Functional ecology of congeneric variation in the leaf economics spectrum.

Variation in resource availability can lead to phenotypic plasticity in the traits comprising the world-wide leaf economics spectrum (LES), potentially impairing plant function and complicating the use of tabulated values for LES traits in ecological studies. We compared 14 Carex (Cyperaceae) species in a factorial experiment (unshaded/shaded × sufficient/insufficient P) to analyze how changes in the network of allometric scaling relationships among LES traits influenced growth under favorable and resource-limited conditions. Changes in leaf mass per area (LMA) shifted the scaling relationships among LES traits expressed per unit area vs mass in ways that helped to sustain growth under resource limitation. Increases in area-normalized photosynthetic capacity and foliar nitrogen (N) were correlated with increased growth, offsetting losses associated with mass-normalized dark respiration and foliar N. These shifts increased the contributions to growth associated with photosynthetic N-use efficiency and the N : P ratio. Plasticity in LMA is at the hub of the functional role of the LES as an integrated and resilient complex system that balances the relationships among area- and mass-based aspects of gas exchange and foliar nutrient traits to sustain at least some degree of plant growth under differing availabilities of above- and below-ground resources.

Complementary crops and landscape features sustain wild bee communities.

Wild bees, which are important for commercial pollination, depend on floral and nesting resources both at farms and in the surrounding landscape. Mass-flowering crops are only in bloom for a few weeks and unable to support bee populations that persist throughout the year. Farm fields and orchards that flower in succession potentially can extend the availability of floral resources for pollinators. However, it is unclear whether the same bee species or genera will forage from one crop to the next, which bees specialize on particular crops, and to what degree inter-crop visitation patterns will be mediated by landscape context. We therefore studied local- and landscape-level drivers of bee diversity and species turnover in apple orchards, blueberry fields, and raspberry fields that bloom sequentially in southern Quebec, Canada. Despite the presence of high bee species turnover, orchards and small fruit fields complemented each other phenologically by supporting two bee genera essential to their pollination: mining bees (Andrena spp.) and bumble bees (Bombus spp.). A number of bee species specialized on apple, blueberry, or raspberry blossoms, suggesting that all three crops could be used to promote regional bee diversity. Bee diversity (rarefied richness, wild bee abundance) was highest across crops in landscapes containing hedgerows, meadows, and suburban areas that provide ancillary nesting and floral resources throughout the spring and summer. Promoting phenological complementarity in floral resources at the farmstead and landscape scales is essential to sustaining diverse wild bee populations.

Similarities and differences in intrapopulation trait correlations of co-occurring tree species: consistent water-use relationships amid widely different correlation patterns.

PREMISE OF THE STUDY: General relationships among functional traits have been identified across species, but the forces shaping these relationships remain largely unknown. Adopting an approach from evolutionary biology, we studied similarities and differences in intrapopulation trait correlations among locally co-occurring tree species to assess the roles of constraints, phylogeny, and the environmental niche in shaping multivariate phenotypes. We tested the hypotheses (1) that intrapopulation correlations among functional traits are largely shaped by fundamental trade-offs or constraints and (2) that differences among species reflect adaptation to their environmental niches. METHODS: We compared pairwise correlations and correlation matrices of 17 key functional traits within and among temperate tree species. These traits describe three well-established trade-off dimensions characterizing interspecific relationships among physiological functions: resource acquisition and conservation; sap transport and mechanical support; and branch architecture. KEY RESULTS: Six trait pairs are consistently correlated within populations. Of these, only one involves dimensionally independent traits: LMA-δ13 C. For all other traits, intrapopulation functional trait correlations are weak, are species-specific, and differ from interspecific correlations. Species intrapopulation correlation matrices are related to neither phylogeny nor environmental niche. CONCLUSIONS: The results (1) suggest that the functional design of these species is centered on efficient water use, (2) highlight flexibility in plant functional design across species, and (3) suggest that intrapopulation, local interspecific, and global interspecific correlations are shaped by processes acting at each of these scales.

Evaluating general allometric models: interspecific and intraspecific data tell different stories due to interspecific variation in stem tissue density and leaf size.

The ability of general scaling models to capture the central tendency or dispersion in biological data has been questioned. In fact, the appropriate domain of such models has never been clearly articulated and they have been supported and challenged using both interspecific and/or intraspecific data. Here, we evaluate several simplifying assumptions and predictions of two prominent scaling models: West, Brown and Enquist's fractal model (WBE) and a null model of geometric similarity (GEOM). Using data for 53 herbaceous angiosperm species from the Songnen Grasslands of Northern China, we compared both the interspecific and intraspecific scaling relationships for plant geometry and biomass partitioning. Specifically, we considered biomass investment in shoots and leaves as well as related several traits not commonly collected in plant allometric analyses: shoot volume, leaf number, and mean leaf mass. At the interspecific level, we find substantial variation in regression slopes, and the simplifying assumptions of WBE and predictions of both the WBE and GEOM models do not hold. In contrast, we find substantial support for the WBE model at the intraspecific level, and to a lesser extent for GEOM. The differences between our results at interspecific and intraspecific levels are due to the fact that leaf size and stem tissue density vary considerably across species in contrast to the simplifying assumptions of WBE. These results highlight the domain within which simplifying model assumptions might be most appropriate, and suggest allometric models may be useful points of departure within some species, growth forms or taxonomic groups.

Experimental test for adaptive differentiation of ginseng populations reveals complex response to temperature.

BACKGROUND AND AIMS: Local climatic adaptation can influence species' response to climate change. If populations within a species are adapted to local climate, directional change away from mean climatic conditions may negatively affect fitness of populations throughout the species' range. METHODS: Adaptive differentiation to temperature was tested for in American ginseng (Panax quinquefolius) by reciprocally transplanting individuals from two populations, originating at different elevations, among temperature treatments in a controlled growth chamber environment. Fitness-related traits were measured in order to test for a population × temperature treatment interaction, and key physiological and phenological traits were measured to explain population differences in response to temperature. KEY RESULTS: Response to temperature treatments differed between populations, suggesting genetic differentiation of populations. However, the pattern of response of fitness-related variables generally did not suggest 'home temperature' advantage, as would be expected if populations were locally adapted to temperature alone. CONCLUSIONS: Failure consistently to detect a 'home temperature' advantage response suggests that adaptation to temperature is complex, and environmental and biotic factors that naturally covary with temperature in the field may be critical to understanding the nature of adaptation to temperature.

Contributions of leaf photosynthetic capacity, leaf angle and self-shading to the maximization of net photosynthesis in Acer saccharum: a modelling assessment.

BACKGROUND AND AIMS: Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ε). METHODS: A functional-structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI(ind)) together with a genetic algorithm to find distributions of leaf angle (L(A)) and leaf photosynthetic capacity (A(max)) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A(max) either unconstrained or constrained to an upper value consistent with reported values for A(max) in A. saccharum. KEY RESULTS: It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ε were simultaneously maximized. Maximization of ε required simultaneous adjustments in L(A) and A(max) along gradients of PPFD in the plants. When A(max) was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ε because PPFD incident on leaves was higher than the PPFD at which ε(max) was attainable. Average leaf ε in constrained plants nonetheless improved with increasing LAI(ind) because of an increase in self-shading. CONCLUSIONS: It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ε at the scale of leaves, which requires a highly integrated response between L(A), A(max) and LAI(ind). The results also suggest that to maximize ε plants have evolved mechanisms that co-ordinate the L(A) and A(max) of individual leaves with PPFD availability.

How do traits vary across ecological scales? A case for trait-based ecology.

Despite the increasing importance of functional traits for the study of plant ecology, we do not know how variation in a given trait changes across ecological scales, which prevents us from assessing potential scale-dependent aspects of trait variation. To address this deficiency, we partitioned the variance in two key functional traits (leaf mass area and leaf dry matter content) across six nested ecological scales (site, plot, species, tree, strata and leaf) in lowland tropical rainforests. In both traits, the plot level shows virtually no variance despite high species turnover among plots and the size of within-species variation (leaf + strata + tree) is comparable with that of species level variation. The lack of variance at the plot level brings substantial support to the idea that trait-based environmental filtering plays a central role in plant community assembly. These results and the finding that the amount of within-species variation is comparable with interspecific variation support a shift of focus from species-based to trait-based ecology.

Disjunct performance and distribution in the sedge Carex prasina.

Efforts to understand species distributions and predict responses to environmental changes depend on specifying how the abiotic environment determines distributions. At landscape scales, it is critical to distinguish effects of environmental factors from other mechanisms such as competition and dispersal limitation. We examined how environmental factors affect the distribution and performance of the sedge Carex prasina across a 10-km(2) old-growth forest in southern Québec. We isolated the effects of soil characteristics by conducting a greenhouse experiment that assessed the performance of C. prasina on soils from a range of wetland habitats where it could potentially occur. This allowed us to compare how the species' performance and its distribution across the landscape relate to the same soil characteristics. In the experiment, the biomass and leaf chlorophyll content of C. prasina increased with increasing soil organic matter (OM). Across the landscape, however, the species' probability of occurrence and abundance decreased with increasing soil OM. C. prasina had similar biomass on soils from sites where it did and did not occur, but it had higher leaf chlorophyll content on soils from sites where it did not occur. We found no evidence that differential performance across environments determines the distribution of this species, as C. prasina tended to occur on soils where it showed reduced performance. Rather, other processes such as competition or dispersal limitation likely override any direct effects of the soil environment on distribution. Our results caution against the common assumption that the environments where a species tends to occur or be most abundant are the environments where it performs best. C. prasina presents a clear example of a species whose performance, at least along edaphic gradients, cannot explain its distribution. This example highlights the importance of distinguishing the relative roles of biotic and abiotic factors that shape species distributions across landscapes.

Codominance of Acer saccharum and Fagus grandifolia: the role of Fagus root sprouts along a slope gradient in an old-growth forest.

We studied how the unusual capacity of mature Fagus grandifolia to form clumps of clonal stems from root sprouts can contribute to its frequent codominance with Acer saccharum in southern Quebec, Canada. In an old-growth forest, the degree of dominance by the two species shifted along topographic gradients spanning a few hundreds of meters, with Fagus more frequent on lower slopes and Acer on upper slopes. The frequency distribution of Fagus stem diameter had an inverse J distribution at all slope positions, which is indicative of continuous recruitment. Acer stem diameter also had an inverse J pattern, except at lower slope positions where size structure was discontinuous. For stems <2 m tall, Fagus regenerated mainly by sprouts at the upper and mid-slopes, while regeneration from seed was more pronounced on the lower slope. This change of regeneration mode affected the spatial pattern of Fagus stems. Understory trees of Fagus were positively correlated with conspecific canopy trees on upper and mid-slopes, but not on lower slopes where Fagus regenerated mainly by seedlings. Understory trees of Acer were positively correlated with conspecific canopy trees only on the mid-slope. There were many Fagus seedlings around Acer canopy trees at the lower slope, suggesting the potential replacement of Acer canopy trees by Fagus. This study suggests that the regeneration traits of the two species changed with slope position and that Fagus patches originating from root sprouts can contribute to the maintenance of Acer-Fagus codominance at the scale of local landscapes.

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content.

BACKGROUND AND AIMS: Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species. METHODS: Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0.5 Hz over 9-17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N(area)), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A(max)), dark respiration (R(d)), apparent quantum yield (phi) and curvature (theta)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A(daily)) for each leaf. KEY RESULTS: Leaf N(area) and A(max) changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A(daily) did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species. CONCLUSIONS: The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple 'big-leaves' in which all constituent 'small leaves' use PPFD with the same photosynthetic efficiency.

Do interspecific differences in sapling growth traits contribute to the co-dominance of Acer saccharum and Fagus grandifolia?

BACKGROUND AND AIMS: Acer saccharum and Fagus grandifolia are among the most dominant late-successional tree species in North America. The influence of sapling growth responses to canopy gaps on the co-dominance of the two species in an old-growth forest in southern Quebec, Canada was examined. Two predictions were evaluated: (a) F. grandifolia is more shade tolerant than A. saccharum due to greater sapling leaf area and net production per sapling in closed-canopy conditions; and (b) the height growth rate of A. saccharum in canopy gaps is greater than that of F. grandifolia due to increased net production per sapling. METHODS: Sapling crown allometry, net production and height growth rates were compared between and within the two species in closed canopy vs. canopy gaps. Standardized major axis regression was used to analyse differences in crown allometry. KEY RESULTS: F. grandifolia had greater crown projection, sapling leaf area and net production rate per sapling than A. saccharum in closed-canopy conditions. In response to canopy gaps, net production per sapling increased to the same degree in both species. The net production per sapling of F. grandifolia thus was much greater than that of A. saccharum in both canopy gap and closed-canopy conditions. The height growth rate of both species increased in canopy gaps, but the degree of increase was greater in F. grandifolia than in A. saccharum. CONCLUSIONS: F. grandifolia regenerated more successfully than A. saccharum in both closed-canopy conditions and canopy gaps, which indicates that the co-dominance of the two species cannot be maintained simply by interspecific differences in shade tolerance and growth in gaps. Previous research showed that although Fagus and Acer shared dominance at this site, their relative dominance shifted with edaphic conditions. This suggests that the widespread co-dominance of the two species in eastern North American forests is maintained by the joint influence of canopy disturbance and species-specific responses to the heterogeneity of moisture and fertility regimes within forested landscapes.

The ecological and functional correlates of nocturnal transpiration.

Contrary to the conventional theory of optimal stomatal control, there is substantial transpiration at night in many tree species, but the functional significance of this phenomenon remains uncertain. To investigate the possible roles of nocturnal transpiration, we compared and contrasted the correlations of both nocturnal and diurnal sap flow with a range of traits in 21 temperate deciduous tree species. These traits included soil water affinity, shade tolerance, cold hardiness, nitrogen concentration of tissues, minimum transpiration rate of excised leaves, growth rate, photosynthetic capacity, stomatal length and density, and the water potential and relative water content of leaves at the wilting point. Nocturnal sap flow was higher in species with higher leaf nitrogen concentrations, higher rates of extension growth and lower shade tolerances. Diurnal sap flow was higher in species with higher leaf nitrogen concentrations and photosynthetic capacities on a leaf area basis. Because leaf metabolism and dark respiration, in particular, are strongly related to leaf nitrogen concentration, our findings suggest that nocturnal transpiration functions to sustain carbohydrate export and other processes driven by dark respiration, and that this function is most important in fast- growing shade-intolerant tree species.

Toward synthesis of relationships among leaf longevity, instantaneous photosynthetic rate, lifetime leaf carbon gain, and the gross primary production of forests.

The assimilation of carbon by plant communities (gross primary production [GPP]) is a central concern in plant ecology as well as for our understanding of global climate change. As an alternative to traditional methods involving destructive harvests or time-consuming measurements, we present a simple, general model for GPP as the product of the lifetime carbon gain by a single leaf, the daily leaf production rate, and the length of the favorable period for photosynthesis. To test the model, we estimated leaf lifetime carbon gain for 26 species using the concept of mean labor time for leaves (the part of each day the leaf functions to full capacity), average potential photosynthetic capacity over the leaf lifetime, and functional leaf longevity (leaf longevity discounted for periods within a year wholly unfavorable for photosynthesis). We found that the lifetime carbon gain of leaves was rather constant across species. Moreover, when foliar biomass was regressed against functional leaf longevity, aseasonal and seasonal forests fell on a single line, suggesting that the leaf production rate during favorable periods is not substantially different among forests in the world. The gross production of forest ecosystems then can be predicted to a first approximation simply by the annual duration of the period favorable for photosynthetic activity in any given region.

Alternative designs and the evolution of functional diversity.

According to conventional wisdom, functional diversity is exclusively a consequence of species having evolved adaptations to fill different niches within a heterogeneous environment. This view anticipates only one optimal combination of trait values in a given environment, but it is also conceivable that alternative designs of equal fitness in the same environment might evolve. To investigate that possibility, we use a genetic algorithm to search for optimal combinations of 34 functional traits in a realistic model of tree seedling growth and survival. We show that separate lineages of seedlings evolving in identical environments result in many alternative functional designs of approximately equal fitness.

The comparative evidence relating to functional and neutral interpretations of biological communities.

Neutral and functional theories provide rival interpretations of community patterns involving distribution, abundance, and diversity. One group of patterns describes the overall properties of species or sites, and derives principally from the frequency distribution of abundance among species. According to neutral theory, these patterns are determined by the number of individuals of novel type appearing each generation in the community, whereas functional theory relates them to the distribution of the extent of niches. A second group of patterns describes the spatial attributes of communities, and derives principally from the decay of similarity in species composition with distance. Neutral theory interprets these patterns as consequences of local dispersal alone, whereas the functional interpretation is that more distant sites are likely to be ecologically different. Neutral theory often provides good predictions of community patterns, yet is at variance with a wide range of experimental results involving the manipulation of environments or communities. One explanation for this discrepancy is that spatially explicit models where selection is generally weak, or where selection acts strongly on only a few species at each site, have essentially the same output as neutral models with respect to the distribution of abundance and the decay of similarity. Detecting a non-neutral signal in survey data requires careful spatial or phylogenetic analysis; we emphasize the potential utility of incorporating phylogenetic information in order to detect functional processes that lead to ecological variation among clades.

Fundamental trade-offs generating the worldwide leaf economics spectrum.

Recent work has identified a worldwide "economic" spectrum of correlated leaf traits that affects global patterns of nutrient cycling and primary productivity and that is used to calibrate vegetation-climate models. The correlation patterns are displayed by species from the arctic to the tropics and are largely independent of growth form or phylogeny. This generality suggests that unidentified fundamental constraints control the return of photosynthates on investments of nutrients and dry mass in leaves. Using novel graph theoretic methods and structural equation modeling, we show that the relationships among these variables can best be explained by assuming (1) a necessary trade-off between allocation to structural tissues versus liquid phase processes and (2) an evolutionary tradeoff between leaf photosynthetic rates, construction costs, and leaf longevity.

Twilight far-red treatment advances leaf bud burst of silver birch (Betula pendula).

Bud development of boreal trees in spring, once initiated, is driven by ambient air temperature, but the mechanism triggering bud development remains unclear. We determined if some aspect of the diurnal or seasonal light regime influences initiation of bud burst once the chilling requirement is met. We grew 3-year-old birch plantlets cloned from a mature tree of boreal origin in light conditions realistically simulating the lengthening days of spring at 60 degrees N. To emulate the reduction in red to far-red light (R:FR) ratio between daylight and twilight, one group of plantlets was subjected to reduced R:FR ratio in the morning and evening in addition to progressively lengthening days, whereas the other group was subjected to the same R:FR ratio throughout the day. The reduced R:FR ratio of twilight advanced bud burst by 4 days compared with the reference group (P = 0.04). To assess the interplay between the fulfillment of the chilling requirement and the subsequent response to warming, we fitted a thermal time model to the data with separate parameterizations for the starting dates of heat sum accumulation in each treatment. Least-squares fitting suggested that bud development started in light regimes corresponding to late March, almost two months after the chilling requirement for dormancy release was satisfied. Therefore, shortening night length or increasing day length, or both, appears to be the cue enabling bud development in spring, with twilight quality having an effect on the photoperiodic response. If twilight alone were the cue, the difference in bud burst dates between the experimental groups would have been greater than 4 days. The result gives experimental support for the use of thermal-time models in phenological modeling.

The sampling characteristics of electivity indices.

Electivity indices measure the utilization of food types (r) in relation to their abundance or availability in the environment (p). Foods that constitute a larger proportion of the diet than of the available foods are considered preferred; conversely those proportionately underrepresented in the diet are avoided. A food is eaten at random if its proportion in the diet equals its proportion in the environment. A family of electivity indices stemming from Ivlev's (1961) classic monograph exist and differ only in the particular algorithm used to calculate electivity from r and p.For each available index I graphed the values of electivity as contours for all combinations of r and p. These graphs are compared to illustrate the strengths and weaknesses of each index on the basis of the following criteria: 1) the value of the index when r=p for a food, 2) the symmetry of the electivity value as feeding deviates from random, 3) the possible range of index values, 4) the linearity of changes in electivity over the full range of r and p, 5) the sensitivity of the index to sampling errors, 6) the statistical testability of the electivity, and 7) the stability of the electivity value for a food type that changes relative abundance or occurs in combination with different food types. No one index ideally satisfies all the criteria.The host preferences of gypsy moth, Lymantria dispar, feeding on tree foliage in an undisturbed deciduous forest in southwestern Quebec, Canada were used to compare the available indices: Ivlev's electivity, E; Ivlev's forage ratio, E'; Jacob's modified electivity, D; Jacob's modified forage ratio, log Q; Chesson's alpha; Strauss' linear index, L; and Vanderploeg and Scavia's relativized electivity, E *. The electivity values calculated by each index differ one from another; host trees shown as preferred by one index will frequently appear avoided according to an alternative index. The rank order electivities for the 19 available host trees, however, are remarkably similar for all but Strauss' linear index, L. Populus grandidentata, Quercus rubra, Ostrya virginiana, and Amelanchier were the most preferred host trees in the sampled forest; Prunus serotina, Acer pensylvanicum, A. rubrum, Betula lutea, and Fraxinus americana were most avoided. The use of Vanderploeg and Scavia's E * index is recommended.

Ecological trends in lichen photosynthesis.

Drawing on data for 42 fruticose and foliose lichen species, four aspects of photosynthetic response have been ex&mined in relation to latitude and climatic variables: 1) the maximal rate of net photosynthesis (Pmax), 2) the tissue temperature at which Pmax occurs (Topt), 3) the photon flux density at which Pmax occurs (PhARsat), and 4) the tissue water status at which Pmax occurs. Tissue water status was measured as either relative water content (RWCopt) or percentage dry weight (DWopt). The Topt decreased significantly with increasing latitude; Topt also increased significantly with increasing July solar radiation, July air temperature, daily hours of bright sunshine, daily solar radiation, annual precipitation, and annual evaporation. Both RWCopt and DWopt increased significantly with annual precipitation. The Topt and RWCopt for a lichen could both be more accurately predicted from multiple regressions on macroclimatic variables describing the lichen's habitat. Both the PhARsat and the Pmax, despite their lack of simple univariate relationships to either latitude or climatic variables, could also be predicted by multiple regression. The limitations on the generality and precision of these ecological trends in lichen photosynthesis are discussed in relation to microclimatic considerations, morphological adaptations, and poikilohydric metabolism.

The effects of climatic pattern on lichen productivity: Cetraria cucullata (Bell.) Ach. in the arctic tundra of northern Alaska.

The climatic control of productivity for two populations of the lichen Cetraria cucullata (Bell.) Ach. growing in the arctic tundra of northern Alaska (70°28'N, 157°23'W) was examined. Respiratory losses of carbon vary with tissue temperature, tissue water content, and time since wetting. Potential net photosynthetic gains of carbon are affected by photon flux density, tissue temperature, and water content. The net CO2 exchange responses of populations growing on ridge tundra and on upland tundra differ and these differences reflect possible adaptation to the normal environmental regimes in the two habitats. Simulation of the lichen's net carbon balance using continuous hourly records of photon flux density, temperature, and water content for the unusually dry period June 28 through July 17, 1977 show that lichen biomass is actually lost during climatic regimes leading to frequent but short periods of lichen metabolic activity. This result is confirmed by the negative relative growth rates measured for C. cucullata over the same monitoring period. This observed loss of biomass may be attributable to depletion of carbon reserves to reactivate dormant metabolism without sufficiently long periods favorable for net photosynthetic activity to replenish the lost reserves. These results illustrate that environmental limits exist on the success of the dormancy strategy characteristic of lichen and moss carbon metabolism.