Asymmetric and frequency‐dependent pollinator‐mediated interactions may influence competitive displacement in two vernal pool plants

A plant species immigrating into a community may experience a rarity disadvantage due to competition for the services of pollinators. These negative reproductive interactions have the potential to lead to competitive displacement or exclusion of a species from a site. In this study, we used one‐ and two‐species arrays of potted plants to test for density and frequency dependence in pollinator‐mediated and above‐ground intraspecific and interspecific competition between two species of Limnanthes that have overlapping ranges, but rarely occur in close sympatry. There were asymmetric competitive effects; the species responded differently to their frequency within 16‐plant replacement series arrays. Limnanthes douglasii rosea experienced stronger reductions in lifetime and per‐flower fertility, likely due to pollinator‐mediated competition with Limnanthes alba. This effect may be linked to asymmetrical competition through heterospecific pollen transfer. This study demonstrates that pollinator‐mediated competition may discourage establishment of L. d. rosea in sites already occupied by its congener.     

Bias in the detection of negative density dependence in plant communities

Regression dilution is a statistical inference bias that causes underestimation of the strength of dependency between two variables when the predictors are error‐prone proxies (EPPs). EPPs are widely used in plant community studies focused on negative density‐dependence (NDD) to quantify competitive interactions. Because of the nature of the bias, conspecific NDD is often overestimated in recruitment analyses, and in some cases, can be erroneously detected when absent. In contrast, for survival analyses, EPPs typically cause NDD to be underestimated, but underestimation is more severe for abundant species and for heterospecific effects, thereby generating spurious negative relationships between the strength of NDD and the abundances of con‐ and heterospecifics. This can explain why many studies observed rare species to suffer more severely from conspecific NDD, and heterospecific effects to be disproportionally smaller than conspecific effects. In general, such species‐dependent bias is often related to traits associated with likely mechanisms of NDD, which creates false patterns and complicates the ecological interpretation of the analyses. Classic examples taken from literature and simulations demonstrate that this bias has been pervasive, which calls into question the emerging paradigm that intraspecific competition has been demonstrated by direct field measurements to be generally stronger than interspecific competition.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Evaluating the effects of pollinator‐mediated interactions using pollen transfer networks: evidence of widespread facilitation in south Andean plant communities

Information about the relative importance of competitive or facilitative pollinator‐mediated interactions in a multi‐species context is limited. We studied interspecific pollen transfer (IPT) networks to evaluate quantity and quality effects of pollinator sharing among plant species on three high‐Andean communities at 1600, 1800 and 2000 m a.s.l. To estimate the sign of the effects (positive, neutral or negative), the relation between conspecific and heterospecific pollen deposited on stigmas was analysed with GLMMs. Network analyses showed that communities were characterised by the presence of pollen hub‐donors and receptors. We inferred that facilitative and neutral pollinator‐mediated interactions among plants prevailed over competition. Thus, the benefits from pollinator sharing seem to outweigh the costs (i.e. heterospecific deposition and conspecific pollen loss). The largest proportion of facilitated species was found at the highest elevation community, suggesting that under unfavourable conditions for the pollination service and at lower plant densities facilitation can be more common.     

Signs of stabilisation and stable coexistence

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant–soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant–soil feedback constitute sufficient conditions for stabilisation of two‐species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence‐invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density‐dependence, frequency‐dependence, and plant–soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.     

Pollinator coupling can induce synchronized flowering in different plant species

Synchronous and intermittent plant reproduction has been identified widely in diverse biomes. While synchronous flowering is normally observed within the same species, different species also flower in synchrony. A well-known example of interspecific synchrony is “general flowering" in tropical rain forests of Southeast Asia. Environmental factors, such as low temperature and drought, have been considered as major trigger of general flowering. However, environmental cues are not enough to explain general flowering because some trees do not flower even when they encounter favorable environmental cues. We propose alternative explanation of general flowering; “pollinator coupling”. When species flower synchronously, the elevated pollen and nectar resource may attract increased numbers of generalist pollinators, with a concomitant enhancement of pollination success (facilitation). However, under these circumstances, plants of different species may compete with one another for limited pollinator services, resulting in declines in pollination success for individual species (competition). Here, we present a model describing resource dynamics of individual trees serviced by generalist pollinators. We analyze combinations of conditions under which plants reproduce intermittently with synchronization within species, and/or (sometimes) between different species. We show that plants synchronize flowering when the number of pollinators attracted to an area increases at an accelerating rate with increasing numbers of flowers. In this case, facilitation of flowering by different species exceeds the negative influence of interspecific plant competition. We demonstrate mathematically that co-flowering of different species occurs under a much narrower range of circumstances than intraspecific co-flowering.     

Accounting for interspecific competition and age structure in demographic analyses of density dependence improves predictions of fluctuations in population size

Understanding species coexistence has long been a major goal of ecology. Coexistence theory for two competing species posits that intraspecific density dependence should be stronger than interspecific density dependence. Great tits and blue tits are two bird species that compete for food resources and nesting cavities. On the basis of long‐term monitoring of these two competing species at sites across Europe, combining observational and manipulative approaches, we show that the strength of density regulation is similar for both species, and that individuals have contrasting abilities to compete depending on their age. For great tits, density regulation is driven mainly by intraspecific competition. In contrast, for blue tits, interspecific competition contributes as much as intraspecific competition, consistent with asymmetric competition between the two species. In addition, including age‐specific effects of intra‐ and interspecific competition in density‐dependence models improves predictions of fluctuations in population size by up to three times.     

When can two plant species facilitate each other's pollination?

Facilitation occurs when an increase in the density of one species causes an increase in the population growth rate or the density of a second species. In plants, ample evidence demonstrates that one species can facilitate another by ameliorating abiotic conditions, but the hypothesis that pollination facilitation – in which the presence of one flowering species increases pollinator visits to a second species – can also occur remains controversial. To identify the necessary conditions for pollination facilitation to occur, we constructed population models of two plant species that share the same pollinator and compete for establishment sites, and we assumed that heterospecific pollen can interfere with successful seed set. We found that facilitation for pollination occurs only when the pollinator visitation rate is an initially accelerating function of the combined numbers of flowering plants of both species in a patch. The presence of a second species can allow populations of a focal species either to persist for a longer amount of time before going extinct ("weak facilitation") or to persist indefinitely at a stable equilibrium density ("strong facilitation"). When only a single plant of either species can occupy a site, the plant species with the higher initial density can experience strong facilitation but will eventually out-compete the other species. However, when site occupancy was not exclusive, strong facilitation sometimes led to coexistence of the two species. Increasing the extent of pollen carryover increased the range of initial population densities leading to strong facilitation. In light of our theoretical results, we discuss the apparent rarity of pollination facilitation in nature.     

Individual flowering phenology shapes plant–pollinator interactions across ecological scales affecting plant reproduction

The balance of pollination competition and facilitation among co-flowering plants and abiotic resource availability can modify plant species and individual reproduction. Floral resource succession and spatial heterogeneity modulate plant–pollinator interactions across ecological scales (individual plant, local assemblage, and interaction network of agroecological infrastructure across the farm). Intraspecific variation in flowering phenology can modulate the precise level of spatio-temporal heterogeneity in floral resources, pollen donor density, and pollinator interactions that a plant individual is exposed to, thereby affecting reproduction. We tested how abiotic resources and multi-scale plant–pollinator interactions affected individual plant seed set modulated by intraspecific variation in flowering phenology and spatio-temporal floral heterogeneity arising from agroecological infrastructure. We transplanted two focal insect-pollinated plant species (Cyanus segetum and Centaurea jacea, n = 288) into agroecological infrastructure (10 sown wildflower and six legume–grass strips) across a farm-scale experiment (125 ha). We applied an individual-based phenologically explicit approach to match precisely the flowering period of plant individuals to the concomitant level of spatio-temporal heterogeneity in plant–pollinator interactions, potential pollen donors, floral resources, and abiotic conditions (temperature, water, and nitrogen). Individual plant attractiveness, assemblage floral density, and conspecific pollen donor density (C. jacea) improved seed set. Network linkage density increased focal species seed set and modified the effect of local assemblage richness and abundance on C. segetum. Mutual dependence on pollinators in networks increased C. segetum seed set, while C. jacea seed set was greatest where both specialization on pollinators and mutual dependence was high. Abiotic conditions were of little or no importance to seed set. Intra- and interspecific plant–pollinator interactions respond to spatio-temporal heterogeneity arising from agroecological management affecting wild plant species reproduction. The interplay of pollinator interactions within and between ecological scales affecting seed set implies a co-occurrence of pollinator-mediated facilitative and competitive interactions among plant species and individuals.     

Effects of multiple competitors for pollination on bumblebee foraging patterns and Mimulus ringens reproductive success

When co‐occurring plant species overlap in flowering phenology they may compete for the service of shared pollinators. Competition for pollination may lower plant reproductive success by reducing the number of pollinator probes or by decreasing the quality of pollen transport to or from a focal species. Pair‐wise interactions between plants sharing pollinators have been well documented. However, relatively few studies have examined interactions for pollination among three or more plant species, and little is known about how the outcomes and mechanisms of competition for pollination may vary with competitor species composition. To better understand how the dynamics of competition for pollination may be influenced by changes in the number of competitors, we manipulated the presence of two competitors, Lythrum salicaria and Lobelia siphilitica, and quantified reproductive success for a third species, Mimulus ringens. Patterns of pollinator preference and interspecific transitions in mixed‐species arrays were significantly influenced by the species composition of competitor plants present. Both pair‐wise and three‐species competition treatments led to a similar ～ 40% reduction in Mimulus ringens seed set. However, the patterns of pollinator foraging we observed suggest that the relative importance of different mechanisms of competition for pollination may vary with the identity and number of competitors present. This variation in mechanisms of competition for pollination may be especially important in diverse plant communities where many species interact through shared pollinators.     

天柱山黄山松种内与种间竞争的研究

采用改进的竞争指数模型研究安徽天柱山黄山松的种内和种间竞争强度。结果发现:(1)随对象木胸径的增大,黄山松种群因自然稀疏过程中密度调节作用,植株距离增加,种内竞争强度降低;(2)黄山松群落内其它物种虽然较多,但个体普遍较小,结果种间竞争相对较弱,种内与种间竞争关系顺序为:黄山松—黄山松>杉木—黄山松>枹栎—黄山松>其它树种—黄山松;(3)竞争强度和对象木胸径的关系服从幂函数关系(CI=AD-B),当黄山松胸径达到20cm以上时,竞争强度变化不大,所得的预测模型能很好地预测黄山松种内和种间的竞争强度;(4)改进的竞争指数模型能很好地度量黄山松的种内和种间竞争强度,改进的确定邻体范围的方法能有效地确定黄山松的竞争范围。     

Behavior influences range limits and patterns of coexistence across an elevational gradient in tropical birds

Does competition influence patterns of coexistence between closely related taxa? Here we address this question by analyzing patterns of range overlap between related species of birds (‘sister pairs’) co‐occurring on a tropical elevational gradient. We explicitly contrast the behavioral dimension of interspecific competition (interference competition) with similarity in resource acquisition traits (exploitative competition). Specifically, we ask whether elevational range overlap in 118 sister pairs that live along the Manu Transect in southeastern Peru is predicted by proxies for competition (intraspecific territorial behavior) or niche divergence (beak divergence and divergence times, an estimate of evolutionary age). We find that close relatives that defend year‐round territories tend to live in non‐overlapping elevational distributions, while close relatives that do not defend territories tend to broadly overlap in elevational distribution. In contrast, neither beak divergence nor evolutionary age was associated with patterns of range limitation. We interpret these findings as evidence that behavioral interactions – particularly direct territorial aggression – can be important in setting elevational range limits and preventing coexistence of closely related species, though this depends upon the extent to which intraspecific territorial behavior can be extended to territorial interactions between species. Our results suggest that interference competition can be an important driver of species range limits in diverse assemblages, and thus highlight the importance of considering behavioral dimensions of the niche in macroecological studies.     

Pollination of Ipomopsis aggregata (Polemoniaceae): effects of intra- vs. interspecific competition

Although plants may simultaneously experience intra- and interspecific competition for pollination, their relative strength has rarely been experimentally evaluated. Yet because intra- and interspecific competition can be caused by different mechanisms, their effect on the ecology and evolution of plants may differ. To determine the relative strength of intra- and interspecific competition for pollination, I manipulated the presence of heterospecifics and density of conspecifics using Ipomopsis aggregata as the focal species. All plots contained I. aggregata and Castilleja linariaefolia, but C. linariaefolia inflorescences were removed from half of the plots to create the heterospecifics-absent treatment. Within each plot, all I. aggregata inflorescences were removed from a 5-m radius around a focal plant to create a low conspecific density experimental unit, and a group of 12 I. aggregata plants/1 m² was designated as a high conspecific density unit. Conspecific pollen deposition was reduced when C. linariaefolia was present but was not influenced by I. aggregata density. Although seed set per fruit was reduced by 17% when C. linariaefolia was present, it was not significantly influenced by either treatment. Interspecific competition for pollination is stronger than intraspecific competition in the I. aggregata–C. linariaefolia system, but neither process appears to influence plant fitness.     

Intrinsic and extrinsic factors acting on the reproductive process in alpine‐snowbed plants: roles of phenology,biological interaction,and breeding system

Pollinator activity and competition for pollinators lead to quantitative and qualitative pollen limitations on seed production and affect the reproductive success of plant species, depending on their breeding system (e.g., self‐compatibility and heterospecific compatibility) and genetic load (e.g., inbreeding depression and hybrid inviability). In alpine ecosystems, snowmelt regimes determine the distribution and phenology of plant communities. Plant species growing widely along a snowmelt gradient often grow with different species among local populations. Their pollinators also vary in their abundance, activity, and behavior during the season. These variations may modify plant–pollinator and plant–plant interactions. We integrated a series of our studies on the alpine dwarf shrub, Phyllodoce aleutica (Ericaceae), to elucidate the full set of intrinsic (species‐specific breeding system) and extrinsic factors (snow condition, pollinator activity, and interspecific competition) acting on their reproductive process. Seasonality of pollinator activity led to quantitative pollen limitation in the early‐blooming populations, whereas in the late‐blooming populations, high pollinator activity ensured pollination service, but interspecific competition for pollinators led to qualitative and quantitative pollen limitation in less competitive species. However, negative effects of illegitimate pollen receipt on seed‐set success might be reduced when cryptic incompatibility systems (i.e., outcross pollen grains took priority over self‐ and heterospecific pollen grains) could effectively prevent ovule and seed discounting. Our studies highlight the importance of species‐specific responses of plant reproduction to changing pollinator availability along environmental gradients to understand the general features of pollination networks in alpine ecosystems.     

Floral density and co-occurring congeners alter patterns of selection in annual plant communities*

Although the evolution and diversification of flowers is often attributed to pollinator-mediated selection, interactions between co-occurring plant species can alter patterns of selection mediated by pollinators and other agents. The extent to which both floral density and congeneric species richness affect patterns of net and pollinator-mediated selection on multiple co-occurring species in a community is unknown and is likely to depend on whether co-occurring plants experience competition or facilitation for reproduction. We conducted an observational study of selection on four species of Clarkia (Onagraceae) and tested for pollinator-mediated selection on two Clarkia species in communities differing in congeneric species richness and local floral density. When selection varied with community context, selection was generally stronger in communities with fewer species, where local conspecific floral density was higher, and where local heterospecific floral density was lower. These patterns suggest that intraspecific competition at high densities and interspecific competition at low densities may affect the evolution of floral traits. However, selection on floral traits was not pollinator mediated in Clarkia cylindrica or Clarkia xantiana, despite variation in pollinator visitation and the extent of pollen limitation across communities for C. cylindrica. As such, interactions between co-occurring species may alter patterns of selection mediated by abiotic agents of selection.     

The interspecific competition presents greater nutrient facilitation compared with intraspecific competition through AM fungi interacting with litter for two host plants in karst soil

AM 真菌和枯落物互作下两种喀斯特植物种间竞争较种内竞争更能促进植物养分利用 枯落物是植物养分获取和土壤养分转化的关键载体。丛枝菌根(Arbuscular mycorrhizae, AM)对植物养分摄取的影响已被广泛认知。然而，在养分亏缺的喀斯特生境中，不同竞争方式的植物如何通过AM真菌和枯落物利用养分尚不清楚。本研究对两种喀斯特适生植物构树(Broussonetia papyrifera)和云贵鹅耳枥(Carpinus pubescens)进行种内竞争和种  间竞争种植处理，并通过幼套球 囊霉(Glomus etunicatum)接种或不接种处理，以及土壤中添加或不添加两物种叶片混合枯落物处理，测定了植物生物量以及氮、磷、钾浓度等指标，研究植物的生长和养分利用。研究结果表明，AM真菌对两种植物养分摄取影响不同，AM真菌显著提高了种内和种间竞争下构树的养分摄取量，但降低了云贵鹅耳枥的养分摄取量。种间竞争下接种AM真菌，枯落物添加促进了云贵鹅耳枥对氮的摄取，抑制了构树对氮的摄取。接种AM真菌和添加枯落物条件下，种间竞争的构树对氮、磷和钾的摄取量及云贵鹅耳枥对氮的摄取量均高于种内竞争；种间竞争下两物种养分竞争力呈现明显差异，即构树对磷和钾养分竞争力显著提高，对氮则不显著；云贵鹅耳枥仅对钾的养分竞争力显著降低，对氮和磷则无显著影响。这些结果说明，在AM真菌与枯落物相互作用下，两种喀斯特植物种间竞争较种内竞争更能促进植物养分利用。     

Relative importance of competition and plant–soil feedback,their synergy,context dependency and implications for coexistence

Plants interact simultaneously with each other and with soil biota, yet the relative importance of competition vs. plant–soil feedback (PSF) on plant performance is poorly understood. Using a meta‐analysis of 38 published studies and 150 plant species, we show that effects of interspecific competition (either growing plants with a competitor or singly, or comparing inter‐ vs. intraspecific competition) and PSF (comparing home vs. away soil, live vs. sterile soil, or control vs. fungicide‐treated soil) depended on treatments but were predominantly negative, broadly comparable in magnitude, and additive or synergistic. Stronger competitors experienced more negative PSF than weaker competitors when controlling for density (inter‐ to intraspecific competition), suggesting that PSF could prevent competitive dominance and promote coexistence. When competition was measured against plants growing singly, the strength of competition overwhelmed PSF, indicating that the relative importance of PSF may depend not only on neighbour identity but also density. We evaluate how competition and PSFs might interact across resource gradients; PSF will likely strengthen competitive interactions in high resource environments and enhance facilitative interactions in low‐resource environments. Finally, we provide a framework for filling key knowledge gaps and advancing our understanding of how these biotic interactions influence community structure.     

Coexistence induced by pollen limitation in flowering-plant species

We report a novel mechanism for species coexistence that does not invoke a trade-off relationship in the case of outbreeding flowering plants. Competition for pollination services may lead to interspecific segregation of the timing of flowering among plants. This, in turn, sets limits on the pollination services, which restrain the population growth of a competitively superior species, thereby allowing an inferior species to sustain its population in the habitat. This explains the often-observed tendency for interspecific differentiation in the timing of flowering between coexisting plants. It also predicts that the introduction of an efficient pollinator to a habitat may cause the extinction of competitively inferior plant species.     

Active dispersal is differentially affected by inter‐ and intraspecific competition in closely related nematode species

Competition is one of the main drivers of dispersal, which can be an important mechanism to achieve permanent or temporal coexistence of multiple species. This coexistence can be achieved by a dispersal‐competition tradeoff, spatial store effects or neutral dynamics. Here we test the effect of inter‐ and intraspecific competition on dispersal of four species of the marine nematode species complex Litoditis marina. A previous study in closed microcosms without a possibility for dispersal had demonstrated pronounced interspecific competition, leading to the exclusion of one species. We now investigated whether 1) the dispersal is affected by interspecific interactions, by intraspecific competition (density) or by food availability, 2) the dispersal dynamics influence assemblage composition and can lead to co‐occurrence of the species, and 3) the abiotic environment (here salinity) can affect these dynamics. We show that density is the main driver for dispersal in two of the four species. Dispersal of a third species always started at the same time irrespective of density, whereas in the fourth species interspecific interactions accelerated dispersal. Remarkably, this fourth species was not a strong competitor, suggesting that a dispersal–competition tradeoff does not explain the observed coexistence. Salinity did not alter the timing of dispersal when interspecific interactions were present but did affect assemblage composition. Consequently, spatial store effects may influence coexistence. All four species co‐occurred in fairly stable abundances throughout the present experiment indicating the importance of species specific dispersal strategies for coexistence. Co‐occurrence can be facilitated because competition is postponed or avoided by dispersal. Neutral dynamics also played a role as intra‐ and interspecific competition were of similar importance in three of the four species. We conclude that dispersal is a driver of the coexistence of closely related nematode species, and that population density and interspecific interactions shape these dynamics.     

Temporal variations in the linkages between plants and flower visitors and the pollination success of Primula modesta along the snowmelt gradient

Although pollination networks between plants and flower visitors are diverse and flexible, seed production of many plant species is restricted by pollen limitation. Obligate outcrossers often suffer from low pollinator activity or severe interspecific competition for pollinator acquisition among co-flowering species. This study focused on seasonal changes in plant–flower visitor linkages in an alpine ecosystem and examined whether and how this seasonality affected the seed-set of Primula modesta, a self-incompatible distylous herb having long-tubed flowers. First, we recorded the linkages between plants and flower visitors along the snowmelt gradient. Then, pollination experiment was conducted to estimate the degree of pollen limitation over the course of flowering season of P. modesta. Flower visitors were classified by their tongue length based on the morphological matching with P. modesta flowers. As the season progressed, plant–visitor linkages became more diverse and generalized, and the visitation frequency to P. modesta flowers increased. In the later part of the season, however, the seed set of P. modesta was significantly reduced due to severe pollen limitation, presumably because of increased competition for long-tongued pollinators among co-flowering species. The present study revealed that pollinator availability for specialist species may be restricted even when plant–visitor linkages are diverse and generalized as a whole. In the case of P. modesta, morphological matching and competition for pollinators might be the main factors explaining this discrepancy.