Strong Intraspecific Competition Research Articles

Dietary Niche Variation in an Invasive Omnivore: The Effects of Habitat on Feral Pig Resource Use in Hawai'i.

Invasive omnivores may have profound impacts on ecological communities through diet selection, particularly when their functional roles differ from those in their native range. While the threat of feral pigs (Sus scrofa) to native plant communities in Hawai'i is well known, their trophic dynamics and the drivers of variation in their diet remain understudied. We investigated the feral pig trophic niche on Hawai'i Island using stable isotopes (13C and 15N) and Bayesian mixing models to identify drivers of variation in resource use. We also reconstructed intra-individual variability for six subsampled individuals to understand temporal variation in resource use and individual diet specialization. Our results revealed that feral pigs on Hawai'i Island exhibit a broad trophic niche characterized by diverse diets, with substantial overlap in resource use across districts and habitats. Differences in dietary composition in the transition from forest to open habitat were driven primarily by a decline in invertebrates and an increasing reliance on resources enriched in 15N, which may reflect a shift in protein sources with habitat. Pigs in forested areas exhibited a smaller trophic niche than those in open habitats, largely driven by differences in feeding strategies and resource availability. Diets for subsampled individuals varied little, suggesting feral pig resource-use strategies in Hawai'i tend to be relatively stable through time. Individual niche width was relatively narrow compared to that of feral pigs in Hawai'i at large, indicating the relatively wide feral pig dietary niche is characterized by substantial intraspecific diet specialization, likely as a result of strong intraspecific competition. Understanding the drivers of feral pig resource use offers key information for management strategies aimed at mitigating their ecological impacts in imperiled systems like Hawai'i.

Resource diversity mitigates the effects of intraspecific competition in co-occurring cryptic nematode species

Intraspecific competition and resource diversity are considered major drivers of niche differentiation, which are expected to promote population niche expansion by driving individuals to feed on alternative resources and/or by enhancing individual diet specialization. Nevertheless, experimental studies on the interaction effects of both factors on animal behavior and population dynamics remain scant. Here, we investigate how resource diversity alters the impact of intraspecific competition on resource preference and fitness of three co-occurring cryptic species of the marine nematode complex Litoditis marina (Pm I, Pm III and Pm IV). For each cryptic species, two competition regimes (‘low nematode density’ and ‘high nematode density’) were established in microcosms with varying resource diversity (E. coli, low-, medium- and high-diversity food). Our results show differences in resource preference and population fitness depending on intraspecific competition and resource diversity, but the response also varied considerably between cryptic species. Pm III did not exhibit resource preference under low intraspecific competition, but preferred the two most diverse food sources under high intraspecific competition. Pm IV also showed preference for medium-diversity food under high competition, whereas no resource preference was observed in Pm I regardless of competition regimes and resource diversity. Nevertheless, all cryptic species exhibited enhanced adult population growth on a more diverse food source under stronger intraspecific competition. These results indicate that resource diversity can alleviate intraspecific competition and affect niche diversification, which may impact diversity maintenance in ecological communities.

Intraspecific competition hinders drought recovery in a resident but not in its range-expanding congener plant independent of mycorrhizal symbiosis

Background and aimsUnderstanding biotic interactions within plant populations and with their symbiotic partners is crucial for elucidating plant responses to drought. While many studies have highlighted the importance of intraspecific plant or mutualistic fungal interactions in predicting drought responses, we know little about the combined effects of these two interactions on the recovery of plants after drought.MethodsWe conducted an experiment to study the recovery after an extreme drought event of a native European plant species (Centaurea jacea) and its range-expanding congener (Centaurea stoebe), across a gradient of plant density and in association with an AM fungal species (Rhizophagus irregularis).ResultsOur results showed strong intraspecific competition in C. jacea, which constrained their post-drought recovery. We further found that AM fungi constrained root biomass recovery of C. jacea after drought under high intraspecific competition. The post-drought recovery in C. stoebe was high potentially due to its greater plasticity in the root diameter under drought conditions.ConclusionStrong intraspecific competition can constrain recovery in plants like C. jacea with lesser root trait plasticity after drought, independent of mycorrhizal symbiosis.

Revisiting the role of behavior-mediated structuring in the survival of populations in hostile environments

Increasing the population density of target species is a major goal of ecosystem and agricultural management. This task is especially challenging in hazardous environments with a high abundance of natural enemies such as parasites and predators. Safe locations with lower mortality have been long considered a beneficial factor in enhancing population survival, being a promising tool in commercial fish farming and restoration of threatened species. Here we challenge this opinion and revisit the role of behavior structuring in a hostile environment in shaping the population density. We build a mathematical model, where individuals are structured according to their defensive tactics against natural enemies. The model predicts that although each safe zone enhances the survival of an individual, for an insufficient number of such zones, the entire population experiences a greater overall mortality. This is a result of the interplay of emergent dynamical behavioral structuring and strong intraspecific competition for safe zones. Non-plastic structuring in individuals’ boldness reduces the mentioned negative effects. We demonstrate emergence of non-plastic behavioral structuring: the evolutionary branching of a monomorphic population into a dimorphic one with bold/shy strains. We apply our modelling approach to explore fish farming of salmonids in an environment infected by trematode parasites.

Gregariousness in lepidopteran larvae.

The gregarious lifestyle of lepidopteran larvae is diverse and shaped by a complex interplay of ecological and evolutionary factors. Our review showed that the larval-aggregation behavior has been reported in 23 lepidopteran families, indicating multiple evolution of this behavior. Some larvae live in sibling groups throughout all larval instars and even pupation stages, which may result from the kin-selection. In contrast, group fusion may occur among different sibling or foraging groups of larvae and form larger aggregates, and the gregariousness of these species might be driven by the group-selection. While group size and foraging patterns vary greatly across species, it is generally associated with improved larval survivorship and accelerated development. However, the advantages of group living, such as facilitating feeding activities, adjusting the temperature, and defending natural enemies, may diminish along with development, with strong intraspecific competition occurring at later instars, even when food is abundant. Therefore, the group sizes and fission-fusion dynamics of certain gregarious lepidopteran larvae may be a consequence of their cost-benefit balance depending on various biotic and abiotic factors. Trail and aggregation pheromones, silk trails, or body contact contribute to collective movement and group cohesion of gregarious lepidopteran larvae. However, frequent contact among group members may cause the horizontal transmission of pathogens and pesticides, which may bring an integrated pest management strategy controlling gregarious lepidopteran pests.

Otolith biochronology for the long-term reconstruction of growth and stock dynamics of fish

Long-term biological time series are essential to evaluate previous responses of organisms to alterations in the environment. Biochronological methods based on archival fish otoliths allow setting such time series, but their predictive potential as proxies of past environmental conditions is still underexploited. In this study, we reconstructed growth variation in European sprat (Sprattus sprattus) in the Baltic Sea from 1956 to 2020 based on measurements of the archived otoliths. We used otolith annual increment widths as a proxy of fish somatic growth. We showed significant negative relationships between sprat growth and sprat spawning stock biomass (SSB) associated with strong intraspecific competition for limited food resources. We also identified a link between sprat growth and water salinity—indicator of the ecosystem's hydrological situation. For the first time, we estimated the SSB prior to the period of available historical data based on the otolith-derived information on the past growth variation. This estimation was based on the strong relationships between SSB and fish growth, complemented with temperature and salinity variables as predictors (R2 = 0.62). A model trained on at least 40 years of data from the more recent past allows us to robustly back-estimate SSB. This study provides new multidecadal data, giving insights into environmental factors affecting the growth of Baltic sprat, and demonstrates the potential of otolith-based biochronology for the provisioning of independent indices of the historical fish stock size. The proposed methodological approach broadens the portfolio of possible applications of the biochronology time series to indicate past changes in the aquatic environment.Graphical abstract

Effects of herbivore on seagrass, epiphyte and sediment carbon sequestration in tropical seagrass bed

Herbivores strongly affect the ecological structure and functioning in seagrass bed ecosystems, but may exhibit density-dependent effects on primary producers and carbon sequestration. This study examined the effects of herbivorous snail (Cerithidea rhizophorarum) density on snail intraspecific competition and diet, dominant seagrass (Thalassia hemprichii) and epiphyte growth metrics, and sediment organic carbon (SOC). The growth rates of the herbivorous snail under low density (421 ind m−2) and mid density (842 ind m−2) were almost two times of those at extremely high density (1684 ind m−2), indicating strong intraspecific competition at high density. Herbivorous snails markedly reduced the epiphyte biomass on seagrass leaves. Additionally, the seagrass contribution to herbivorous snail as food source under high density was about 1.5 times of that under low density, while the epiphyte contribution under low density was 3 times of that under high density. A moderate density of herbivorous snails enhanced leaf length, carbon, nitrogen, total phenol and flavonoid contents of seagrasses, as well as surface SOC content and activities of polyphenol oxidase and β-glucosidase. However, high density of herbivorous snails decreased leaf glucose, fructose, detritus carbon, and total phenols contents of seagrasses, as well as surface SOC content and activities of polyphenol oxidase and β-glucosidase. Therefore, the effects of herbivorous snail on seagrass, epiphyte and SOC were density-dependent, and moderate density of herbivorous snail could be beneficial for seagrasses to increase productivity. This provided theoretical guidance for enhancing carbon sink in seagrass bed and its better conservation.

Coexistence of chaotic and non-chaotic attractors in a three-species slow–fast system

The dynamical complexity of conceptual few-species systems has long been attracting considerable attention. In particular, significant attention has been paid to the three trophic level systems to demonstrate that even a baseline prey/predator/top-predator system can exhibit complex dynamics. However, previous studies left many open questions. In this paper, we consider a generalization of the Hastings–Powell model that include intraspecific competition among the predators and top predators. To enhance the ecological relevance of the model, we incorporate different timescales for different species. Specifically, we aim to study how the dynamics of the system is affected in the presence of multiple timescales along the trophic levels. We employ the geometric singular perturbation approach to analyze the change in species abundance over three different timescales, slow, fast, and intermediate. We analytically determine the entry and exit points from the vicinity of the critical manifold; the corresponding values act as critical thresholds for a sudden, fast transition in population density. We show that the properties of this generalized form of the Hastings–Powell model are dynamically more rich than it was observed in previous studies; particularly, the system can exhibit bi-stability or tri-stability in certain parameter intervals. The existence of a homoclinic orbit in a limiting subsystem (prey–predator) indicates period-doubling cascades to chaos in the full system (prey–predator–top predator). We further investigate the impact of the intraspecific competition of predators and top predators on the system’s dynamics. Strong intraspecific competition among the predators and top predators shows periodic coexistence of all species. In contrast, weak competition can drive the system to exhibit tri-stability, which includes the coexistence of chaotic and periodic (of different periods) attractors.

Cover Picture and Issue Information

Journal of EcologyVolume 110, Issue 10 p. 2265-2267 COVER PICTURE AND ISSUE INFORMATIONFree Access Cover Picture and Issue Information First published: 09 October 2022 https://doi.org/10.1111/1365-2745.13705AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat Graphical Abstract Cover image: A drone photo of the iconic “Red Beach” coastal ecosystem located in Yancheng National Nature Reserve, Jiangsu Province, China. The dominant plant species are Suaeda salsa (red) and Spartina alterniflora (green). The ring-like pattern of Spartina alterniflora may be an indication of strong intra-specific competition that induces mortality of inner-circle individuals. Photograph by Guoyuan Chen. For full details see pages 2268–2287. Volume110, Issue10October 2022Pages 2265-2267 RelatedInformation

Biomass Allocation and Competitive Ability of a Semiarid Perennial Grass and a Legume in Mixtures under Periodical Soil Water Decreasing Conditions

Soil moisture is the key factor controlling plant growth in semiarid grasslands. Here, we sought to evaluate the effects of soil moisture decreasing at different growth stages on biomass accumulation, water use efficiency, and plant-plant interaction of a C3 leguminous subshrub Lespedeza davurica (L) and a C4 perennial grass Bothriochloa ischaemum (B) when sown singly and as a mix in five different ratios in a pot experiment. Results showed that soil water decrease significantly reduced total biomass production of the mixtures by 3.7–53.8% compared with well-watered conditions, and plants at the heading and flowering periods were more vulnerable to soil water decline than those at the late stage. The relative yield total (RYT) of the mixtures was mostly greater than those sown singly. Soil water decreasing increased root/shoot ratio and water use efficiency (WUE) of the mixtures, and such effects were mediated by mixture ratio and/or growth stage. In the mixtures, a strong intraspecific competition was observed in B. ischaemum, whereas interspecific competition in L. davurica. The highest overall biomass (86.47 g pot−1) and WUE (6.33 g kg−1) were observed when the mixture ratio was B:L = 10:2 regardless of soil moisture, and thus could be considered an optimal mixture ratio for establishing restored grassland using the two species. Our results suggest that sown seed mixtures of the two species with an appropriate ratio could sustain a relatively high total biomass production and improve WUE under soil water decreasing conditions in the semiarid Loess Plateau.

Niche overlap in rodents increases with competition but not ecological opportunity: A role of inter-individual difference.

Niche variation at population level mediates niche packing (i.e. patterns of species' spread within the niche space) and species coexistence at community level. Competition and ecological opportunity (resource diversity) are two of the main mechanisms underlying niche variation. Dense niche packing could occur through increased niche partitioning or increased niche overlap. In this study, we used stable carbon and nitrogen isotope data of 635 individual rodents from four species across nine sites in the montane region of a subtropical island to test the effects of competition and ecological opportunity on population isotope niche size, inter-individual niche difference within population and interspecific niche overlap within community. We used the Bayesian Standard Ellipse Area (SEAB, the ellipse area enclosed by carbon and nitrogen isotope values of organisms on a bi-plot) to estimate population niche size and interspecific niche overlap. Inter-individual niche difference within population was quantified as isotopic divergence and isotopic uniqueness. We used rodent abundance (the number of unique individuals captured) to measure competition and plant isotope niche size (plant SEAB) to measure ecological opportunity. The rodents experienced competition as evidenced by a negative relationship between population change rate and conspecific abundance. Rodent population niche size increased with ecological opportunity but not competition. The inter-individual niche difference (isotopic uniqueness) increased with competition (interspecific competition only) but not ecological opportunity. At community level, interspecific niche overlap (herbivore-omnivore pair only) increased with competition (the combined abundance of the pair) but not ecological opportunity. This study demonstrated that isotope niche variation of the rodents could be hierarchically influenced by ecological opportunity and competition, with the former setting the limit of population niche size across communities and the latter shaping inter-individual niche difference and interspecific niche overlap within communities. Under strong intraspecific competition and limited ecological opportunity for niche expansion, individuals may choose to increase their isotopic uniqueness from conspecifics at the cost of overlapping with heterospecifics of different trophic roles within the community niche space as overall competition increases. Denser niche packing of these rodent communities might be achieved through increased niche overlap.

Body size is associated with yearling breeding and extra-pair mating in the Island Scrub-Jay

Abstract Large body size is an important determinant of individual fitness in many animal species, especially in island systems where habitat saturation may result in strong intraspecific competition for mates and breeding territories. Here we show that large body size is associated with benefits to yearling breeding and extra-pair mating in the Island Scrub-Jay (Aphelocoma insularis), endemic to Santa Cruz Island, California. This species is ~20% larger than its mainland congener, consistent with the island syndrome, indicating that body size may be a trait under selection. From 2009 to 2013, we quantified the reproductive success of a marked population of Island Scrub-Jays, tracked which yearlings acquired a breeding territory and bred, and measured the occurrence of extra-pair paternity. Two potential contributors to fitness were positively related to body size. Larger yearling males were more likely to breed, possibly due to greater behavioral dominance during aggressive encounters. Larger males were also less likely to lose paternity to extra-pair males and, anecdotally, extra-pair males were larger than the social male cuckolded. This study provides evidence that larger males may have a fitness advantage over smaller males by breeding earlier and avoiding paternity loss, but estimates of lifetime reproductive success are ultimately needed for Island Scrub-Jays and other long-lived species.

Size‐mediated priority effects are trait‐dependent and consistent across latitudes in a damselfly

Variation in hatching time (phenology) might cause size differences within populations resulting in size‐mediated priority effects (SMPEs) shaping intraspecific interactions. These phenology‐driven effects potentially can be strengthened by seasonal time constraints caused by a short growth season, and depend on latitude. Here the single and combined effects of phenology and latitude‐associated time constraints on SMPEs in larvae of an aquatic insect, the damselfly Lestes sponsa, are studied. We did so by rearing larvae in groups of 16 individuals with different phenology (hatching date) thereby imposing strong intraspecific competition, resulting in cannibalism. We thereby manipulated in a fully crossed way time constraints (combination of temperature and photoperiod: thermo‐photoperiod) in larvae from low‐latitude and more time constrained high‐latitude populations, and examined effects on life history (survival, development, growth) and physiology (fat and protein contents, and phenoloxidase activity as a measure of immune function). Phenology, time constraints and latitude of origin had strong effects on life history, but only the time constraint affected the physiology. We detected a SMPE for survival that, however, was not stronger under time constraints and was consistent in strength between latitudes. Phenology and time constraints interacted for development and growth in a direction suggesting adaptive responses to time constraints but these life history traits did not show SMPEs. We provided important insights in the study of SMPEs thereby showing these to be trait‐dependent and not more pronounced under experimentally manipulated or latitude‐associated time constraints. Our study thereby makes an important addition to geographic variation in SMPEs, a largely neglected topic.

Planting Density Affects Panax notoginseng Growth and Ginsenoside Accumulation by Balancing Primary and Secondary Metabolism.

Adjusting planting density is a common agricultural practice used to achieve maximum yields. However, whether the quality of medicinal herbs can be improved by implementing appropriate planting densities is still uncertain. The medicinal crop Panax notoginseng was used to analyze the effects of planting density on growth and ginsenoside accumulation, and the possible mechanisms of these effects were revealed through metabonomics. The results showed that P. notoginseng achieved high ginsenoside accumulation at high planting densities (8 × 8 and 10 × 10 cm), while simultaneously achieved high biomass and ginsenoside accumulation at moderate planting density of 15 × 15 cm. At the moderate planting density, the primary metabolism (starch and sucrose metabolism) and secondary metabolism (the biosynthesis of phytohormone IAA and ginsenoside) of the plants were significantly enhanced. However, the strong intraspecific competition at the high planting densities resulted in stress as well as the accumulation of phytohormones (SA and JA), antioxidants (gentiobiose, oxalic acid, dehydroascorbic acid) and other stress resistance-related metabolites. Interestingly, the dry biomass and ginsenoside content were significantly lower at low densities (20 × 20 and 30 × 30 cm) with low intraspecific competition, which disturbed normal carbohydrate metabolism by upregulating galactose metabolism. In summary, an appropriate planting density was benefit for the growth and accumulation of ginsenosides in P. notoginseng by balancing primary metabolism and secondary metabolism.

Burrow occupation rates and spatial distribution within habitat of the ghost crab Ocypode quadrata (Fabricius, 1787): Implications for impact assessments

The most conspicuous characteristic of ghost crabs (Crustacea: Ocypodidae) is the construction of semi-permanent burrows on the sand, widely used as tools for impact assessments. Thus, it is now critical to understand the factors driving burrowing activity and burrow occupation rates, including the effects of natural and anthropogenic disturbances. We tested three hypotheses in the present study with the ghost crab Ocypode quadrata: (i) overestimation of population size by traditional burrow counts depends on disturbance level at local scale; (ii) burrow occupation rate is density-dependent; and (iii) the predominant spatial distribution of burrows within the habitat is uniform. The last two predictions were assessed using a meta-analysis for a macro scale assessment. The comparison of distinct methods for burrow counting confirmed that ghost crab density is overestimated on non-urbanized beaches at local scale. A non-linear relationship between burrow occupation rates and burrow density was found at local and regional scales. When burrow density increases, the burrow occupation rate is also higher. However, from approximately 20 to 30 burrows/100 m2, the occupation rate reduces with increased density. Thus, the overestimation of the ghost crab density on non-urbanized beaches can be partially explained by a density-dependence in burrowing activity probably driven by a stronger intraspecific competition on non-urbanized areas. Finally, the distribution pattern of the ghost crab is predominantly uniform in most of its distribution range, despite of urbanization level and sampling design, hence suggesting intra-specific interactions as a driver of burrowing behaviour. In conclusion, burrow density, occupation rate and spatial distribution within the habitat evidenced at various spatial scales that burrowing activity of the ghost crab is density-dependent and this constitutes an issue never considered before in impact assessments inherent to coastal monitoring and management.

Species coexistence in resource‐limited patterned ecosystems is facilitated by the interplay of spatial self‐organisation and intraspecific competition

The exploration of mechanisms that enable species coexistence under competition for a sole limiting resource is widespread across ecology. Two examples of such facilitative processes are intraspecific competition and spatial self‐organisation. These processes determine the outcome of competitive dynamics in many resource‐limited patterned ecosystems, classical examples of which include dryland vegetation patterns, intertidal mussel beds and subalpine ribbon forests. Previous theoretical investigations have explained coexistence within patterned ecosystems by making strong assumptions on the differences between species (e.g. contrasting dispersal behaviours or different functional responses to resource availability). In this paper, I show that the interplay between the detrimental effects of intraspecific competition and the facilitative nature of self‐organisation forms a coexistence mechanism that does not rely on species‐specific assumptions and captures coexistence across a wide range of the environmental stress gradient. I use a theoretical model that captures the interactions of two generic consumer species with an explicitly modelled resource to show that coexistence relies on a balance between species' colonisation abilities and their local competitiveness, provided intraspecific competition is sufficiently strong. Crucially, the requirements on species' self‐limitation for coexistence to occur differ on opposite ends of the resource input spectrum. For low resource levels, coexistence is facilitated by strong intraspecific dynamics of the species superior in its colonisation abilities, but for larger volumes of resource input, strong intraspecific competition of the locally superior species enables coexistence. Results presented in this paper also highlight the importance of hysteresis in understanding tipping points, in particular extinction events. Finally, the theoretical framework provides insights into spatial species distributions within single patches, supporting verbal hypotheses on coexistence of herbaceous and woody species in dryland vegetation patterns and suggesting potential empirical tests in the context of other patterned ecosystems.

The Intraspecific competition as a driver for true production potential of soybean

Phenotypic plasticity of agricultural plants is considered to be one of the main means by which plants cope with the variability of environmental factors. A major contributor to plant plasticity is sowing density, which has a relevant impact on competitive intensity concerning plant density in different environments (CI) and absolute severity of competition (ASC) concerning plant-plant responses to each other in canopy. A field experiment with soybean was set up at the Experimental Station in Prusy, Krakow, to determine the impact of intraspecific competition on growth, plant architecture, nitrogen accumulation, and yield of soybean as an effect of seven different plant densities and weather conditions. The study showed that intraspecific competition in soybean was conditioned by sowing density and access to water, thus revealing the true plant productive potential. Low intraspecific competition increased with plant density causing an increase in the yield of plants. In the wet year of 2014, strong intraspecific competition resulted in high yield and nitrogen accumulation only up to a density of 42 plants m–2, compared to dry years when nitrogen uptake of soybean increased with plant density in full range. The CI and ASC competition indices were sensitive to the varying amount of rainfall. Greater rainfall during crop vegetation increased the intensity of competition as well as the absolute severity of competition and decreased the relative yield with increasing density. In contrast, drought reduced intraspecific competition, eliminating it entirely at over 52 plants m–2.
 Highlights - It is recommended to increase the sowing density in areas with less rainfall to compensate number of pods and number of seeds. - Low intraspecific competition increased yield and nitrogen accumulation in the seeds as the plant density increased.- In the wet year, strong intraspecific competition resulted in high yield and nitrogen accumulation only up to a density of 42 plants m–2. - At very low crop density, the intensity of intraspecific competition is very low and blocked the manifestation of the plants’ true production potential. - In the dry year, nitrogen uptake of soybean increased with plant density in the canopy up to the maximum density.

Interspecific facilitation mediates the outcome of intraspecific interactions across an elevational gradient.

Where interspecific facilitation favors the establishment of high densities of a beneficiary species, strong intraspecific competition may subsequently impede beneficiary performance. Consequently, the negative influence of intraspecific competition between beneficiary individuals could potentially outweigh the positive influence of interspecific facilitation when, for example, higher densities of a beneficiary are negated by the negative effect of crowding on beneficiary reproduction. The aim of this study was, therefore, to examine the impact of an interspecific interaction on the outcome of intraspecific interactions within the context of plant-plant facilitation. We used the cushion-forming Azorella selago and a commonly co-occurring dominant perennial grass species, Agrostis magellanica, on sub-Antarctic Marion Island as a model system. We assessed the impact of an interspecific interaction (between A. selago and A. magellanica) on the outcome of intraspecific interactions (between A. magellanica individuals), by testing if the impact of A. magellanica density on A. magellanica performance is mediated by its interaction with A. selago. We observed evidence for competition among A. magellanica conspecifics, with a decreasing proportion of A. magellanica individuals being reproductive under higher conspecific density. This negative intraspecific effect was greater on A. selago than on the adjacent substrate, suggesting that the facilitative effect of A. selago changes the intensity of intraspecific interactions between A. magellanica individuals. However, experimentally reducing A. magellanica density did not affect the species' performance. We also observed that the effect of A. selago on A. magellanica was positive, and despite the negative effect of intraspecific density on the proportion of reproductive A. magellanica individuals, the net reproductive effort of A. magellanica (i.e., the density of reproductive individuals) was significantly greater on A. selago than on the adjacent substrate. These results highlight that, in abiotically severe environments, the positive effects of interspecific facilitation by a benefactor species may outweigh the negative effects of intraspecific competition among beneficiaries. More broadly, these results suggest that both positive inter- and intraspecific biotic interactions may be key to consider when examining spatial and temporal variation in species' performance.

Why are there so many bee-orchid species? Adaptive radiation by intra-specific competition for mnesic pollinators.

Adaptive radiations occur mostly in response to environmental variation through the evolution of key innovations that allow emerging species to occupy new ecological niches. Such biological innovations may play a major role in niche divergence when emerging species are engaged in reciprocal ecological interactions. To demonstrate coevolution is a difficult task; only a few studies have confirmed coevolution as driver of speciation and diversification. Herein we review current knowledge about bee orchid (Ophrys spp.) reproductive biology. We propose that the adaptive radiation of the Mediterranean orchid genus Ophrys, comprising several hundred species, is due to coevolutionary dynamics between these plants and their pollinators. We suggest that pollination by sexual swindling used by Ophrys orchids is the main driver of this coevolution. Flowers of each Ophrys species mimic a sexually receptive female of one particular insect species, mainly bees. Male bees are first attracted by pseudo-pheromones emitted by Ophrys flowers that are similar to the sexual pheromones of their females. Males then are lured by the flower shape, colour and hairiness, and attempt to copulate with the flower, which glues pollen onto their bodies. Pollen is later transferred to the stigma of another flower of the same Ophrys species during similar copulation attempts. In contrast to rewarding pollination strategies, Ophrys pollinators appear to be parasitized. Here we propose that this apparent parasitism is in fact a coevolutionary relationship between Ophrys and their pollinators. For plants, pollination by sexual swindling could ensure pollination efficiency and specificity, and gene flow among populations. For pollinators, pollination by sexual swindling could allow habitat matching and inbreeding avoidance. Pollinators might use the pseudo-pheromones emitted by Ophrys to locate suitable habitats from a distance within complex landscapes. In small populations, male pollinators would disperse once they have memorized the local diversity of sexual pseudo-pheromone bouquets or if all Ophrys flowers are fertilized and thus repel pollinators via production of repulsive pheromones that mimic those produced by fertilized female bees. We propose the following evolutionary scenario: Ophrys radiation is driven by strong intra-specific competition among Ophrys individuals for the attraction of species-specific pollinators, which is a consequence of the high cognitive abilities of pollinators. Male bees record the pheromone signatures of kin or of previously courted partners to avoid further copulation attempts, thereby inducing strong selection on Ophrys for variation in odour bouquets emitted by individual flowers. The resulting odour bouquets could by chance correspond to pseudo-pheromones of the females of another bee species, and thus attract a new pollinator. If such pollinator shifts occur simultaneously in several indivuals, pollen exchanges might occur and initiate speciation. To reinforce the attraction of the new pollinator and secure prezygotic isolation, the following step is directional selection on flower phenotypes (shape, colour and hairiness) towards a better match with the body of the pollinator's female. Pollinator shift and the resulting prezygotic isolation is adaptive for new Ophrys species because they may benefit from competitor-free space for limited pollinators. We end our review by proritizing several critical research avenues.

Genotypic diversity weakens competition within, but not between, plant species

Abstract Intraspecific diversity can affect the ecological function of communities. Because community function is the outcome of species interactions, understanding how intraspecific diversity affects species interactions may shed light on the mechanism by which intraspecific diversity affects communities. Competition among and within plant species may be particularly important, as the relative strengths of these interactions can influence species coexistence and community stability. We established a field experiment in a New England salt marsh to determine how plant intraspecific diversity affects both intra‐ and interspecific competition. Focal plants of two species (Spartina alterniflora and Spartina patens) were planted with S. alterniflora neighbours that were either genotypically identical to one another (monoculture) or genotypically distinct (polyculture). After two growing seasons, S. alterniflora performance was enhanced in the presence of polyculture conspecific neighbours compared to monoculture neighbours, whereas S. patens was not strongly affected by diversity of its S. alterniflora neighbours. This suggests that intraspecific competition was weakened by intraspecific diversity, potentially even shifting to facilitation, but we did not find strong evidence of intraspecific diversity impacting interspecific competition. By weakening competition within species, intraspecific diversity may promote coexistence of genotypes within that species. Synthesis. The community‐level effects of intraspecific diversity may be mediated in part by its effect on intraspecific competition/facilitation. Furthermore, stronger intraspecific competition and decreased coexistence resulting from reduced diversity may exacerbate ongoing declines in biodiversity in response to global change.