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.

Intra- and interspecific interactions in the two coexisting Locustella warblers revealed by song playback experiments.

Males usually come into conflict due to competition for territories and females. However, interference competition can also occur between males of congeneric species when their ecological requirements are overlapping. Using acoustic playback experiments, we investigated male-male interactions within and between Grasshopper (Locustella naevia; GW) and River Warbler (L. fluviatilis; RW). Our objective was to evaluate the song and behavioural response of tested males of both species to conspecific song stimuli in order to compare this with the response to congeneric stimulus, based on which we could assess whether these two commonly co-existing species show interspecific territorialism. A total of nine GW and 11 RW males were tested in May and June 2019 in western Slovakia. The ability to differentiate between the heterospecific (control), congeneric, and conspecific stimuli was similar between the two species. Conspecific playback elicited the strongest non-vocal response and a significant change in vocalization. The GW males shortened the songs, while the RW males shortened the songs and also increased their syllable rate. The congeneric playback elicited a lower intensity of behavioural response than conspecific playback and no change in vocalization in either species. We conclude that interspecific interference competition between GW and RW is rather low, suggesting that the species' ecological requirements are separated, although these two congeneric species commonly share habitat.

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.

Interspecific killing of wolverines by one wolf pack.

Interactions between different species of predators are not uncommon, yet they are generally understudied in North America. Across their range, gray wolves (Canis lupus) and wolverines (Gulo gulo) occupy similar habitats and dietary niches. However, due to the elusiveness and relatively low density of these two species, interactions between them are not well documented. Here, we describe three instances of a single wolf pack killing a wolverine in the span of 13 months. None of the wolverines killed by wolves were consumed, suggesting that food was not the primary motivation behind the killings. Alternatively, defense of a food resource, territoriality, interspecific competitive killing, or some combination of those behaviors appear to be the cause of these actions. Documentation of these occurrences improves our understanding of wolf and wolverine ecology, interspecific predator interactions, and potential future changes to this aspect of community ecology.

Investigating avian competition for surface water in an arid zone bioregion.

Interference competition has the potential to alter avian assemblages at long-lasting arid zone waterholes, particularly in a warming world, as more potentially aggressive species frequent these sites to drink. We used camera traps and observational surveys to investigate interference competition between terrestrial avian species at six long-lasting waterholes across three sampling seasons (two summers and one winter) within the MacDonnell Ranges Bioregion in central Australia. The proportion of individuals drinking for each of four dietary classes (granivores, nectarivores, omnivores, and insectivores) was modelled in relation to their abundance in the immediate waterhole habitat, which informed the potential for competition in each season. We then used the temporal overlap estimators to quantify the degree of competition between species at waterholes with species grouped into families (Meliphagidae, Ptilonorhynchidae, Estrildidae, and Rhipiduridae). We found the proportion of individuals drinking at waterholes was greatest during hot and dry periods, suggesting the potential for interference competition is greatest during these times. This was particularly the case for nectarivores where, in hot and dry conditions, the proportion of drinking individuals increased significantly as their abundance also increased in the waterhole habitat. We predicted that subordinate species would alter their activity periods to avoid competitive interactions with meliphagids (honeyeaters), however, we found there was a high degree of temporal overlap between all families sampled across all seasons. These results suggest subordinate species are unlikely to be excluded from long-lasting waterholes by potentially aggressive species, such as honeyeaters. However, some species may face trade-offs between foraging and accessing waterholes to stay hydrated as they shift their activity to avoid the hottest parts of the day during the summer months. Under global warming, extended hot and dry periods will likely create conditions where balancing energy and hydration requirements becomes increasingly difficult and results in the loss of body condition.

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.

Size-Mediated Trophic Interactions in Two Syntopic Forest Salamanders.

Exploitative competition and interference competition differ in the way they affect re-source availability for competitors: in the former, organisms reduce resource availability for the competitors; in the latter, one organism actively prevents the competitor from accessing resources, independently of their availability. Our aim is to test for the presence of foraging competition in two forest-dwelling salamanders in Italy: Speleomantes strinatii and Salamandrina perspicillata. We also aim at testing for size-mediated competition. We obtained stomach contents from 191 sampled individuals by means of stomach flushing at 8 sampling sites where both species occur. We focused our analysis on the core prey taxa shared by both species: Collembola and Acarina. We found that the foraging activity of S. perspicillata is positively affected by body size and negatively affected by potential competitor's activity on the forest floor during the sampling, which also significantly weakened the positive relationship with body size. These results suggest the presence of an interference/interaction occurring between the two species and affecting the foraging activity of S. perspicillata. This competitive interaction is size mediated and configured as interference competition rather than exploitative competition.

A coevolution experiment between Flavobacterium johnsoniae and Burkholderia thailandensis reveals parallel mutations that reduce antibiotic susceptibility.

One interference mechanism of bacterial competition is the production of antibiotics. Bacteria exposed to antibiotics can resist antibiotic inhibition through intrinsic or acquired mechanisms. Here, we performed a coevolution experiment to understand the long-term consequences of antibiotic production and antibiotic susceptibility for two environmental bacterial strains. We grew five independent lines of the antibiotic-producing environmental strain, Burkholderia thailandensis E264, and the antibiotic-inhibited environmental strain, Flavobacterium johnsoniae UW101, together and separately on agar plates for 7.5 months (1.5 month incubations), transferring each line five times to new agar plates. We observed that the F. johnsoniae ancestor could tolerate the B. thailandensis-produced antibiotic through efflux mechanisms, but that the coevolved lines had reduced susceptibility. We then sequenced genomes from the coevolved and monoculture F. johnsoniae lines, and uncovered mutational ramifications for the long-term antibiotic exposure. The coevolved genomes from F. johnsoniae revealed four potential mutational signatures of reduced antibiotic susceptibility that were not observed in the evolved monoculture lines. Two mutations were found in tolC: one corresponding to a 33 bp deletion and the other corresponding to a nonsynonymous mutation. A third mutation was observed as a 1 bp insertion coding for a RagB/SusD nutrient uptake protein. The last mutation was a G83R nonsynonymous mutation in acetyl-coA carboxylayse carboxyltransferase subunit alpha (AccA). Deleting the 33 bp from tolC in the F. johnsoniae ancestor reduced antibiotic susceptibility, but not to the degree observed in coevolved lines. Furthermore, the accA mutation matched a previously described mutation conferring resistance to B. thailandensis-produced thailandamide. Analysis of B. thailandensis transposon mutants for thailandamide production revealed that thailandamide was bioactive against F. johnsoniae, but also suggested that additional B. thailandensis-produced antibiotics were involved in the inhibition of F. johnsoniae. This study reveals how multi-generational interspecies interactions, mediated through chemical exchange, can result in novel interaction-specific mutations, some of which may contribute to reductions in antibiotic susceptibility.

Relative influence of inter- and intraspecific competition in an ungulate assemblage modified by introduced species.

Interspecific competition from introduced and naturally colonizing species has potential to affect resident populations, but demographic consequences for vertebrates have rarely been tested. We tested hypotheses of interspecific and intraspecific competition for density, body mass, and fertility of adult female Roe Deer (Capreolus capreolus) across a heterogeneous forest landscape occupied by two introduced deer species: Mediterranean Fallow Deer (Dama dama); and subtropical Reeve's Muntjac (Muntiacus reevesi). Species-specific deer densities in buffers around culling locations of 492 adult female Roe Deer, sampled over seven years, were extracted from spatially explicit models calibrated through annual nocturnal distance sampling. Roe Deer fertility and body mass were related to species-specific deer densities and extent of arable lands using piecewise structural equation models. Reeve's Muntjac density was lower at higher Fallow Deer densities, suggesting interspecific avoidance via interference competition, but greater when buffers included more arable land. Roe Deer body mass was marginally greater when buffers included more arable land and was independent of deer densities. However, Roe Deer fertility was unrelated to female body mass, suggesting that fertility benefits exceeded an asymptotic threshold of body condition in this low-density population. However, Roe Deer fertility was slightly greater rather than reduced in areas with greater local Roe Deer density, suggesting negligible intraspecific competition. In contrast, Roe Deer was less fertile in areas with greater Reeve's Muntjac densities; thus, interspecific exceeded intraspecific competition in this assemblage. In contrast, we found no support for any effects of Fallow Deer density on Roe Deer density, body mass, or fertility. Complex networks of interspecific competition operating in this deer assemblage include: interspecific interference from Fallow Deer exceeded habitat effects for Reeve's Muntjac; and interspecific competition from introduced, smaller sedentary Reeve's Muntjac reduced fertility, unlike intraspecific, or potential competition with larger, more mobile, Fallow Deer for native Roe Deer. Mechanisms driving Roe Deer fertility may include interspecific behavioral interference or stress-resource depletion is considered less likely because Roe Deer fertility was independent of body mass. Findings emphasize the importance of ensuring appropriate management strategies for controlling invasive species.

Litter, Plant Competition, and Ecosystem Dynamics: A Theoretical Perspective.

AbstractCommunity structure depends jointly on species' responses to, and effects on, environmental factors. Many such factors, including detritus, are studied in ecosystem ecology. Detritus in terrestrial ecosystems is dominated by plant litter (nonliving organic material), which, in addition to its role in material cycling, can act as a niche factor modulating interactions among plants. Litter thus links traditional community and ecosystem processes, which are often studied separately. We explore this connection using population dynamics models of two plant species and a litter pool. We first find conditions determining the outcome of interactions between these species, highlighting the role that litter plays and the role of broader ecosystem parameters, such as decomposition rate. Species trade-offs in tolerance to direct competition and litter-based interference competition allow for coexistence, provided the litter-tolerant species produces more litter at the population level; otherwise, priority effects may result. When species coexist, litter-mediated interactions between plants disrupt the traditional relationship between biomass accumulation and decomposition. Increasing decomposition rate may have no effect on standing litter density and, in some cases, may even increase litter load. These results illustrate how ecosystem variables can influence community outcomes that then feed back to influence the ecosystem.

Eco-evolutionary dynamics of interference competition.

Theorists have identified several mechanisms through which species that compete exploitatively for resources could coexist. By contrast, under the current theory, interference competitors could coexist only in rare circumstances. Yet, some types of interference competition, such as interspecific territoriality, are common. This mismatch between theory and nature inspired us to model interference competition in an eco-evolutionary framework. We based the model on the life cycle of territorial birds and ran simulations to examine whether natural selection could rescue a superior interference competitor from extinction without driving a superior exploitative competitor extinct. We found that coexistence between interference competitors can occur over a wide range of ecologically plausible scenarios, and up to the highest levels of resource overlap. An important caveat is that coexistence requires the species to co-evolve. Reductions in population size and levels of genetic variation could destabilise coexistence between interference competitors, and thereby increase extinction rates over current estimates.

Coexistence under Hierarchical Resource Exploitation: The Role of the R*-Preemption Trade-Off.

AbstractResource competition theory predicts coexistence and exclusion patterns based on species' R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesized that coexistence under preemption requires an R*-preemption trade-off-that is, the species with the priority access should have a higher R* (lower "efficiency"). Thus, we developed an extension of resource competition theory to investigate partial and total preemption (in the latter, the preemptor is unaffected by species with lower preemption rank). We found that an R*-preemption trade-off is a necessary condition for coexistence in all models. Moreover, under total preemption, the trade-off alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discuss the implications of our finding for seemingly distinct trade-offs, which we view as special cases of the R*-preemption trade-off. These trade-offs include the digger-grazer trade-off, the competition-colonization trade-off, and trade-offs related to light competition between trees and understories.

Niche partitioning in a guild of invasive mammalian predators.

Predators compete aggressively for resources, establishing trophic hierarchies that influence ecosystem structure. Competitive interactions are particularly important in invaded ecosystems where introduced predators can suppress native prey species. We investigated whether niche partitioning exists within a guild of invasive mammalian predators and determined the consequences for native species. Over 4405 camera-trap days, we assessed interactions among three invasive predators: two apex predators (feral cats Felis catus and ferrets Mustela furo) and a mesopredator (stoats Mustela erminea), in relation to their primary prey (lagomorphs, rodents and birds) and habitat use. Further, we tested for mesopredator release by selectively removing cats and ferrets in a pulse perturbation experiment. We found compelling evidence of niche partitioning; spatiotemporal activity of apex predators maximized access to abundant invasive prey, with ferrets targeting lagomorphs and cats targeting rodents. Mesopredators adjusted their behavior to reduce the risk of interference competition, thereby restricting access to abundant prey but increasing predation pressure on diurnal native birds. Stoats were only recorded at the treatment site after both larger predators were removed, becoming the most frequently detected predator at 6 months post-perturbation. We suggest there is spatial and resource partitioning within the invasive predator guild, but that this is incomplete, and avoidance is achieved by temporal partitioning within overlapping areas. Niche partitioning among invasive predators facilitates coexistence, but simultaneously intensifies predation pressure on vulnerable native species.

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.

Habitat Selection and Activity Patterns of Japanese Serows and Sika Deer with Currently Sympatric Distributions.

The Japanese serow (Capricornis crispus) and sika deer (Cervus nippon) in Japan are usually allopatric. However, a recent expansion in the distribution range of sika deer, combined with an increase in abundance, has resulted in an overlap of the distribution ranges of the two species. We examined the habitat selection and activity patterns of Japanese serows and sika deer using camera traps placed at 83 sites within a 210 km2 study area, where the distribution range of these two species has recently overlapped. Although both species were photographed throughout the study area, we observed a low spatial overlap between them. The Japanese serows selected steep slopes, and the sika deer selected areas far away from human settlements. In addition, the Japanese serows and sika deer exhibited temporal partitioning with diurnal and crepuscular activity patterns, respectively. The observed partitioning could be explained by differences in their species-specific habitat selections, rather than competition, because the photographic capture rate of the Japanese serows was not affected by that of the sika deer and vice versa. These partitioning behaviors are likely to reduce the rate of encounters between the two ungulates, which enables their coexistence considering the sika deer density remains low.

A genetically based ecological trade-off contributes to setting a geographic range limit.

Understanding the ecological factors that shape geographic range limits and the evolutionary constraints that prevent populations from adaptively evolving beyond these limits is an unresolved question. Here, we investigated why the euryhaline fish, Poecila reticulata, is confined to freshwater within its native range, despite being tolerant of brackish water. We hypothesised that competitive interactions with a close relative, Poecilia picta, in brackish water prevents P. reticulata from colonising brackish water. Using a combination of field transplant, common garden breeding, and laboratory behaviour experiments, we find support for this hypothesis, as P. reticulata are behaviourally subordinate and have lower survival in brackish water with P. picta. We also found a negative genetic correlation between P. reticulata growth in brackish water versus freshwater in the presence of P. picta, suggesting a genetically based trade-off between salinity tolerance and competitive ability could constrain adaptive evolution at the range limit.

Competition-based screening helps to secure the evolutionary stability of a defensive microbiome.

BACKGROUND: The cuticular microbiomes of Acromyrmex leaf-cutting ants pose a conundrum in microbiome biology because they are freely colonisable, and yet the prevalence of the vertically transmitted bacteria Pseudonocardia, which contributes to the control of Escovopsis fungus garden disease, is never compromised by the secondary acquisition of other bacterial strains. Game theory suggests that competition-based screening can allow the selective recruitment of antibiotic-producing bacteria from the environment, by providing abundant resources to foment interference competition between bacterial species and by using Pseudonocardia to bias the outcome of competition in favour of antibiotic producers. RESULTS: Here, we use RNA-stable isotope probing (RNA-SIP) to confirm that Acromyrmex ants can maintain a range of microbial symbionts on their cuticle by supplying public resources. We then used RNA sequencing, bioassays, and competition experiments to show that vertically transmitted Pseudonocardia strains produce antibacterials that differentially reduce the growth rates of other microbes, ultimately biassing the bacterial competition to allow the selective establishment of secondary antibiotic-producing strains while excluding non-antibiotic-producing strains that would parasitise the symbiosis. CONCLUSIONS: Our findings are consistent with the hypothesis that competition-based screening is a plausible mechanism for maintaining the integrity of the co-adapted mutualism between the leaf-cutting ant farming symbiosis and its defensive microbiome. Our results have broader implications for explaining the stability of other complex symbioses involving horizontal acquisition.

Microgeographic evolution of metabolic physiology in a salamander metapopulation.

The Metabolic Theory of Ecology explains ecological variation spanning taxonomic organization, space, and time based on universal physiological relationships. The theory depends on two core parameters: the normalization constant, a mass-independent measure of metabolic rate expected to be invariant among similar species, and the scaling coefficient, a measure of metabolic change with body mass commonly assumed to follow the universal 3/4 scaling law. However, emerging evidence for adaptive microevolution of metabolic rates led us to hypothesize that metabolic rate might exhibit evolved variation among populations on microgeographic scales. To evaluate our hypothesis, we explored evidence for evolved variation in the scaling coefficient and normalization constant within a spotted salamander (Ambystoma maculatum) metapopulation in Connecticut, USA. We measured standard metabolic rate in common-garden raised spotted salamanders from 22 different populations and tested for the effects of six ecological variables suspected in advance to select for divergent physiology. We found that metabolic rate rose with body mass with a log-log slope of 0.97 that was statistically different from the expected 3/4 scaling law. Although we found no evidence for interpopulation variation in the scaling coefficient, we found evidence for interpopulation variation in the normalization constants among populations. Metabolic variation was best explained by differences in population density among ponds. Our results provide mixed support for Metabolic Theory of Ecology assumptions about parameter invariance and illustrate how fundamental physiological processes such as metabolic rate can evolve across microgeographic spatial scales.

Resource availability and heterogeneity shape the self-organisation of regular spatial patterning.

Explaining large-scale ordered patterns and their effects on ecosystem functioning is a fundamental and controversial challenge in ecology. Here, we coupled empirical and theoretical approaches to explore how competition and spatial heterogeneity govern the regularity of colony dispersion in fungus-farming termites. Individuals from different colonies fought fiercely, and inter-nest distances were greater when nests were large and resources scarce-as expected if competition is strong, large colonies require more resources and foraging area scales with resource availability. Building these principles into a model of inter-colony competition showed that highly ordered patterns emerged under high resource availability and low resource heterogeneity. Analysis of this dynamical model provided novel insights into the mechanisms that modulate pattern regularity and the emergent effects of these patterns on system-wide productivity. Our results show how environmental context shapes pattern formation by social-insect ecosystem engineers, which offers one explanation for the marked variability observed across ecosystems.