Dynamics and spatial organization of plant communities in water-limited systems

A mathematical model for plant communities in water-limited systems is introduced and applied to a mixed woody-herbaceous community. Two feedbacks between biomass and water are found to be of crucial importance for understanding woody-herbaceous interactions: water uptake by plants' roots and increased water infiltration at vegetation patches. The former acts to increase interspecific competition while the latter favors facilitation. The net interspecific interaction is determined by the relative strength of the two feedbacks. The model is used to highlight new mechanisms of plant-interaction change by studying factors that tilt the balance between the two feedbacks. Factors addressed in this study include environmental stresses and patch dynamics of the woody species. The model is further used to study mechanisms of species-diversity change by taking into consideration tradeoffs in species traits and conditions giving rise to irregular patch patterns.     

Competition, traits and resource depletion in plant communities

Although of primary importance to explain plant community structure, general relationships between plant traits, resource depletion and competitive outcomes remain to be quantified across species. Here, we used a comparative approach to test whether instantaneous measurements of plant traits can capture both the amount of resources depleted under plant cover over time (competitive effect) and the way competitors perceived this resource depletion (competitive response). We performed a large competition experiment in which phytometers from a single grass species were transplanted within 18 different monocultures grown in a common-garden experiment, with a time-integrative quantification of light and water depletion over the phytometers’ growing season. Resource-capturing traits were measured on both phytometers (competitive response traits) and monocultures (competitive effect traits). The total amounts of depleted light and water availabilities over the season strongly differed among monocultures; they were best estimated by instantaneous measurements of height and rooting depth, respectively, performed when either light or water became limiting. Specific leaf area and leaf water potential, two competitive response traits measured at the leaf level, were good predictors of changes in phytometer performance under competition, and reflected the amount of light and water, respectively, perceived by plants throughout their lifespan. Our results demonstrated the relevance of instantaneous measures of plant traits as indicators of resource depletion over time, validating the trait-based approach for competition ecology. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Individuals have limitations,not communities — A response to Marrs,Weiher and Lortie et al.

A reply is presented on the comments by Marrs, Weiher, and particularly Lortie et al. on an earlier Forum paper. The main point is that adapted alpine plants are not stressed, which follows, i.a., from their productivity which is equal to that in tropical systems when the length of the growing season is taken into account. Another point is that individual‐based and community‐based considerations should not be confused.     

Individual-based model of spatio-temporal dynamics of mixed forest stands

This research presents the results of constructing and parameterizing an individual-based model of spatiotemporal dynamics of mixed forest stands. The model facilitates computerized experiments with forest stands having different combinations of species and age structures. These forest stands grow on temperate areas where light is the main system-forming factor that shapes and develops forest ecosystems. The model TEMFORM (TEMperate FORests Model) is developed with few equations and parameters, most of which can be estimated using standard forest inventory data. Parameterization of the model used the growth tables of a set of basic forest-forming species in Far East Russia. Simulation results of the development of the natural single- and mixed-species stands and the effects of different types of disturbances on the stand dynamics and compositions are presented.     

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.     

Shedding light on plant competition: modelling the influence of plant morphology on light capture (and vice versa)

A plant's morphology is both strongly influenced by local light availability and, simultaneously, strongly influences this local light availability. This reciprocal relationship is complex, but lies at the heart of understanding plant growth and competition. Here, we develop a sub-individual-based simulation model, cast at the level of interacting plant components. The model explicitly simulates growth, development and competition for light at the level of leaves, branches, etc., located in 3D space. In this way, we are able to explore the manner in which the low-level processes governing plant growth and development give rise to individual-, cohort-, and community-level phenomena. In particular, we show that individual-level trade-offs between growing up and growing out arise naturally in the model, and robustly give rise to cohort-level phenomena such as self-thinning, and community processes such as the effect of ecological disturbance on the maintenance of biodiversity. We conclude with a note on our methodology and how to interpret the results of simulation models such as this one.     

Effects of spatial pattern and relatedness in an experimental plant community

Many plant species show limited dispersal resulting in spatial and genetic substructures within populations. Consequently, neighbours are often related between each other, resulting in sibling competition. Using seed families of the annuals Capsella bursa-pastoris and Stachys annua we investigated effects of spatial pattern (i.e. random versus aggregated) on total and individual performance at the level of species and seed families under field conditions. At the level of species, we expected that inferior competitors increase, while superior competitors decrease their performance within neighbourhoods of conspecifics. Thus, we expected a species by spatial pattern interaction. Sibling competition, however, might reduce the performance of competitors, when genetically related, rather than non-related individuals are competing. Therefore, aggregations at the level of seed families could decrease the performance of competitors. Alternatively, if the opposite outcome would be observed, kin selection might be hypothesized to have occurred in the past. Because heavy seeds are expected to disperse less than light seeds, we further hypothesized that kin selection might be more likely to occur in superior competitors with heavy, locally dispersed seeds (e.g. Stachys) compared to inferior competitors with light, more distantly dispersed seeds (e.g. Capsella). We found a significant species by spatial pattern interaction. Indeed, the inferior competitor, Capsella, showed increased reproductive biomass production in aggregated compared to random patterns. Whereas, the performance of the superior competitor, Stachys, was to some extent decreased by intraspecific aggregation. Although statistically not significant, effects of intrafamily aggregations tended to be rather negative in Capsella but positive in Stachys. Our results confirmed that spatial patterns affect growth and reproduction of plant species promoting coexistence in plant communities. Although, we could not provide strong evidence for sibling competition or kin selection, our results suggested that competition among relatives was more severe for Capsella (lighter seeds) compared to Stachys (heavier seeds).     

A mathematical approach to spatial distribution and temporal succession in plant communities

The Volterra equations which represent competitions between two species are utilized to examine the phenomenon of boundary formation between two species of plants. The set of stable stationary points for these equations is determined and is illustrated in a product space of parameters and dynamical variables. The stages of boundary appearance and succession are visualized by considering slow changes of the parameters as functions of time and space.     

Long‐term compositional responses of a South African mesic grassland to burning and mowing

Question: What is the long‐term compositional response of grass and forb species to various combinations of burning and mowing? Can these responses be predicted from simple plant traits? Location: Ukulinga research and training farm (24°24′E, 30°24′S), Pietermaritzburg, KwaZulu‐Natal, South Africa. Methods: Grass species relative abundance in 1996 in various burning and mowing treatments of a long‐term (> 50 a) experiment was calculated from data obtained using a point sampling method, whereas forb species abundance in 1999 was determined using the importance score method. The experiment consisted of different frequencies (annual, biennial and triennial) of burning and mowing in winter or spring in combination with different frequencies of summer mowing (none, early, late or both). Results: Grasses responded to the type of disturbance (burn or mow) and frequency of burning, whereas forbs responded primarily to the presence or absence of any form of disturbance and secondarily to the timing of burning (spring versus winter). Summer mowing and annual or biennial dormant‐period burning maintained communities dominated by short grasses, whereas tall grasses dominated under annual dormant‐period mowing, triennial burning or protection from disturbance. Grass tillering strategy (below‐ or above‐ground) influenced response to burning frequency. Many erect herbaceous dicot species with aerial meristems were reduced in abundance by summer mowing whereas most small creeping herbaceous dicot species appeared to be dependent upon summer mowing. Conclusions: This long‐term experiment demonstrated that: (1) grasses and forbs responded differently to burning and mowing; (2) simple plant traits, such as height, position of tiller initiation, and position of meristems have potential for predicting the response of species to the timing and frequency of burning and mowing.     

Temporal trends in species composition and plant traits in natural grasslands of Uruguay

Abstract. We report the successional trends of the major life‐forms (graminoids and forbs) in natural grasslands of Uruguay over a 9‐yr period after the removal of domestic herbivores. For the whole community, species richness and diversity decreased over the successional period. In graminoids we observed clear temporal trajectories in floristic composition; the rate of floristic change decreased with time and was associated with a shift in plant traits. The exclusion of large herbivores promoted erect and tall grasses with narrow leaves and greater seed length, vegetative growth constrained to the cool season and increased frequency of annual species. Forbs did not show a clear temporal trend in species composition, but there was, nevertheless, a plot‐specific species turnover of this functional group that was reflected in their attributes. Species spreading by means of rhizomes, with vegetative growth restricted to the warm season. Species with larger seeds increased under grazing exclusion, as did annual and nitrogen‐fixing forbs. The floristic changes induced by cattle exclusion occurred early in the succession. This early high rate of change has practical implications for management and conservation programs of the natural grasslands of Uruguay. Additionally, the shift in plant traits may be helpful in devising simple indicators of grazing impact.     

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.     

Modelling potential impacts of climate change on the spatial distribution of zonal forest communities in Switzerland

Abstract. A spatially explicit, climate-sensitive vegetation model is presented to simulate both present and future distribution of potential natural vegetation types in Switzerland at the level of zonal forest communities. The model has two versions: (1) a ‘basic’ version using geographical region, aspect, bedrock (represented by soil pH), and elevation, and (2) a ‘climate-sensitive’ version obtained by replacing elevation (complex environmental gradient) with temperature (climatic factor). Version 2 is used to predict vegetation response under different (today's and projected) climatic conditions. Two regional climate scenarios are applied: (1) assuming an annual mean temperature increase of 1.1 — 1.4 °C, and (2) assuming an increase of 2.2 — 2.75 °C. Both scenarios result in significant changes of the spatial vegetation patterns as compared with today's climatic conditions. In scenario 1, ca. 33 % of the sample points remain unchanged in terms of the simulated zonal forest community; in scenario 2, virtually all sample points change. The most noticeable changes occur on the Swiss Plateau with Carpinion forests (zonal vegetation of present colline belt) expanding to areas that are occupied today by submontane and low-montane Fagus forests. To estimate the reliability of the simulation, quantitative (comparison with field mapping) and qualitative (comparison with climate types in the Alpine region) tests are performed and the main limitations of the approach are evaluated.     

Plant zonation at salt marshes of the endangered cordgrass <Emphasis Type="Italic">Spartina maritima</Emphasis> invaded by <Emphasis Type="Italic">Spartina densiflora</Emphasis>

The South American cordgrass Spartina densiflora is invading European salt marshes getting into contact with the indigenous and endangered low-marsh dominant, Spartina maritima. This work describes the evolution of the plant zonation during 7 years in a marsh of S. maritima invaded by S. densiflora. S. maritima appeared throughout the whole intertidal gradient from 1.72 to 3.33 m over Spanish Hydrographic Zero (SHZ), showing its higher biomasses and shoot densities at low elevations. In contrast, S. densiflora only invaded upper areas (>+2.59 m SHZ) at the centre of circular tussocks of S. maritima. Above-ground biomass of S. maritima dropped drastically at maximum occupation of space by the alien, and its shoot density and above-ground biomass decreased at S. densiflora zone during the study. The competitive potential of S. densiflora was reflected in high above- and below-ground biomass and shoot densities, accompanied by elevated wrack accumulation and the absence of other marsh plants presented together with S. maritima from areas dominated by S. densiflora. S. densiflora altered the native vegetational zonation pattern through the invasion of the centre of S. maritima tussocks; however, the alien invasion may be limited by the presence of the autochthonous cordgrass at lower elevations. Handling editor: Luis Mauricio Bini     

A spatial simulation model for vegetation dynamics

A fine-resolution, spatially explicit, stochastic model was developed to simulate the dynamics of species cover abundance and pattern in a single vegetation layer wherein neighbouring individuals are assumed to compete for growing space. Each species in the model is characterized by a small number of morphological and life-history parameters, which enter into equations that stand for a minimal set of vegetation processes. The model performed well in reproducing post-fire successional trends among the three codominant dwarf shrubs in a Dutch heathland community as recorded in an annually mapped permanent quadrat. Program inputs, outputs and an example of sensitivity analysis are illustrated. With suitable changes, the model could potentially describe any plant community in which the vertical structure is simple and community dynamics are determined by spatial interactions among neighbouring plants.Jacques de Smidt provided data, information and stimulus for this project. The model was developed during ICP's visit to Utrecht, arranged by Marinus Werger and financed by The Netherlands Science Research Council (ZWO). We also thank David Glenn-Lewin and Ernst Lippe for discussion and cooperation.     

Effects of temporal and spatial heterogeneities created by consumer-driven nutrient recycling on algal diversity

A spatially explicit plant-herbivore model composed of planktonic herbivores, algal preys and nutrients was constructed to examine the effects of consumer-driven nutrient recycling (CNR) on the algal species richness with and without spatial structure. The model assumed that either of two essential nutrients (N and P) limited growth of algal populations and that consumer individuals moved randomly in the lattice and grazed all the algal species with the same efficiency. The results showed that when there was no CNR, the number of persistent algal species was affected by neither supply rates of external nutrients nor spatial structure and was consistently low. When consumers recycled nutrients according to their stoichiometry, the algal species richness changed with supply rates of external nutrients depending on spatial structure: the algal species richness decreased with increasing nutrient loadings when there were no spatial structure because CNR increased the probability of stochastic extinction of algal species by amplifying the oscillation of algae-consumer dynamics. However, when spatial structures were created by the migration of consumers, CNR increased the algal species richness in a range of nutrient loadings because spatial variation of grazing pressure functioned to stabilize the algal-consumer dynamics. The present study suggests that through grazing and nutrient recycling, consumer individuals can create ephemeral heterogeneity in growth environments for algal species and that this ephemerality is one of the keys to understanding algal species in nature.     

镇江内江湿地不同演替阶段植物群落小气候日动态

2005年5月,选择镇江内江湿地具有代表性的裸地、虉草群落和芦苇群落,分别代表植被群落的不同演替阶段,测定不同群落、不同层次的光照强度、气温、土温和空气相对湿度,研究其植物群落小气候的日动态.结果表明,随演替由裸地到虉草群落到芦苇群落进行,群落内光照强度、气温和土壤温度均明显降低,日变幅减小.其中,日均光照强度由1 204.7 μmol·m^-2·s^-1降至141.28 μmol·m^-2·s^-1,日均变幅由1 126μmol·m^-2·s^-1降至265 μmol·m^-2·s^-1;日均气温由32.2 ℃降至24.9 ℃,日均变幅由12.75 ℃降至4.8 ℃;日均土温由21.83 ℃降至19.47 ℃,日均变幅由4.5 ℃降至2.1 ℃.群落内空气相对湿度明显升高(由58.95%增至87.3%）,变幅减小(由29.75%降至5.15%）.生境具有早期的开放性和后期的封闭性,小气候环境朝着更为阴、凉、湿的环境变化,且波动性减弱,稳定性增强.各群落内的光强、气温、湿度及土温之间均存在一定相关,但不同演替阶段各因子间相关程度各异.     

Effects of local interaction range and mobility on the spatio-temporal dynamics of competing animals in uniform habitats

Reduction of oscillations in population size is of fundamental importance to both theoretical and applied ecology. Spatial variability in population rates among different habitat regions is known to be an important mechanism that inhibits oscillations in population size. In the current study we used an individual-based model to simulate a single population of animals whose individual members are sensitive to competition only within their vicinity (i.e., within their competition neighborhood, CN). Our model extends previous studies by exploring how local interactions reduce population oscillations in competitive systems of animals, rather than in systems of plants. Our simulations explored the effects of animal mobility and interaction range separately on population dynamics. In our model, a decrease in CN dimensions tended to reduce population oscillations at all tested animal movement speeds. Yet, movement speed affected animal distribution patterns; an increase in movement speed led to more random distributions. We also found that mean population size was affected more by CN dimensions at lower mobility levels than when it was high.