the seasonal occupancy of coccinellids within gardens across a wide range of landscape contexts. Following the Habitat Compression Hypothesis, we predicted that gardens within agricultural landscapes would be alien-dominated, whereas captures of natives would be higher within landscapes encompassing a high concentration of natural habitat.Within the state of Ohio, USA, community scientists collected lady beetles for a 7-day period across 4 years in June and August using yellow sticky card traps. All identifications were verified by professional scientists and beetles were classified by three traits: status (alien or native), mean body length, and primary diet. We compared the relative abundance and diversity of coccinellids seasonally and determined if the distribution of beetles by size, status, and diet was related to landscape features.Alien species dominated the aphidophagous fauna. Native aphidophagous coccinellid abundance was positively correlated with forest habitat while alien species were more common when gardens were embedded within agricultural landscapes. Urbanization was negatively associated with both aphidophagous alien and native coccinellids. Synthesis and Applications: Our census of native coccinellid species within residential gardens-a widespread and understudied habitat-was enabled by volunteers. These data will serve as an important baseline to track future changes within coccinellid communities within this region. We found that native coccinellid species richness and native aphidophagous coccinellid abundance in gardens were positively associated with forest habitat at a landscape scale of 2 km. However, our understanding of when and why (overwintering, summer foraging, or both) forest habitats are important remains unclear. Our findings highlight the need to understand how declining aphidophagous native species utilize forest habitats as a conservation priority.

The evolution of the irreversible transition from a free-swimming state to an immobile sessile state in aquatic invertebrates modelled in a chemostat.

To better understand the environmental factors and ecological processes underlying the evolution of the irreversible transition from a free-swimming state to an immobile sessile state as seen in many aquatic invertebrates, we study the adaptive dynamics of the settling rate of a hypothetical microorganism onto the wall of a chemostat. The two states, floating or settled, differ in their nutrient ingestion, reproduction and death rate. We consider three different settling mechanisms involving competition for space on the wall: (i) purely exploitative competition where free-swimming individuals settle in vacant space only, (ii) mixed exploitative and interference competition where individuals attempt to settle in any place but fail and die if the space is already occupied, and (iii) mixed exploitative and interference competition, but now settling in occupied space is successful and the former occupant dies. In the simplified environment of the chemostat, the input concentration of nutrients and the dilution rate of the tank are the main environmental control variables. Using the theory of adaptive dynamics, we find that the settling mechanisms and environmental control variables have qualitatively different effects on the evolution of the settling rate in terms of the direction of evolution as well as on species diversity. In the case of purely exploitative competition a small change in the settings of the environmental control variables can lead to an abrupt reversal of the direction of evolution, while in the case of mixed exploitative and interference competition the effect is gradual. For all three settling mechanisms, periodic fluctuations in the nutrient input open the possibility of evolutionary branching leading to the long-term coexistence of an intermediate and an infinitely high settling rates (in the case of low-frequency fluctuations), and an intermediate and a zero settling rates (in the case of high-frequency fluctuations).

High honeybee abundances reduce wild bee abundances on flowers in the city of Munich.

The increase in managed honeybees (Apis mellifera) in many European cities has unknown effects on the densities of wild bees through competition. To investigate this, we monitored honeybees and non-honeybees from 01 April to 31 July 2019 and 2020 at 29 species of plants representing diverse taxonomic and floral-functional types in a large urban garden in the city of Munich in which the same plant species were cultivated in both years. No bee hives were present in the focal garden, and all bee hives in the adjacent area were closely monitored by interviewing the relevant bee keepers in both 2019 and 2020. Honeybee numbers were similar in April of both years, but increased from May to July 2020 compared to 2019. The higher densities correlated with a significant increase in shifts from wild bee to honeybee visits in May/June/July, while visitor spectra in April 2019 and 2020 remained the same. Most of the species that experienced a shift to honeybee visits in 2020 were visited mostly or exclusively for their nectar. There were no shifts towards increased wild bee visits in any species. These results from a flower-rich garden have implications for the discussion of whether urban bee keeping might negatively impact wild bees. We found clear support that high honeybee densities result in exploitative competition at numerous types of flowers.

Plant-animal interactions between carnivorous plants, sheet-web spiders, and ground-running spiders as guild predators in a wet meadow community.

Plant-animal interactions are diverse and widespread shaping ecology, evolution, and biodiversity of most ecological communities. Carnivorous plants are unusual in that they can be simultaneously engaged with animals in multiple mutualistic and antagonistic interactions including reversed plant-animal interactions where they are the predator. Competition with animals is a potential antagonistic plant-animal interaction unique to carnivorous plants when they and animal predators consume the same prey.The goal of this field study was to test the hypothesis that under natural conditions, sundews and spiders are predators consuming the same prey thus creating an environment where interkingdom competition can occur.Over 12 months, we collected data on 15 dates in the only protected Highland Rim Wet Meadow Ecosystem in Kentucky where sundews, sheet-web spiders, and ground-running spiders co-exist. One each sampling day, we attempted to locate fifteen sites with: (a) both sheet-web spiders and sundews; (b) sundews only; and (c) where neither occurred. Sticky traps were set at each of these sites to determine prey (springtails) activity-density. Ground-running spiders were collected on sampling days. DNA extraction was performed on all spiders to determine which individuals had eaten springtails and comparing this to the density of sundews where the spiders were captured.Sundews and spiders consumed springtails. Springtail activity-densities were lower, the higher the density of sundews. Both sheet-web and ground-running spiders were found less often where sundew densities were high. Sheet-web size was smaller where sundew densities were high.The results of this study suggest that asymmetrical exploitative competition occurs between sundews and spiders. Sundews appear to have a greater negative impact on spiders, where spiders probably have little impact on sundews. In this example of interkingdom competition where the asymmetry should be most extreme, amensalism where one competitor experiences no cost of interaction may be occurring.

Contrasting forms of competition set elevational range limits of species.

How abiotic and biotic factors constrain distribution limits at the harsh and benign edges of species ranges is hotly debated, partly because macroecological experiments testing the proximate causes of distribution limits are scarce. It has long been recognized - at least since Darwin's On the Origin of Species - that a harsh climate strengthens competition and thus sets species range limits. Using thorough field manipulations along a large elevation gradient, we show the mechanisms by which temperature determines competition type, resulting in a transition from interference to exploitative competition from the lower to the upper elevation limits in burying beetles (Nicrophorus nepalensis). This transition is an example of Darwin's classic hypothesis that benign climates favor direct competition for highly accessible resources while harsh climates result in competition through resources of high rivalry. We propose that identifying the properties of these key resources will provide a more predictive framework to understand the interplay between biotic and abiotic factors in determining geographic range limits.

Beyond replicator dynamics: From frequency to density dependent models of evolutionary games.

Game theoretic models of evolution such as the Hawk-Dove game assume that individuals gain fitness (which is a proxy of the per capita population growth rate) in pair-wise contests only. These models assume that the equilibrium distribution of phenotypes involved (e.g., Hawks and Doves) in the population is given by the Hardy-Weinberg law, which is based on instantaneous, random pair formation. On the other hand, models of population dynamics do not consider pairs, newborns are produced by singles, and interactions between phenotypes or species are described by the mass action principle. This article links game theoretic and population approaches. It shows that combining distribution dynamics with population dynamics can lead to stable coexistence of Hawk and Dove population numbers in models that do not assume a priori that fitness is negative density dependent. Our analysis shows clearly that the interior Nash equilibrium of the Hawk and Dove model depends both on population size and on interaction times between different phenotypes in the population. This raises the question of the applicability of classic evolutionary game theory that requires all interactions take the same amount of time and that all single individuals have the same payoff per unit of time, to real populations. Furthermore, by separating individual fitness into birth and death effects on singles and pairs, it is shown that stable coexistence in these models depends on the time-scale of the distribution dynamics relative to the population dynamics. When explicit density-dependent fitness is included through competition over a limited resource, the combined dynamics of the Hawk-Dove model often lead to Dove extinction no matter how costly fighting is for Hawk pairs.

Resisting annihilation: relationships between functional trait dissimilarity, assemblage competitive power and allelopathy.

Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within-assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real-world time-series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.

Competition between sympatric wolf taxa: an example involving African and Ethiopian wolves.

Carnivore populations are declining globally due to range contraction, persecution and prey depletion. One consequence of these patterns is increased range and niche overlap with other carnivores, and thus an elevated potential for competitive exclusion. Here, we document competition between an endangered canid, the Ethiopian wolf (EW), and the newly discovered African wolf (AW) in central Ethiopia. The diet of the ecological specialist EW was dominated by rodents, whereas the AW consumed a more diverse diet also including insects and non-rodent mammals. EWs used predominantly intact habitat, whereas AWs used mostly areas disturbed by humans and their livestock. We observed 82 encounters between the two species, of which 94% were agonistic. The outcomes of agonistic encounters followed a territory-specific dominance pattern, with EWs dominating in intact habitat and AWs in human-disturbed areas. For AWs, the likelihood of winning encounters also increased with group size. Rodent species consumed by EWs were also available in the human-disturbed areas, suggesting that these areas could be suitable habitat for EWs if AWs were not present. Increasing human encroachment not only affects the prey base of EWs, but also may impact their survival by intensifying competition with sympatric AWs.

An interaction-driven cannibalistic reaction norm.

Cannibalism is induced in larval-stage populations of the Hokkaido salamander, Hynobius retardatus, under the control of a cannibalism reaction norm. Here, I examined phenotypic expression under the cannibalism reaction norm, and how the induction of a cannibalistic morph under the norm leads to populational morphological diversification. I conducted a set of experiments in which density was manipulated to be either low or high. In the high-density treatment, the populations become dimorphic with some individuals developing into the cannibal morph type. I performed an exploratory analysis based on geometric morphometrics and showed that shape characteristics differed between not only cannibal and noncannibal morph types in the high-density treatment but also between those morph types and the solitary morph type in the low-density treatment. Size and shape of cannibal and noncannibal individuals were found to be located at either end of a continuum of expression following a unique size-shape integration rule that was different from the rule governing the size and shape variations of the solitary morph type. This result implies that the high-density-driven inducible morphology of an individual is governed by a common integration rule during the development of dimorphism under the control of the cannibalism reaction norm. Phenotypic expression under the cannibalism reaction norm is driven not only by population density but also by social interactions among the members of a population: variation in the populational expression of dimorphism is associated with contingent social interaction events among population members. The induced cannibalistic morph thus reflects not only by contest-type exploitative competition but also interference competition.

Examining spatial patterns of selection and use for an altered predator guild.

Anthropogenic disturbances have altered species' distributions potentially impacting interspecific interactions. Interference competition is when one species denies a competing species access to a resource. One mechanism of interference competition is aggression, which can result in altered space-use of a subordinate species due to the threat of harm, otherwise known as a 'landscape of fear'. Alternatively, subordinates might outcompete dominant species in resource-poor environments via a superior ability to extract resources. Our goal was to evaluate spatial predictions of the 'landscape of fear' hypothesis for a carnivore guild in Newfoundland, Canada, where coyotes recently immigrated. Native Newfoundland carnivores include red foxes, Canada lynx, and black bears. We predicted foxes and lynx would avoid coyotes because of their larger size and similar dietary niches. We used scat-detecting dogs and genetic techniques to locate and identify predator scats. We then built resource selection functions and tested for avoidance by incorporating predicted values of selection for the alternative species into the best supported models of each species. We found multiple negative relationships, but notably did not find avoidance by foxes of areas selected by coyotes. While we did find that lynx avoided coyotes, we also found a reciprocal relationship. The observed patterns suggest spatial partitioning and not coyote avoidance, although avoidance could still be occurring at different spatial or temporal scales. Furthermore, Newfoundland's harsh climate and poor soils may swing the pendulum of interspecific interactions from interference competition to exploitative competition, where subordinates outcompete dominant competitors through a superior ability to extract resources.

A window of opportunity: Subdominant predators can use suboptimal prey.

Introduced species have been linked to declines of native species through mechanisms including intraguild predation and exploitative competition. However, coexistence among species may be promoted by niche partitioning if native species can use resources that the invasive species cannot. Previous research has shown that some strains of the aphid Aphis craccivora are toxic to a competitively dominant invasive lady beetle, Harmonia axyridis. Our objective was to investigate whether these aphids might be an exploitable resource for other, subdominant, lady beetle species. We compared larval development rate, survival, and adult weight of five lady beetle species in no-choice experiments with two different strains of A. craccivora, one of which is toxic to H. axyridis and one that is nontoxic. Two lady beetle species, Cycloneda munda and Coleomegilla maculata, were able to complete larval development when feeding on the aphid strain that is toxic to H. axyridis, experiencing only slight developmental delays relative to beetles feeding on the other aphid strain. One species, Coccinella septempunctata, also was able to complete larval development, but experienced a slight reduction in adult weight. The other two lady beetle species, Hippodamia convergens and Anatis labiculata, demonstrated generally low survivorship when consuming A. craccivora, regardless of aphid strain. All five species showed increased survival and/or development relative to H. axyridis on the "toxic" aphid strain. Our results suggest that this toxic trait may act as a narrow-spectrum defense for the aphids, providing protection against only some lady beetle enemies. For other less-susceptible lady beetles, these aphids have the potential to provide competitive release from the otherwise dominant H. axyridis.