The role of biotic interactions in altering tree seedling responses to an extreme climatic event

Questions: We addressed two poorly understood aspects of plant response to climate change: the impact of extreme climatic events and the mediating role of biotic interactions, through a study of heatwave effects on tree seedling survival rates and ability of the tree canopy to alter seedling responses. Location: Mountain belt of the northern French Alps (Maurienne Valley). Methods: The survival rates of two seedling cohorts from four tree species (Abies alba, Acer pseudoplatanus, Fraxinus excelsior and Picea abies) were measured during both the 2003 European heatwave and an average summer (2004) in deciduous broadleaf mountain forests. Seedlings were transplanted into two soil moisture conditions, and in experimental gaps or under the tree canopy. Results: The heatwave strongly decreased tree seedling survival rates, while there was an important species‐specific mediating role of biotic interactions. In the wettest conditions, the tree canopy strongly increased survival of Abies, buffering the negative impact of the heatwave. In contrast, in the driest conditions, the tree canopy decreased survival of Picea and Acer, amplifying the negative impact of the heatwave. We found evidence of increasing soil water stress in the understorey of the driest community, but further studies including vapour pressure deficit measurements are needed to elucidate the driving mechanism of facilitation. Conclusions: The high species specificity of the mediating role of biotic interactions and its variation along stress gradients leads to questions on our ability to predict large‐scale responses of species to climate changes.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Positive interactions in plant communities and the individualistic-continuum concept

The individualistic nature of communities is held as a fundamental ecological tenet by many ecologists. The empirical rationale for the individualistic hypothesis is largely based on gradient analyses in which plant species are almost always found to be arranged independently of one another in “continua” along environmental gradients. However, continua are correlative patterns and do not identify the processes that determine them, and so they do not necessarily preclude the possibility of interdependent interactions within plant communities. For example, the common occurrence of positive interactions suggests that plant species may not always be distributed independently of each other. If the distributions and abundances of species are enhanced by the presence of other species, their organization is not merely a coincidence of similar adaptation to the abiotic environment. Interpretations of gradient analyses also appear to assume that interactions among species should be similar at all points along environmental axes, and that groups of species should be associated at all points on a gradient if interdependence is to be accepted. However, virtually all types of ecological interactions have been shown to vary with changes in the abiotic environment, and a number of field experiments indicate that positive effects become stronger as abiotic stress increases. Furthermore, interactions among plants have been shown to shift from competition to facilitation along environmental continua. Thus, significant interdependence may occur even when species do not fully overlap in distribution. Higher-order, indirect interactions between animals and plants, and among plants, also suggest that interdependence within communities occurs. Eliminating a species involved in an indirect interaction may not necessarily mean that its beneficiary will be eliminated from a community, but the prospect that the distribution and abundance of any species in a plant community may be positively affected by the effects that other species have on their competitors suggests that communities are organized by much more than “the fluctuating and fortuitous immigration of plants and an equally fluctuating and variable environment” as stated by Henry Gleason. The ubiquity of direct and indirect positive interactions within plant communities provides a strong argument that communities are more interdependent than current theories allow. Received: 17 February 1997 / Accepted: 23 May 1997     

The role of mycorrhiza in plant populations and communities

Summary There is a large body of literature concerning the value of mycorrhiza to plant growth. Recent emphasis on the potential benefits of the fungi to natural ecosystems conservation and productive agriculture has focused attention on the roles and underlying mechanisms of the association. In parallel, recognition that isolates/ species of vesicular arbuscular mycorrhizal fungi have variable life-history traits has resulted in investigations focused on the symbiotic fungi rather than the host, a perspective which may reflect the fungal evolutionary strategies. This paper discusses progress in understanding interactions amongst hyphae, which in one form or another are a major component of mycorrhiza and each phase of the life-history.     

Seed dispersal spectra of plant communities in the Iberian Peninsula

We studied the seed dispersal spectra of 46 Iberian plant communities of five types: potential woodland, forest fringe, substitutional scrubland, nitrophile communities and montane communities. There were no significant differences between the spectra of communities of the same type from Mediterranean and Eurosiberian regions. Biotic dispersal is most common at mature stages of succession, and abiotic dispersal at immature stages.     

Determinants of coastal treeline and the role of abiotic and biotic interactions

Recent die-off of coastal forests has been attributed primarily to the effects of sea level rise by correlation with tide-gage records. Due to the temporal and spatial scales involved, direct monitoring of sea level rise impacts is challenging and its attribution can be confounded by both land-use history and species interactions. Here we present experimental evidence for a micro-tidal, oligohaline estuarine system that the location of coastal treeline is determined by both environmental controls and positive and negative species interactions. We conducted field surveys and a transplant experiment to determine the controls on pine seedling establishment and survival along a salinity and flooding gradient. Using a two-way changepoint model we determined that sawgrass cover (Cladium jamaicense) and salinity concentrations interact to control natural pine regeneration (Pinus taeda). The long-term removal of sawgrass resulted in increased soil salinity and high rates of (planted) pine seedling mortality. In contrast, pine seedlings planted directly under sawgrass were able to survive at the same level as upland forest plots because of reduced salinity levels. This research provides evidence that sawgrass can facilitate pine seedling survival, but also suggests that either competitive exclusion by sawgrass or dispersal limitations prevent initial pine seedling regeneration. We propose that forest dynamics are closely coupled to fire, which consumes sawgrass biomass and reduces competition between pine seedlings and grass. Following fire, pine seedling establishment and the regrowth of sawgrass facilitates long-term pine seedling survival. Under this scenario, recent marsh invasion into coastal forests may not necessarily represent a permanent state change in locations where abiotic stress is not the only determinant for community composition.     

Plant traits: Their role in the regeneration of alpine plant communities in sub‐arctic Finland

Abstract. Question: How do the relative frequencies of plant traits (clonality, growth form, seed weight, diaspore morphology) vary during the life cycle and how does this affect regeneration? Location: Alpine meadow and heath communities at Kilpisjärvi, sub‐Arctic Finland. Methods: Control plots and three treatments were used to measure relative species abundances for five life cycle stages: standing vegetation, seed rain, seed bank and seedlings emerging in gaps and in closed vegetation. Results: The relative frequencies of plant traits varied between the life cycle stages. The meadows were dominated by weakly clonal herbs, small or intermediate seeds and unappendaged diaspores, while the heaths were dominated by clonal dwarf shrubs, small seeds and fleshy fruits. In the meadows, species with small seeds dominated during the seed rain and in the seedling stage in gaps, while species with intermediate seeds dominated the seed bank and the seedling stage in closed vegetation. Species with unappendaged diaspores dominated throughout the life cycle. In the heaths, seed bank and seedling stage were practically absent. Conclusions: The observed differences in plant trait spectra between life cycle stages indicate that important environmental factors differ among the stages. Small seeds are advantageous for dispersal, whereas intermediate seeds have a greater probability of germinating and establishing in closed vegetation. Appendages facilitate dispersal, whereas unappendaged diaspores favour seed burial. Although the plant growth form spectrum largely reflects environmental constraints during the regeneration cycle, information on seed weight and diaspore morphology improves our knowledge of the relative importance of morphological adaptations of sexual structures in different stages during the life cycle.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

Convergence patterns and multiple species interactions in a designed plant mixture of five species

It is known that convergence and divergence can occur in complex plant communities, but the relative importance of biotic and abiotic factors driving these processes is less clear. We addressed this issue in an experiment using a range of mixed stands of five species that are common in Swiss fens (Carex elata, C. flava, Lycopus europaeus, Lysimachia vulgaris and Mentha aquatica) and two levels of water and nutrients. One hundred and seventy-six experimental mixtures were maintained in large pots (75 l) for two consecutive growing seasons in an experimental garden. The stands varied systematically in the initial relative abundance of each of the five species and in overall initial stand abundance. The changes in biomass over 2 years were modelled as linear functions of treatments and the initial biomass of each species. The dynamics of the system were mainly driven by differences in the identity of species and by a negative feedback mechanism but also by different abiotic conditions. In all mixtures, C. elata became more dominant over time, which caused an overall convergence of community composition. In addition, the rate of change of each species’ biomass was negatively related to its own initial abundance. Thus, a negative feedback further contributed to the convergence of communities. Species responded differently to water level and nutrient supply, causing community dynamics to differ among treatments. However, the different abiotic conditions only slightly modified the overall convergence pattern. Competitive interactions between more than two species were weaker than the negative feedback but still significantly influenced the species’ final relative abundance. The negative feedback suggests that there is niche partitioning between the species, which permits their coexistence. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Resource limitation,habitat segregation,and species interactions of british tree-hole mosquitoes in nature

Summary The insect fauna of water-filled tree holes in southern Britain consists primarily of the mosquitoes Aedes geniculatus, Anopheles plumbeus, Culex torrentium, and a benthic detritivorous fauna that includes primarily the scirtid beetle Prionocyphon serricornis and the chironomid midge Metriocnemus martinii. Culex torrentium has been documented only relatively recently in tree holes but all three species of mosquitoes partition the resource in space and time. When mosquito larvae were forced to coexist in natural tree holes at limiting densities and at higher than natural levels of interspecific encounter, there was no evidence that Aedes geniculatus or Anopheles plumbeus affected pupation success, pupal weight, or development time of the other or that either Aedes geniculatus or C. torrentium affected the survivorship, pupation success, pupal weight, and biomass yield of the other. When A. geniculatus at limiting densities were forced in natural tree holes to live without or to coexist with natural or twice natural densities of P. serricornis and M. martinii, the presence, absence, or superabundance of the benthic insects did not affect pupation success or pupal weight of A. geniculatus; development time of A. geniculatus was faster when a superabundance of the benthic fauna was present. Effects of the benthic fauna on A. geniculatus are slight and the only significant interaction is facilitative, not competitive. The pattern of habitat segregation among treehole mosquitoes in southern Britain is characteristic of their respective genera and we propose that this pattern is more likely (but not certain) to have arisen through a process of independent evolution than through competitively driven niche shifts among already coexisting species.     

The relative importance of positive and negative interactions for pollinator attraction in a plant community

Plant–pollinator interactions provide ideal frameworks for studying interactions in plant communities. Despite the large potential influence of such interactions on plant community structure, biodiversity and evolutionary processes, we know surprisingly little about the relative importance of positive and negative interactions among plant species for pollinator attraction. Therefore, we explored the relationships between conspecific and heterospecific floral densities and the flower visitation rates of nine plant species mainly visited by bumble bees, and six plant species mainly visited by flies, in a temperate grassland, through stepwise multiple regressions. Significant relationships were interpreted as interactions for pollinator attraction. Our results revealed that positive intra- and interspecific interactions for pollinator attraction were far more frequent than negative ones. Seventeen interspecific interactions were revealed of which 14 were significantly positive, whereas three of four significant intraspecific interactions were positive. Seven species experienced only positive interactions and two species experienced only negative interactions. The results presented here indicate that negative interactions are not necessarily the dominant ecological interaction for pollination among plants within a community, and the study represents a straightforward approach to study intra- and interspecific interactions among multiple species within a community. We discuss which mechanisms may drive the positive interactions for pollinator attraction and whether this may result in facilitative effects on reproductive success. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The roles of stochasticity and biotic interactions in the spatial patterning of plant species in alpine communities

Questions

Plant community composition can be influenced by multiple biotic, abiotic, and stochastic factors acting on the local species pool to determine their establishment success and abundance and subsequently the diversity of the community. We asked if the influences of biotic interactions on the composition of plant species in communities, as indicated by patterns of plant species spatial associations (independent, positive or negative), vary across a productivity gradient within a single ecosystem type. Do dominant species of communities show spatial patterning suggestive of competitive interactions with interspecific neighbors? Do species that span multiple community types exhibit the same heterospecific interactions with neighbours in each community?

Location

Three alpine communities in the southern Rocky Mountains.

Methods

We measured the occurrence of species in a 1‐cm spatial grid within 2 m × 2 m plots to determine the spatial patterns of species pairs in the three communities. A null model of independent species spatial arrangements was used to determine whether species pairs were positively, negatively or independently associated, and how these patterns differed among the communities across the gradient of resource supply and environmental stress.

Results

Positive associations, indicative of facilitation between species, were most common in the most resource‐poor and least productive community. However negative associations, suggestive of competitive interactions among species, were not more common in the two more resource‐rich, productive communities. The dominant species of these communities did exhibit higher negative than positive associations with neighbours relative to positive patterning. Independent interspecific patterning was equally common relative to positive and negative patterns in all communities. Species that previously were shown to either facilitate other species or compete with neighbours exhibited spatial patterning consistent with the earlier experimental work.

Conclusions

A large number of species exhibit a lack of net biotic interactions, and stochastic factors appear to be as important as competition and facilitation in shaping the structure of the three alpine plant communities we studied.

     

The role of birds and mammals in Korean pine (Pinus koraiensis) regeneration dynamics

This study examined the interrelationships of the fall seed-foraging guild with Korean pine (Pinus koraiensis) and its regeneration. The study took place in old-growth and secondary-growth forests in Northeastern China. Foraging behavior, seed transportation distance, and cache location of various birds and mammals were observed. Regeneration characteristics of Korean pine were also analyzed using plots in various vegetation cover types, successional stages, and topographical situations. Ten species of vertebrates were able to harvest seed from the closed-cone Korean pine. Of these, the Eurasian nutcracker, Eurasian nuthatch, red squirrel, and Siberian chipmunk were found to be potential seed dispersal agents. The nutcracker was the most important dispersal agent, easily acquiring seed with its large pointed bill, carrying up to 62 seeds in one trip, carrying seed at least 4 km, and placing seed in a variety of sites 2.5–3 cm deep in the soil. The Eurasian nuthatch carried a single seed per trip at distances less than 50 m, needed nutcrackers to open the cones and expose seed before they could acquire seed, and occasionally cached seed in the soil. Red squirrels were uncommon visitors to the tree tops of Korean pine, carried cones shorter distances than nutcrackers, and were only found under forest canopies. Human harvest of cones by knocking off branches also affected squirrel behavior and reduced future cone crops. Siberian chipmunks also collected seed from cones in trees and appeared to transport seed less than 50 m. Six other species were observed in this study successfully harvesting seed from cones but were not potential seed dispersers. Natural seedling establishment was found to be over 1000 seedlings/ha except on old-growth pine-hardwood sites. Squirrels were commonest here, but few seedlings survived past the 2nd year due to the intense shading. Second-growth forest types, including aPicea plantation where nutcrackers cached seed daily, and an old-growth pine-hardwood selective-harvest site, had the greatest regeneration. In conclusion, most natural regeneration of Korean pine in this part of its range is due primarily to the Eurasian nutcracker. Nutcrackers can aid forest managers in reaching desired stocking levels after disturbance, as well as a more natural-appearing forest. Squirrels, chipmunks, and nuthatches are minor seedling establishment agents. Korean pine seed is an important food source used by at least 22 species of forest wildlife.     

Extreme climatic events change the dynamics and invasibility of semi-arid annual plant communities

Extreme climatic events represent disturbances that change the availability of resources. We studied their effects on annual plant assemblages in a semi-arid ecosystem in north-central Chile. We analysed 130 years of precipitation data using generalised extreme-value distribution to determine extreme events, and multivariate techniques to analyse 20 years of plant cover data of 34 native and 11 exotic species. Extreme drought resets the dynamics of the system and renders it susceptible to invasion. On the other hand, by favouring native annuals, moderately wet events change species composition and allow the community to be resilient to extreme drought. The probability of extreme drought has doubled over the last 50 years. Therefore, investigations on the interaction of climate change and biological invasions are relevant to determine the potential for future effects on the dynamics of semi-arid annual plant communities.     

A theory of the spatial and temporal dynamics of plant communities

An individual-based model of plant competition for light that uses a definition of plant functional types based on adaptations for the simultaneous use of water and light can reproduce the fundamental spatial and temporal patterns of plant communities. This model shows that succession and zonation result from the same basic processes. Succession is interpreted as a temporal shift in species dominance, primarily in response to autogenic changes in light availability. Zonation is interpreted as a spatial shift in species dominance, primarily in response to the effect of allogenic changes in water availability on the dynamics of competition for light. Patterns of succession at different points along a moisture gradient can be used to examine changes in the ecological roles of various functional types, as well as to address questions of shifts in patterns of resource use through time.Our model is based on the cost-benefit concept that plant adaptations for the simultaneous use of two or more resources are limited by physiological and life history constraints. Three general sets of adaptive constraints produce inverse correlations in the ability of plants to efficiently use (1) light at both high and low availability, (2) water at both high and low availability, and (3) both water and light at low availabilities.The results of this type of individual-based model can be aggregated to examine phenomena at several levels of system organization (i.e., subdisciplines of ecology), including (1) plant growth responses over a range of environmental conditions, (2) population dynamics and size structure, (3) experimental and field observations on the distribution of species across environmental gradients, (4) studies of successional pattern, (5) plant physiognomy and community structure across environmental gradients, and (6) nutrient cycling.     

The role of Namibian inselbergs in contributing to local and regional plant species richness

This study investigated four inselberg landscapes in Namibia's aridNama Karoo. Inselbergs of different size and geology were investigated (a) todetermine environmental variables influencing the number of inselberg restrictedplant species and (b) to investigate seed dispersal spectra as one parameterdescribing functional properties of inselberg specialists. With regard to therole of inselbergs in contributing to local and regional species richness, manyspecies restricted to inselberg habitats occurred (up to 79 species recorded inthis survey) and so contribute to local and regional species richness. As nospecies endemic to an inselberg were recorded in this study, inselbergspecialists may provide an indicator for conservation value of these mountains.Elevation rather than surface area provided a measure for predicting the numberof inselberg specialists. Besides elevation, underlying geology, and resultinglandforms and edaphic conditions, as well as distance to potential mainland alsoinfluenced the number of inselberg specialists. Functional relationships betweeninselbergs based on seed dispersal spectra of inselberg specialists were notconclusive.     

The role of seed traits in determining the phylogenetic structure of temperate plant communities

Background and Aims

Phylogenetic clustering of species within plant communities can be expected to result from environmental filtering acting on an evolutionary-conserved plant trait. One such a candidate trait is the embryo to seed-size ratio (E:S). A high E:S may allow faster germination immediately after imbibition, and is therefore assumed to be advantageous in dry habitats. In this study the hypothesis was tested that habitat filtering driven by soil moisture conditions and acting on seed germination and seedling establishment is an important ecological mechanism in structuring temperate plant communities.

Methods

Vegetation samplings were performed in three habitats located within 200 km of each other in western Europe: Ellenberg indicator values showed that the habitats selected differed substantially in terms of soil moisture and light availability. E.S ratio and seed mass data for all genera were obtained from literature. Data were analysed using recently developed phylogenetic methods.

Key Results

Genera with a similar E:S tend to co-occur, as low and high E:S genera dominate in moist and dry habitats, respectively. A phylogenetically clustered pattern of community structure was evident, and dispersion of E:S was positively related to phylogenetic dispersion.

Conclusions

The phenotypically and phylogenetically clustered pattern indicates that E:S-mediated habitat filtering is an important assembly process structuring the plant community of the temperate climate habitats studied.