A trait‐based trade‐off between growth and mortality: evidence from 15 tropical tree species using size‐specific relative growth rates

A life‐history trade‐off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged‐over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species‐specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed‐effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed‐effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade‐off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade‐off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

Strategic light manipulation as a restoration strategy to reduce alien grasses and encourage native regeneration in Hawaiian mesic forests

Question: Is there a light level at which alien grass biomass is reduced while still supporting growth and survival of native woody species, allowing for native species regeneration in abandoned pastures? Location: Island of Hawaii, USA. Methods: In a two‐part study we examined the effect of light availability on common native woody and alien grass species found in secondary forests in Hawaii. A field survey was conducted to examine the relationship between light availability and canopy type (open pasture, planted canopy and secondary forest) on understory grass biomass and litter accumulation. We then experimentally manipulated light levels to determine the effect of light availability on growth and survival of six native woody species and three alien grasses. Low‐light (5%), medium‐light (10%) and high‐light (20‐30%) treatments were created using shade structures erected beneath the existing secondary koa canopy. Results: In the field survey, alien grass biomass was greatest under the open pasture and lowest in the secondary forest. There was a positive correlation between understory light availability and alien grass biomass. In the experimental study, large reductions in relative growth rates were documented for all of the grass species and four of the six woody species under the lowest light level. Although growth at 5% light is substantially reduced, survival is still high (84‐100%), indicating that these species may persist under closed canopy. Conclusion: Low‐light conditions result in the greatest reduction in alien grass biomass while creating an environment in which native woody species can grow and survive.     

Differing Life History Characteristics Support Coexistence of Tree Soil Generalist and Specialist Species in Tropical Rain Forests

Niche differentiation is a prominent mechanism for explaining tree species coexistence in tropical rain forests. Theory predicts that species that are specialized on a set of environmental conditions should competitively exclude generalists in those conditions, and that environmental heterogeneity allows the coexistence of many different species based on niche specialization. Yet, many tropical tree species of the family Dipterocarpaceae have broadly similar habitat preferences, with some occurring widely across several soil types. These soil generalists clearly persist despite the presence of other dipterocarp species that show clear soil specialization. We evaluate comparative seedling performance (growth and survival) of six Shorea species (Dipterocarpaceae) which differ in their adult tree habitat associations within the Sepilok Forest Reserve (Borneo, Malaysia). We tested the hypothesis that seedlings of species associated with a particular soil type perform better on that soil type than seedlings of soil generalist species, with generalists in turn performing better than specialists on a different soil type. We conducted a reciprocal transplant experiment including two soil types (alluvial and mudstone) and two light treatments (gap and understory). The soils differed in soil acidity, Al and P concentration. Observed species‐specific differences in seedling responses to soil, light, and occasional flooding events could partially explain observed adult distribution, although not all species could be fully differentiated along these axes. Other trade‐offs, such as investment in defense against herbivores and tolerance to soil waterlogging, may play additional roles in explaining coexistence of these species.     

Leaf litter decomposition rates in degraded and fragmented tropical rain forests of Borneo

Previously extensive tracts of primary rain forest have been degraded by human activities, and we examined how the effects of forest disturbance arising from habitat fragmentation and commercial selective logging affected ecosystem functioning in these habitats by studying leaf litter decomposition rates in litter bags placed on the forest floor. The rain forests of Borneo are dominated by trees from the family Dipterocarpaceae, and we compared leaf litter decomposition rates of three dipterocarp species at eight forest fragment sites (area 3–3529 ha) that had different histories of disturbance pre‐fragmentation: four fragments had been selectively logged prior to fragmentation and four had been formed from previously undisturbed forest. We compared these logged and unlogged forest fragments with sites in continuous forest that had been selectively logged (two sites) and fully protected and undisturbed (two sites). After 120 d, undisturbed continuous forest sites had the fastest rates of decomposition (52% mass loss). Forest fragments formed from unlogged forest (32% mass loss) had faster decomposition rates than logged forest fragments (28% mass loss), but slower rates than continuous logged forest (39% mass loss). Leaves of a light‐demanding species (Parashorea malaanonan) decomposed faster than those of a shade‐tolerant species (Hopea nervosa), but decomposition of all three dipterocarp species that we studied responded similarly to logging and fragmentation effects. Reduced decomposition rates in logged and fragmented forest sites may affect nutrient cycling and thus have detrimental consequences for forest regeneration. Conservation management to improve forest quality should be a priority, particularly in logged forest fragments.     

Performance Trade-offs Driven by Morphological Plasticity Contribute to Habitat Specialization of Bornean Tree Species

Growth-survival trade-offs play an important role in niche differentiation of tropical tree species in relation to light-gradient partitioning. However, the mechanisms that determine differential species performance in response to light and soil resource availability are poorly understood. To examine responses to light and soil nutrient availability, we grew seedlings of five tropical tree species for 12 mo at < 2 and 18 percent full sunlight and in two soil types representing natural contrasts in nutrient availability within a lowland dipterocarp forest in North Borneo. We chose two specialists of nutrient-rich and nutrient-poor soils, respectively, and one habitat generalist. Across all species, growth was higher in high than low light and on more nutrient rich soil. Although species differed in growth rates, the ranking of species, in terms of growth, was consistent across the four treatments. Nutrient-rich soils improved seedling survival and increased growth of three species even under low light. Slower-growing species increased root allocation and reduced specific leaf area (SLA) and leaf area ratio (LAR) in response to decreased nutrient supply. All species increased LAR in response to low light. Maximum growth rates were negatively correlated with survival in the most resource-limited environment. Nutrient-poor soil specialists had low maximum growth rates but high survival at low resource availability. Specialists of nutrient-rich soils, plus the habitat generalist, had the opposite suite of traits. Fitness component trade-offs may be driven by both light and belowground resource availability. These trade-offs contribute to differentiation of tropical tree species among habitats defined by edaphic variation.     

Recovery of tree and mammal communities during large‐scale forest regeneration in Kibale National Park,Uganda

Tropical landscapes are changing rapidly as a result of human modifications; however, despite increasing deforestation, human population growth, and the need for more agricultural land, deforestation rates have exceeded the rate at which land is converted to cropland or pasture. For deforested lands to have conservation value requires an understanding of regeneration rates of vegetation, the rates at which animals colonize and grow in regenerating areas, and the nature of interactions between plants and animals in the specific region. Here, we present data on forest regeneration and animal abundance at four regenerating sites that had reached the stage of closed canopy forest where the average dbh of the trees was 17 cm. Overall, 20.3 percent of stems were wind‐dispersed species and 79.7 percent were animal‐dispersed species, while in the old‐growth forest 17.3 percent of the stems were wind‐dispersed species. The regenerating forest supported a substantial primate population and encounter rate (groups per km walked) in the regenerating sites was high compared to the neighboring old‐growth forests. By monitoring elephant tracks for 10 yr, we demonstrated that elephant numbers increased steadily over time, but they increased dramatically since 2004. In general, the richness of the mammal community detected by sight, tracks, feces, and/or camera traps, was high in regenerating forests compared to that documented for the national park. We conclude that in Africa, a continent that has seen dramatic declines in the area of old‐growth forest, there is ample opportunity to reclaim degraded areas and quickly restore substantial animal populations.     

Trade‐offs in seedling growth and survival within and across tropical forest microhabitats

For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade‐offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade‐offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade‐offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51–53%) followed by substrate type (35–37%) and light (11–12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade‐offs among species during regeneration are common in low‐diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.     

Positive effects of ectomycorrhizal colonization on growth of seedlings of a tropical tree across a range of forest floor light conditions

In a shadehouse experiment we tested the effects of light, nutrients and ectomycorrhizal fungi (EMF) on the growth of Vatica albiramis van Slooten (Dipterocarpaceae) seedlings. We hypothesised that it is more advantageous for plants to form connections with EMF and to trade carbon for nutrients with EMF under high light than low light. The relationship between seedling growth and the proportion of ectomycorrhizal root tips was expected as positive in high light and as negative in low light. Light conditions simulated the forest understory (low; 3% full sunlight), a small gap (medium; 11%) and a large gap (high; 33%) and a fully factorial combination of nutrients (F?/+) and ectomycorrhizal colonization (EMF?/+) treatments were applied within light conditions. The application of EMF and nutrients did significantly alter seedling growth across the range of forest floor light conditions, however the key hypothesis was rejected as seedling growth under low light was not affected by increased EMF colonization of root tips (light:EMF colonization χ²?=?2.97, p?=?0.23). In addition, the lack of difference in morphotype abundance across light conditions indicated that light changes may not favour the association to specific EMF in seedlings of this particular dipterocarp species. Our results suggest that antagonistic (non-beneficial to the plant) effects due to ectomycorrhizal colonization under a light constrained environment may not affect seedling growth of Vatica albiramis.     

Growth and photosynthetic responses of the federally endangered shrub, Lindera melissifolia (Lauraceae), to varied light environments

Photosynthetic, morphological, and growth responses to light environment can be useful measurements to determine favorable habitat conditions for the conservation of endangered species. For Lindera melissifolia (Walt.) Blume, we compared morphological and photosynthetic responses under natural and controlled light regimes, and growth under three light treatments: 100%, 42%, and 19% full sunlight. Typical sun-shade morphological responses to decreasing light levels included decreased stomatal density, increased specific leaf area, and increased leaf area ratio. Photosynthetic capacity (3-6 μmol CO(2) · m(-2) · s(-1)) was consistent with other shade-tolerant species. Light-saturated rates of photosynthesis of experimental plants increased with increasing light up to 42% sunlight, but declined at 100% sunlight. The 100% light treatment also resulted in lower plant biomass, primarily from a reduction in root biomass. Results indicate that canopy conditions at levels below 40% sunlight are optimal for plant growth and should be considered in management and reintroduction efforts for this species.     

米心水青冈幼苗对光照和养分的响应

在模拟郁闭林下(L1,大约1%~2%的全日照)、林窗(L2,大约18%的全日照)、开阔地(L3,全日照)3个光照水平和每个光照水平下进行施肥(F1)和不施肥(F0)对照的6个实验处理条件下,研究了2年龄米心水青冈(FagusenglerianaSeem)幼苗在随后的两个生长季里的生长对光照和土壤养分的响应。结果显示:光照和养分对幼苗高度、基径和生物量有显著的影响。经过两个生长季,L1处理下幼苗高度增量极显著地小于L2和L3处理下幼苗高度增量。L1处理下基径和生物量的增量在处理当年秋就极显著地小于L2和L3处理下的增量,并在第二年差异继续扩大。L2和L3处理下的幼苗间的生长没有显著差异。施肥明显地促进了L2和L3处理下的幼苗的生长,但对L1处理下的幼苗没有明显的作用。这些结果说明,虽然2年龄米心水青冈幼苗能够在林下的弱光条件下生存,但生长受到了极大的抑制。幼苗在林窗的中等光照条件下能够与在开阔地全日照条件下生长的一样好或更好,这与许多耐阴的落叶树的响应一样。在比林内光照强度较高的条件下施肥或较高的土壤养分才对米心水青冈幼苗的生长和生存起作用。     

Differential Responses of Dipterocarp Seedlings to Soil Moisture and Microtopography

Niche diversification is prominent among the mechanisms proposed to explain tropical rain forest tree diversity, with many studies focusing on trade‐offs among shade tolerance and growth. Less obvious is the impact of occasional, ephemeral and often minor disturbances on tree seedling survival. We propose that differential tolerances to soil waterlogging can contribute to the distribution of tree seedling communities along microtopographical gradients. We test this hypothesis experimentally by evaluating survival and performance of planted seedlings across microtopographical gradients in a periodically inundated tropical rain forest environment. Survival and relative growth rates were assessed for six Shorea (Dipterocarpaceae) species in Sepilok Forest Reserve (Sabah, Malaysia) over a 2‐yr period, during which seedlings were subjected to two brief flooding events. The species were selected on the basis of soil habitat affinities, with two species being primarily associated with low‐lying alluvial flats subject to inundation, two being associated with non‐flooded mudstone hills, and two species occurring in both habitats. Seedling performance was related to microtopographic elevation within and among plots and to soil moisture among plots. The faster growing species, Shorea argentifolia, Shorea leprosula and Shorea parvifolia, tended to be more vulnerable to high soil moisture in terms of mortality than the three species with lower growth rates. Within plots, soil moisture was inversely correlated with microelevation, and seedlings located at higher microelevations had an increased probability of survival. Microtopographical differences in seedling position could therefore contribute to species assembly processes through differential mortality, particularly in areas subject to minor and ephemeral flooding events.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Beta‐Diversity of Termite Assemblages Among Primary French Guiana Rain Forests

Termites are an important component of tropical rain forests, and have been included in many studies focusing on the influence of human disturbance. Their distribution among primary rain forest has, however, rarely been investigated. Here we studied the termite fauna in seven mostly undisturbed forest sites, representing several rain forest types. Overall, approximately 70 percent of species were soil‐feeders and 25 percent were wood‐feeders, the remaining 5 percent being classified here as litter‐feeders. Termite species richness did not differ significantly among sites, but sites differed in termite abundance. The palm swamp and the low forest situated on the foothills of an inselberg, hosted different termite communities to the other sites. These two sites presented a singular physiognomy suggesting that forest type is an important factor influencing species composition. We found no correlation between termite species composition and distance between sites, highlighting that at the scale of our study (about 100 km), forest sites share a similar species pool.     

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

Question: 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.     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

Coexistence of two congeneric tree species of Lauraceae in a secondary warm‐temperate forest on Miyajima Island,south‐western Japan

We analyzed the spatial distributions of two congeneric tree species, Neolistea aciculata and Neolistea sericea (Lauraceae), in a warm‐temperate forest on Miyajima Island, south‐western Japan. Both species were mainly found in valley sites on the island. Hence, these species shared the same topographic habitat niche. However, we found a clear difference between the spatial distributions of the two species in relation to the light environment. Neolistea aciculata was predominantly found in stands with low light, such as beneath the canopy of dense evergreen broadleaved forest. In contrast, N. sericea was predominantly associated with ample light, such as in secondary Pinus densiflora forest. In stands with moderate light conditions, both species were found. This habitat niche segregation in relation to light conditions presumably allows the coexistence of these two species in the predominantly successional forest on Miyajima Island.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

4种木本植物叶片的光合电子传递和吸收光能分配特性对光强的适应

以气体交换和叶绿素荧光测定相结合的方法研究了亚热带自然林乔木荷树、黧蒴和林下灌木九节、罗伞幼苗的光合电子传递及激发能利用的分配对生长光强的适应特性。4种植物生长于100%、36%和16%的自然光下8个月,叶片的光化学速率和热能耗散速率随光强增大而提高,热能耗散占总的光能吸收的比例也因光强不同而改变,16%光下的相对热耗散率约为40%～45%,100%自然光下增大至50%～75%。叶片总的非环式电子流速率及其分配到光呼吸的比例在100%光强下最高。乔木和灌木的电子传递和光能分配特性在16%光下相似,在100%光下差别较明显。除灌木种有较高的热耗散比例之外,其余的参数皆比乔木的低。结果表明乔木与灌木皆可通过提高激发能热耗散比例和提高光合电子传递向光呼吸的比例来适应于高光强条件。     

Shade tolerance,canopy gaps and mechanisms of coexistence of forest trees

The belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche differentiation in response to gaps has focused on evidence for an interspecific tradeoff between low light survival and high light growth. The objectives of this study are first to distinguish the possible mechanisms allowing coexistence based on this tradeoff, and second, to explore their limitations. We present a theory of forest dynamics driven by small‐scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non‐linearity’, ‘the successional niche’ and the ‘storage effect’. We find that temporal fluctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. The rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.