Gradient Of Light Availability Research Articles

Functional traits determine trade-offs and niches in a tropical forest community.

How numerous tree species can coexist in diverse forest communities is a key question in community ecology. Whereas neutral theory assumes that species are adapted to common field conditions and coexist by chance, niche theory predicts that species are functionally different and coexist because they are specialized for different niches. We integrated biophysical principles into a mathematical plant model to determine whether and how functional plant traits and trade-offs may cause functional divergence and niche separation of tree species. We used this model to compare the carbon budget of saplings across 13 co-occurring dry-forest tree species along gradients of light and water availability. We found that species ranged in strategy, from acquisitive species with high carbon budgets at highest resource levels to more conservative species with high tolerances for both shade and drought. The crown leaf area index and nitrogen mass per leaf area drove the functional divergence along the simulated light gradient, which was consistent with observed species distributions along light gradients in the forest. Stomatal coordination to avoid low water potentials or hydraulic failure caused functional divergence along the simulated water gradient, but was not correlated to observed species distributions along the water gradient in the forest. The trait-based biophysical model thus explains how functional traits cause functional divergence across species and whether such divergence contributes to niche separation along resource gradients.

Long-term physiological and morphological acclimation by the evergreen shrub Buxus sempervirens L. to understory and canopy gap light intensities

Physiological and morphological plasticity are essential for growth and reproduction in contrasting light environments. In dry forest ecosystems, light generalists must also cope with the trade-offs involved in synchronous acclimation to light availability and drought. To understand how the broadleaf evergreen tree-shrub Buxus sempervirens L. (common box) inhabits both understory and successional terrain of Mediterranean forest, we measured photosynthesis–fluorescence light response, morphological traits and architectural characteristics across a light gradient. Our results show that B. sempervirens exhibits stress resistance syndrome, with little change in net photosynthesis rate across a light availability gradient, due to compensatory physiological and morphological acclimation. Light energy processing and dissipation potential were highest in leaves of well-illuminated plants, with higher electron transport rate, fraction of open photosystem II reaction centres, non-photochemical quenching, photorespiration and dark respiration. In contrast, traits reducing light capture efficiency were observed in high light shrubs, including higher leaf mass per unit area, leaf clumping, leaf inclination and branch inclination. We suggest that both physiological and morphological plasticity are required for B. sempervirens to survive across a light gradient in a dry forest ecosystem, while exhibiting homoeostasis in photosynthetic gas exchange. We further speculate that the low growth rate of B. sempervirens is effective in full sun only due to a lack of competition in low resource microsites.

Levantamento e características ecológicas de orchidacea e da mata ciliar do rio dourados, Dourados-MS

A família Orchidaceae é uma das maiores entre as plantas com flores e apresenta grande importância do ponto de vista da conservação. Apesar disso, não existe nenhum trabalho sobre as orquídeas de Mato Grosso do Sul. Nesse sentido, o objetivo desta pesquisa foi realizar o levantamento das espécies de Orchidaceae e de aspectos de sua ecologia na Mata Ciliar de um fragmento de Floresta Estacional Semidecidual, às margens do rio Dourados (Dourados, MS). O levantamento foi realizado em sistema de varredura, sendo, nas epífitas, avaliadas as disposições vertical e horizontal sobre os forófitos. Para caracterização do microclima foram utilizados termo-higrômetro e luxímetro. Identificaram-se 17 espécies, pertencentes a 13 gêneros. Dos gêneros levantados, os de maior abundância foram: Acianthera, Macradenia e Capanemia. Relacionaram-se, ainda, as disposições vertical e horizontal das Orchidaceae com os gradientes inversos de disponibilidades hídrica e luminosa. Algumas espécies mostraram-se sensíveis à categorização em níveis de altura, enquanto outras pareciam ocorrer com frequência similar ao longo de todo o forófito. Em relação à orientação cardeal, a aparente resposta preferencial pelas faces sul e leste foi associada à insuficiência amostral e à menor disponibilidade hídrica que poderia ocorrer em razão de a face norte estar oposta ao espelho d'água.

Photosynthetic performance of the aquatic macrophyte Althenia orientalis to solar radiation along its vertical stems

We have studied the plasticity of the photosynthetic apparatus in the endangered aquatic macrophyte Althenia orientalis to the gradient of light availability within its meadow canopy. We determined diurnal change in situ irradiance, light quality, in vivo chlorophyll a fluorescence, ex situ oxygen evolution rates, respiration rate and pigment concentration. The levels of photosynthetic photon flux density (PFD) and ultraviolet radiation (UVR) and the red/far-red ratio decreased with depth within the canopies of A. orientalis. Apical leaves had a greater decrease of the maximal quantum yield (F(v)/F(m)) in the morning and a faster recovery rate in the afternoon than those in the basal ones. The relative electron transport rate (ETRr) was not saturated at any time of the day, even in the apical leaves that received the highest light. The maximum light-saturated rate of gross photosynthesis (GP(max)) took place in apical leaves around noon. The chlorophyll a/b ratio values were higher, and the chlorophyll/carotenoid ratio values lower, in apical leaves than basal ones. The highest concentrations in total carotenoids were reached in the apical leaves around noon. A. orientalis has a high capacity to acclimatize to the changes in the light environment, both in quality and quantity, presenting sun and shade leaves in the same stem through the vertical gradient in the canopy.

Eco‐evolutionary differences in light utilization traits and distributions of freshwater phytoplankton

We compiled light utilization traits for 56 species of freshwater phytoplankton to analyze group differences, trait trade‐offs, and allometric scaling relationships. We also used these traits to explain differences in major group distributions along the light availability gradient in 527 lakes in the continental United States. Major taxonomic groups differed significantly in their light utilization traits. Cyanobacteria had the highest initial slope of the growth‐irradiance curve (ɑ) and low irradiance at the onset of photoinhibition, indicating adaptation to low light environments. Green algae had the highest maximal growth rates and low ɑ, indicating adaptation to higher light environments. Groups capable of mixotrophy had traits indicative of poor light competitive abilities and high light requirements. Key light utilization traits scaled allometrically with cell size and exhibited trade‐offs leading to contrasting ecological strategies; ɑ and cell size were conserved at the highest taxonomic level (domain), indicating a fundamental trait divergence between prokaryotic and eukaryotic phytoplankton. In line with these trait differences, major groups showed different responses to light availability in natural conditions. The relative abundances of low light–adapted groups declined with increasing light availability and vice versa. The genera mean values of the initial slopes of the growth‐irradiance curves were significantly negatively correlated with the slopes of the relationships between the genus's relative abundance and light availability characterized by Secchi depth in 527 lakes. This indicates that light utilization traits can be used to explain phytoplankton distributions in nature.

Influence of light intensity and nutrient source on δ13C and δ15N signatures in Ulva pertusa

Stable isotopes are increasingly used to infer sources of nutrient enrichment and trophic linkages in coastal marine systems, although the utility of these tools often depends upon a pre- dictable expression of δ 15 N and δ 13 C signatures by primary producers. Accordingly, we examined how tissue δ 15 N and δ 13 C values change in the common coastal marine alga Ulva pertusa Kjellman under contrasting light and nutrient treatments. In Expt 1, we manipulated nitrogen and phosphorus enrichment and light availability in a factorial design and found that: (1) δ 13 C values in the tissue of U. pertusa depended upon interactions between light and nutrient availability, and there was no clear, overarching relationship between tissue δ 13 C values and growth rate; and (2) these effects yielded a substantial (11.6‰) range of variation in δ 13 C values. In Expt 2, we manipulated natural light (shaded versus unshaded) and nitrogen form (nitrate versus ammonium) in a factorial design and found that (3) δ 15 N of U. pertusa tissue was closely tied to δ 15 N of source nitrogen under all treat- ments and (4) δ 15 N differences between high and low light treatments were largest when U. pertusa was supplied with ammonium (3.7‰), relative to nitrate (0.8‰). The variation in δ 13 C values has implications for studies that use stable isotopes to infer trophic relationships in coastal marine envi- ronments, where gradients in nutrient concentration and light availability are common. The compar- atively small range of δ 15 N values expressed in U. pertusa supplied with nitrate confirms that this spe- cies represents a good proxy for δ 15 N of biologically available nitrogen in nitrate-dominated coastal seawater.

Simulating regeneration and vegetation dynamics in Mediterranean coniferous forests

This study aims to provide a quantitative framework to model the dynamics of Mediterranean coniferous forests by integrating existing ecological data within a generic mathematical simulator. We developed an individual-based vegetation dynamics model, constrained on long-term field regeneration data, analyses of tree-rings and seed germination experiments. The simulator implements an asymmetric competition algorithm which is based on the location and size of each individual. Growth is parameterized through the analysis of tree-rings from more than thirty individuals of each of the three species of interest. A super-individual approach is implemented to simulate regeneration dynamics, constrained with available regeneration data across time-since-disturbance and light-availability gradients. The study concerns an insular population of an endemic to Greece Mediterranean fir ( Abies cephalonica Loudon) on the island of Cephalonia (Ionian Sea) and two interacting populations of a Mediterranean pine ( Pinus brutia Ten.) and a more temperate-oriented pine ( Pinus nigra Arn. ssp. pallasiana) on the island of Lesbos (NE Aegean Sea), Greece. The model was validated against plot-level observations in terms of species standing biomass and regeneration vigour and adequately captured regeneration patterns and overall vegetation dynamics in both study sites. The potential effects of changing climatic patterns on the regeneration dynamics of the three species of interest were subsequently explored. With the assumption that a warmer future would probably cause changes in the duration of cold days, we tested how this change would affect the overall dynamics of the study sites, by focusing on the process of cold stratification upon seed germination. Following scenarios of a warmer future and under the current model parameterization, changes in the overall regeneration vigour controlled by a reduction in the amount of cold days, did not alter the overall dynamics in all plant populations studied. No changes were identified in the relative dominance of the interacting pine populations on Lesbos, while the observed reduction in the amount of emerging seedlings of A. cephalonica on Cephalonia did not affect biomass yield at later stages of stand development.

Leaf morphology correlates with water and light availability: What consequences for simple and compound leaves?

Leaves are organs sensitive to environmental changes in the process of evolution and may exhibit phenotypic plasticity as a response to abiotic stress. However, affirmation of leaf morphological plasticity and its regulations in different environments are still unclear. We performed a simulated experiment to study the variations of leaf morphology in different gradients of water and light availability. Considering different types of leaves and venation, we chose pinnate-veined simple leaves of Quercus acutissima and compound leaves of Robinia pseudoacacia as the study objects. The morphological parameters we investigated include leaf size, shape and venation pattern which can be easily measured in the field. Significant variations occurred in many parameters due to the effects of the environment and/or allometry. There were broadly consistent trends for leaf morphological variations along the gradients. The leaf size became smaller with a short supply of resources. Leaf elongation and fractions of the lamina area altered to enhance resources acquisition and conservation. Trade-offs between investments in support and functional structures optimized the venation pattern of major and minor veins. Leaflets partially played a role such as leaf teeth, for they are not only individual units, but also a part of the compound leaf. We suggest that more or less the same trends in morphological variations may be an important explanation for coexisting species to adapt to similar habitats and form the niche differentiation.

Species patterns in foliar nitrogen concentration, nitrogen content and 13C abundance for understory saplings across light gradients

Soil nitrogen (N) supply and uptake by regenerating trees is an important ecosystem attribute but difficult to quantify in partial-cut forests where light availability varies. The foliar attributes of N concentration (N%) and N per unit area (Na) may help characterize the influence of soil nutrition, but ideally the relationship between soils and foliage would be tested separately by species across well-defined light gradients. To do this, we examined foliar attributes of four tree species across gradients of light availability in 12 year-old partially-cut forests in northwest British Columbia, Canada. There were no differences in forest floor or mineral soil N mineralization rates across the light gradients, and for western hemlock (Tsuga heterophylla) and hybrid white spruce (Picea glauca x sitchensis), this consistent level of soil N supply corresponded with unchanging foliar N%. In contrast, foliar N% of Betula papyrifera (paper birch) and Thuja plicata (western redcedar) declined with shading, perhaps due to shifts in root-shoot biomass allocation for B. papyrifera, and climatic constraints on shade tolerance for T. plicata. Leaf δ13C approached an asymptote at approx. 40% full light for the coniferous species, but increased linearly with light for B. papyrifera. Foliar Na was linearly correlated with leaf δ13C for three species, reflecting the dual effect of light and nutrition on photosynthesis processes, and suggesting that foliar Na may be a simple parameter to integrate both resource constraints on regenerating saplings. These results demonstrate both support for and limits to foliar attributes among species in isolating soil N effects against light constraints in partial-cut forests.

Estimating colonization potential of migrant tree species

AbstractPlant populations migrating in response to climate change will have to colonize established communities. Even if a population disperses to a new region with a favorable climate, interactions with other species may prevent its establishment and further spread. The potential of these species to grow along with residents will be a critical factor controlling their response to climate change. To determine the capacity of migrating species to colonize established communities we conducted extensive long‐term transplant experiments where potential tree migrant species, i.e. species within ‘migration range,’ were planted side by side with resident ones. Potential immigrants were selected to be representative species of their native communities. For both groups, residents and potential migrants (17 species), we compared their growth response along gradients in soil moisture and light availability. Rather than manipulate climate directly, we exploited natural microclimatic gradients and the fluctuations in climate that occurred during the 5‐year experiment. Experimental results were used to estimate growth in the context of novel climate and relevant establishment factors. Results suggest that potential immigrant species had similar growth rates in the new environment than those from resident species ensuring their ability to establish in the area. However, contrary to our expectations, the soil moisture requirements for the immigrant group were similar to those of the resident species. These results could have major implications for vegetation changes under the predicted drier climate for the region. If it is the case that neither resident species nor potential migrants are able to maintain stable populations, the region may experience a decline in local biodiversity.

Modeling rule-based behavior: habitat selection and the growth-survival trade-off in larval cod

Environmental variation can cause significant fluctuations in the survival of larval fish and plankton. Understanding these fluctuations is critical for developing more accurate fisheries models, which are needed for both scientific and socioeconomic research. Growth, survival, and dispersal of marine planktonic larvae rely strongly on their behavior. Larval fish change their vertical positioning due to strong vertical gradients in light, temperature, predation pressure, and prey availability. Here, we explore how various behavioral rules predict vertical distribution, growth, and survival of larval cod (Gadus morhua) in a numerical model. The rules determine the trade-offs between larval growth, feeding rate, and predation rate, including their dependence on gut fullness and body mass. We evaluated the survival through size classes for different rules and random behavior and compared model predictions with observed larval distribution patterns. The rules predicted the correct average depth position with larval size, but failed to predict the timing of the observed vertical distribution pattern. However, model simulations revealed significant increases in survival for larval and juvenile cod with active behavior compared with larvae with random behavior. Behavior was important across all sizes of fish, and this study illustrates the value or added information of incorporating behavior in biophysical models. Copyright 2009, Oxford University Press.

Effects of Severe Tropical Cyclone Larry on rainforest vegetation and understorey microclimate near a road, powerline and stream

Abstract Severe Tropical Cyclone Larry damaged a large swathe of rainforest to the west of Innisfail in north‐eastern Queensland on 20 March 2006. Within the path of the most destructive core of the cyclone were sites previously established along human‐made (powerlines and highways) and natural (streams) linear canopy openings for a study of edge effects on adjacent rainforest plant communities and associated microclimates. Vegetation damage and understorey microclimate parameters were measured 6 months after the passage of Cyclone Larry and compared with results before the cyclone. We examined the spatial patterns of vegetation damage in relation to natural and artificial linear clearing edges and the vegetation structural factors influencing these patterns as well as resulting alterations to microclimate regimes experienced in the rainforest understorey. Vegetation damage was spatially patchy and not elevated near linear clearing edges relative to the forest interior and did not differ between edge types. Vegetation damage was influenced, albeit relatively weakly, by structural traits of individual trees and saplings, especially size (diameter at breast height, d.b.h.) and successional status: tree damage was greater in pioneer species and in larger trees, while sapling damage was greater in canopy tree species than in understorey tree or shrub species. Changes in the understorey microclimate mirrored the degree of damage to vegetation. Where vegetation damage appeared greater, the understorey microclimate was brighter, warmer, drier and windier than below less‐damaged areas of the forest canopy. Overall, understorey light availability, wind speed and the diurnal ranges of air temperature and vapour pressure deficit increased dramatically after Cyclone Larry, while pre‐cyclone edge gradients in light availability were lost and temperature and vapour pressure deficit gradients were reversed.

Physiological characteristics of the seagrass Posidonia sinuosa along a depth-related gradient of light availability

Physiological plasticity has the capacity to prolong seagrass survival under reduced light conditions. However, when light is attenuated across a depth gradient, the relative importance of this over other mechanisms for tolerating long-term light reduction (such as morphological adjust- ments and shoot loss) has been questioned. This study aimed to describe a number of the physio- logical characteristics of Posidonia sinuosa Cambridge et Kuo along a depth-related gradient of light availability (1.6 to 9.0 m depth) and infer how these characteristics are important for the long-term maintenance of the meadow. Rapid light curve-derived parameters, light harvesting pigments, photoprotective pigments and nutrient and carbohydrate concentrations exhibited few differences among depth strata, but showed some (albeit limited) adjustment between the seasons. It was inferred that some physiological plasticity is possible in P. sinuosa but that differences in the depth- related gradient of long-term light availability were not sufficient to induce physiological differences, even at the depth limit. Shoot density reductions, which reduce the effects of self-shading, possibly offset depth-related light reductions. Because the physiological characteristics we examined did not explain the adaptations by P. sinuosa to the long-term gradient of light availability, they may not be useful indicators of long-term light reduction.

Photosynthesis and respiration decline with light intensity in dominant and suppressed Eucalyptus globulus canopies.

Within canopy gradients in light-saturated photosynthesis (Amax), foliar nitrogen ([N]area) and leaf dark respiration (R15) were studied in the canopies of dominant and suppressed trees within an even-aged (4-year-old) Eucalyptus globulus (Labill) stand in southern Tasmania. Despite being an even-aged stand growing in a relatively uniform environment with respect to nutrient and water availability, the stand exhibited considerable structural complexity. Diameter at 1.3 m ranged between 3 cm and 21 cm, trees average 12 m height and stand leaf area index was ~6 m2 m-2 leading to strong gradients in light availability. We were interested in understanding the processes governing canopy production in trees of contrasting dominance classes. Vertical gradients in photosynthesis and foliar respiration were studied within the canopies of dominant and suppressed trees during 2006 and 2007. Amax varied from ~18 μmol m-2 s-1 in the upper canopy to 3 μmol m-2 s-1 at lower canopy positions. On average, Amax were higher in the dominant trees than in the suppressed trees. However, at any given height, Amax were similar in suppressed and dominant trees and were strongly related to leaf nitrogen content. Dark respiration varied from ~1.4 μmol m-2 s-1 in the upper canopy to 0.2 μmol m-2 s-1 in the lower canopy positions. In contrast to the patterns for Amax, dark respiration rates in the suppressed trees were higher than dominant trees at similar canopy positions. Respiration rates were also strongly related to [N]area and to Amax.

Morphological, growth and meadow characteristics of the seagrass Posidonia sinuosa along a depth-related gradient of light availability

Morphological and growth characteristics of the meadow-forming seagrass Posidonia sinuosa (Cambridge et Kuo), were measured along a depth-related gradient of light to infer its response to long-term differences in light availability. Morphometric measurements were carried out at 6 depths between 1.6 and 9.0 m in summer and winter at Cockburn Sound and summer only at Warnbro Sound in south-western Australia. The minimum light requirement for P. sinuosa of 8.5% sub-surface light was among the lower range reported for seagrasses. Its slow growth rate (0.5–1.5 mgdry shoot^–1 d^–1), relative to similarly sized species, may contribute to the low light requirements of this species. Shoot density, leaf area index and biomass showed pronounced and consistent differences among depths (up to 88-fold reduction of above-ground biomass from shallow to deep sites). At the deeper sites, the reduced shoot density probably reduces respiratory demand and alleviates self-shading. Morphological differences (leaf length, width and thickness and number of leaves per shoot) did not follow a clear and consistent trend with depth. Despite a 70% reduction in light availability at the canopy level between the shallowest and deepest sites, leaf growth rate was unaffected by depth during summer, and in winter differed between only a few depths. We propose that the reduction in shoot density partially alleviates the effects of self-shading and permits comparable leaf growth rates across the depth range. These results suggest that for interpreting long-term responses to light availability, shoot density is the most sensitive of the morphological characteristics measured here.

Variation in juvenile survival and related physiological traits among dipterocarp species co‐existing in a Bornean forest

AbstractQuestion: What environmental factors and physiological traits determine the juvenile survival rates of 11 dipterocarp species (Dipterocarpaceae) co‐occurring in a Bornean tropical rain forest?Location: Tropical northwestern Borneo.Methods: In 248 quadrats, distributed over 12 ha, the juvenile (height< 2.0 m) survival of 11 shade‐tolerant dipterocarp species was monitored for one year. The probability of survival for each species was expressed as a logistic function of plant height and three environmental factors: canopy openness, soil water potential and conspecific individual density. The correlation between survival under shaded conditions and several physiological traits of juveniles was tested by normal Pearson correlation and a phylogeny based manner using independent contrasts.Results: Seven species had increased survival under more open canopies, one species had increased and two species had decreased survival at drier sites. Six species had higher survival as the density of conspecifics increased, whereas two species had lower survival. The magnitudes of the effects for the three environmental factors were similar. However, the interspecific difference in survival was largely determined by a dependence of small seedlings on the light environment. Survival under low light was correlated positively with root dry mass and negatively with total leaf area of juveniles.Conclusions: Responses of seedlings to the fine‐scale gradient of light availability were considerably different among the 11 species. Species that suffered increased mortality under low light had a set of structural traits that would potentially allow rapid growth under bright conditions. Differential performance along the light gradient may contribute to the stable coexistence of these species.

Counteracting gradients of light and soil nutrients in the understorey of Mediterranean oak forests

Abstract. The forest canopy modifies the availability of resources (light, water, and soil nutrients) in the understorey. In this paper we analyze the relationships between woody canopy density, litter accumulation, and topsoil N and P availability in the understorey of two oak forests: one in southern Portugal and the other in southern Spain. Both forests persist on low-nutrient soils, particularly poor in P. We hypothesize that direct and indirect effects of the canopy overstorey cause opposite gradients in the availability of essential resources (light and key soil nutrients) in the understorey. In both studied forests we found significant relationships between the overall canopy density, light availability, topsoil litter accumulation, and the availability of N and P, which frequently limit plant growth. Path analysis (by Shipley’s d-sep method) showed that the available data were consistent with the proposed causal model. The average values of soil variables at the end quartiles of the light-availability gradient were compared. Results showed large differences in litter accumulation (~30×) and available-N and -P topsoil concentrations (~3×) in the Spanish forest (with the wider environmental gradient). Furthermore, P increased from the “very low” range to the “low” or even the “optimum” range of availability (according to standard plant growth criteria), which suggests potential effects on the growth of the understorey plant species. We conclude that the counteracting gradients of the essential resources – light and nutrients – in the forest understorey resulted from direct and indirect effects of the canopy overstorey, respectively. We suggest that these counteracting effects of the woody canopy on essential resources of different nature must be considered when interpreting the patterns of understorey plant populations and communities.

Phenotypic plasticity and biomass allocation pattern in three Dryopteris (Dryopteridaceae) species on an experimental light-availability gradient

We were interested in whether the contrasting regional distribution patterns of three congeneric, frequently co-occurring fern species (Dryopteris carthusiana, D. dilatata and D. expansa) could be explained by differential biomass allocation strategies and different phenotypic plasticities to light availability. The morphology and habitat preference of these ferns are known to be very similar, but in Estonia, their frequencies of occurrence differ sharply––Dryopteris carthusiana is common, D. expansa grows in scattered localities, and D. dilatata is rare. We grew the species under different levels of illumination (100, 50, 25 and 10% of full daylight) in an experimental garden to compare their autecological responses to shading. After one growing season there were clear interspecific differences in total plant biomass accumulation––D. carthusiana > D. expansa > D. dilatata––indicating the possible competitive inferiority of the latter at the young sporophyte stage. D. expansa was the least shade-tolerant, with biomass decreasing sharply under less than 50% illumination; D. dilatata was the most shade-tolerant, with similar growth at all illumination levels. In relative biomass allocation patterns, the most notable differences among species were in the relative shares of biomass stored in rhizomes. In D. carthusiana and D. expansa this share was nearly constant and independent of the illumination conditions. D. dilatata allocated very little biomass into rhizome in deep shade, but was able to increase this share more than twofold in 50% light. Dryopteris dilatata was clearly shown to be morphologically the most plastic of the three. In four traits––rhizome mass, frond:below-ground biomass ratio, stipe length and specific leaf area––its degree of ontogenetic plasticity to light was significantly higher than that of D. expansa and D. carthusiana. While the general performance (biomass production) of species in the experiment coincided with that observed in nature, the results of plasticity estimation were somewhat surprising––it is difficult to explain the inferior performance of a species (D. dilatata) through high morphological plasticity. Probably, the species is rare either because of certain climatic restrictions, or because it is presently expanding its distribution and is in the phase of invading Estonian understory communities.

THE ECONOMICS OF MUTUALISMS: OPTIMAL UTILIZATION OF MYCORRHIZAL MUTUALISTIC PARTNERS BY PLANTS

Can choice of mutualistic partners and the degree of their utilization determine (1) mutualistic partner coexistence, (2) relative abundance of mutualistic partners, and (3) environment-dependent changes in relative abundance? We investigate these questions in the context of the plant-mycorrhizal fungal mutualism by building a biological market model potentially applicable to other mutualisms as well. We examine the situation where a single plant selectively utilizes member(s) of a group of ectomycorrhizal potential trading partners. Under biologically realistic circumstances, the plant may simultaneously utilize multiple partners, its degree of utilization determining the community structure of the fungi. If utilization of multiple partners is optimal, the marginal cost of acquiring additional nitrogen from every trading partner must be equal while the marginal cost of acquiring it from any unutilized partner must be larger. Because the plant's nitrogen demand is light dependent, the composition of the fungal species among its trading partners changes along light-availability gradients. We discuss the design of an experiment to test the key prediction of our model, the equalization of marginal cost.

A bounded null model explains juvenile tree community structure along light availability gradients in a temperate rain forest

Ecologists have proposed that tree species may coexist by specialising on light environments associated with gaps of different sizes. Remarkably few community‐level studies, however, have actually examined juvenile tree distributions along light availability gradients. Here we describe distributions of juvenile trees in relation to canopy openness in a temperate rainforest, and test the hypothesis that competitive sorting causes coexisting species to overlap less in light environment occupancy than would be expected by chance. Average overlap of species’ interquartile ranges on the canopy openness gradient was tested against a bounded domain null model of community structure which used range‐size criteria to constrain random placement of species optima. Microsite availability was strongly skewed towards low light, with 43% of microsites occurring at <5% canopy openness. We therefore transformed canopy openness values to ranks, so that equal intervals on the transformed gradient represented equal areas of microsite availability. We then calculated the interquartile range (25–75%) of sample ranks occupied by juveniles of each species. About half the assemblage was non‐randomly distributed in relation to canopy openness, providing evidence of niche expression. Average overlap of species’ interquartile ranges did not depart significantly from that predicted by the bounded null model, indicating that community structure in relation to canopy openness was mainly explained by a mid‐domain effect. As predicted by the null model, species’ interquartile range mid‐points were concentrated in the centre of the rank‐transformed gradient, and species richness (overlap of interquartile ranges) peaked close to the median light environment. Most species therefore had intermediate light requirements. The apparent lack of constraints on pairwise overlap suggest that differences in light use are not a prerequisite for tree species coexistence. As far as we are aware, this is the first study to identify a mid‐domain effect on a resource availability axis.