The confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Seed Size,Seed Germination,and Seedling Survival of Brazilian Tropical Tree Species Differing in Successional Status1

Seed germination and seedling establishment patterns have been used to classify species as shade tolerant or intolerant. The main objective of this research was to investigate, under controlled conditions, seed germination of species from different successional positions as well as to follow seed germination and seedling survival under natural shade in the field. The species studied were Solarium granuloso‐leprosum, Trema micrantha, Cecropia pachystachya, Croton piptocalyx, Bauhinia forficata subsp. pruinosa. Senna macranthera, Schizolobium parahyba, Piptadenia gonoacantha, Chorisia speciosa, Pseudobombax grandiflorum, Ficus guaranitica, Esenbeckia leiocarpa, Pachystroma longifolium, Myroxylon peruiferum, and Hymenaea courbaril. Field trials were carried out at Santa Genebra Municipal Reserve, Campinas, SP, Brazil, at the forest edge and in the understory. No significant correlations were detected between successional status and seed size or seed water content. Light‐regulated germination was present only in small‐seeded species. In field experiments, most species, including the light‐sensitive ones, were able to germinate under the canopy, where a low red/far‐red ratio predominates. Most species, mainly those of early‐ and intermediate successional positions, presented low seedling survival rates under shade. Myroxylon peruiferum was the most shade tolerant species, while 5. granuloso‐leprosum, C. speciosa, P. gonoacantha, F. guaranitica, T. micrantha, and 5. parahyba were the most shade intolerant. These latter species showed little or no survival under the shade conditions.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.     

Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality

Questions: How does woody vegetation abundance and diversity differ after natural disturbances causing different levels of mortality? Location: Abies balsamea–Betula papyrifera boreal mixed‐wood stands of southeast Quebec, Canada. Methods: Woody vegetation abundance and diversity were quantified and compared among three disturbance‐caused mortality classes, canopy gap, moderate‐severity disturbances, and catastrophic fire, using redundancy analysis, a constrained linear ordination technique, and diversity indices. Results: Substantial changes in canopy tree species abundance and diversity only occurred after catastrophic fire. Shade‐tolerant, late‐successional conifer species remained dominant after canopy gap and moderate‐severity disturbances, whereas shade‐intolerant, early‐successional colonizers dominated canopy tree regeneration after catastrophic fire. Density and diversity of mid‐tolerant and shade‐intolerant understory tree and shrub species increased as the impact of disturbance increased. Highest species richness estimates were observed after catastrophic fire, with several species establishing exclusively under these conditions. Relative abundance of canopy tree regeneration was most similar after canopy gap and moderate‐severity disturbances. For the sub‐canopy tree and shrub community, relative species abundances were most similar after moderate‐severity disturbances and catastrophic fire. Vegetation responses to moderate‐severity disturbances thus had commonalities with both extremes of the disturbance‐caused mortality gradient, but for different regeneration layers. Conclusions: Current spatio‐temporal parameters of natural disturbances causing varying degrees of mortality promote the development of a complex, multi‐cohort forest condition throughout the landscape. The projected increase in time intervals between catastrophic fires may lead to reduced diversity within the system.     

Floristic changes and regeneration patterns for a 12‐year period during the 3rd and 4th decades following selection logging in a subtropical rainforest

Abstract Changes in regeneration patterns in a subtropical rainforest in north‐east New South Wales (Australia) are presented for a 12‐year period during the 3rd and 4th decades following repeated single‐tree selection logging. Changes were investigated using multivariate and univariate approaches. There were no significant differences in floristic assemblages within and between censuses. However, two contrasting trends of changes in plant groups were detected. In trees with a diameter at breast height (d.b.h.; that is, 1.3 m above the ground level) ≥ 10 cm, both the density and species richness increased in the shade‐tolerant group, while density increased and species richness decreased in the shade‐intolerant group. Among smaller sized regenerating species including trees (1.3 m in height < 10 cm d.b.h.), a general decrease in species richness was observed along with significant changes in stem densities where the number of stems in the shade‐tolerant species increased while that of both shade‐intolerant and vine species decreased. Excluding the vines and understorey species from the broader regenerating species, revealed a decrease in species richness in juvenile canopy tree, and a significant change in densities with the number of stems in the shade‐tolerant increasing while that of shade‐intolerant trees decreased. A comparison between the canopy trees ≥ 10 cm d.b.h. and the juvenile canopy trees group showed that these groups were tending towards similar floristic assemblages. These results suggest gradual replacement of shade‐intolerant by shade‐tolerant species as stands tend toward later stages of regeneration. This study shows that the inclusion of regenerating species in long‐term studies is both complementary to the larger plant component and more revealing of both trends and changes.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Regeneration niche of three epiphytic species of Gesneriaceae from Chilean rainforests: implications for the evolution of growth habits in Coronanthereae

Ecological and evolutionary studies of the epiphytic growth habit in angiosperms are limited. In this article, we assess the relationship between growth habit and regeneration niche in Coronanthereae (Gesneriaceae) and discuss its implications for the evolution of epiphytism in this lineage. In the temperate rainforest of southern Chile, we quantified the vertical distribution and experimentally examined the regeneration niche of three endemic species of Coronanthereae. One species was a holoepiphyte, which was more frequent in the upper canopy, and two species were secondary hemiepiphytes, which decreased in abundance with tree height. Seed germination of the holoepiphyte was higher on tree bark substrates and under open canopy than on forest soil and in the shade. In contrast, seed germination of both secondary hemiepiphytes did not differ between substrates (bark vs. soil) or light conditions (light vs. shade). Seedling survival percentage of secondary hemiepiphytes was higher on forest soil and under a closed canopy, thus behaving as shade‐tolerant species. In turn, the holoepiphyte behaved as a shade‐intolerant species. The reconstruction of the ancestral growth habits and regeneration niches on the inferred phylogenetic tree of Coronanthereae revealed that the specialized regeneration niche of Sarmienta repens, characterized by requirements of shade intolerance and germination on tree bark, was coupled with the evolution of the holoepiphytic growth habit. We conclude that differentiation in the regeneration niche is a key process in the evolution of epiphytic growth habits in Coronanthereae. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 79–92.     

Artificial canopy gaps accelerate restoration within an exotic Pinus radiata plantation

We created small‐scale artificial canopy gaps to accelerate the growth of mature indigenous forest canopy species for restoration of an 18‐year‐old exotic Pinus radiata plantation forest, in the Marlborough Sounds, New Zealand. Small and large circular gaps were formed by felling. Seedlings of two indigenous forest canopy species, Podocarpus totara (Podocarpaceae) and Beilschmiedia tawa (Lauraceae), were planted within artificial gaps and undisturbed plantation canopy. Seedling height growth, mortality, and occurrence of animal browse were monitored at approximately 6‐month intervals over 17 months. Both P. totara and B. tawa differed significantly in height growth and in animal browse occurrence among artificial gap treatments. Growth of the light‐demanding P. totara was better under large canopy gaps, whereas growth of the shade‐tolerant B. tawa increased under gaps of any size but was most consistent under small gaps. For P. totara, any significant restoration benefit of gap formation on height growth was lost when browsed seedlings were taken into account. Animal browse significantly limited B. tawa height growth in large but not in small gaps. Small‐scale canopy gap creation is an effective method of modifying light transmission to the plantation understorey and accelerating seedling growth rates. Canopy gap size can be used to optimize understorey illumination according to species‐specific light requirements. The increased occurrence of animal browse in gaps requires consideration. Artificial canopy gaps within planted monocultures create structural heterogeneity that would otherwise take an extended period of time to develop. These results further support the role of plantations as indigenous forest restoration sites.     

Gap regeneration and replacement patterns in an old-growth <Emphasis Type="Italic">Fagus</Emphasis>–<Emphasis Type="Italic">Abies</Emphasis> forest of Bosnia–Herzegovina

We examined the influence of small-scale gap disturbances on stand development and tree species coexistence in an old-growth Fagus sylvatica–Abies alba forest in the Dinaric Mountains of Bosnia–Herzegovina. The structure and composition of tree regeneration in gaps were compared to the forest as a whole, and the influence of gap size on the density and composition of regeneration was assessed. Transition probabilities were also calculated from gapfillers in different life stages to examine canopy replacement patterns. The structure and composition of tree regeneration were similar between gaps and the forest as a whole, and there was no relationship between overall regeneration density and gap size, indicating most individuals established prior to gap formation. Likewise, there was no strong evidence of gap-size partitioning for shade tolerant F. sylvatica and A. alba, although less tolerant Acer pseudoplatanus only recruited to taller life stages in larger gaps. Transition probabilities calculated from the seedling and sapling data suggest that most gaps will be captured by F. sylvatica, while probabilities based on pole-sized gapmakers indicate both A. alba and F. sylvatica will be maintained in the canopy. We suggest that gaps primarily play a role in reorganizing advance regeneration, and that coexistence of shade tolerant F. sylvatica and A. alba is more likely related to their differential ability to tolerate shaded understory conditions, particularly during larger life stages, rather than gap-size partitioning.     

Vegetative Life History of a Dominant Rain Forest Canopy Tree1

Tetramerista glabra has a remarkable combination of life history traits. It is a dense-wooded, large, common canopy tree in primary peat swamp rain forest. Its seedlings, although shade tolerant, can grow rapidly in high light conditions, but frequently lack structural stability. Most juvenile stems (94% in the understory and 38% in canopy gaps) collapse under their own weight or from branchfalls. Fallen stems then ramify into vegetative sprouts, which in turn may collapse, perpetuating a vegetative juvenile cycle. Most recruitment is from sprouts rather than from seed. Structural analysis of stem dimensions shows that stems 2–8 cm DBH (diameter at breast height) are close to the theoretical buckling limit, especially for those dependent on neighboring vegetation to maintain vertical form. Trees > 4 cm DBH persisting as upright stems develop stilt root support and become structurally independent at ca 8 cm DBH. Eventually, as stems thicken, stilt roots anastamose and trees adopt the cylindrical growth form of mature canopy trees (up to 150 cm DBH). Thus, the vegetative life history strategy of the species is to: (i) regenerate a large “ramet bank” from the majority of juveniles that fail structurally while suppressed in the understoty, and (ii) to maximize height growth at the expense of diameter growth in high light conditions. The growth pattern and plastic form of T. glabra shows how a shade tolerant species may adapt to utilize the ephemeral light resource in canopy gaps. The growth strategy of this species allows it to circumvent the normal trade-off between shade tolerance and rapid growth in canopy gaps.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

The role of canopy gaps in the regeneration of coastal dune forest

In regenerating coastal dune forest, the canopy consists almost exclusively of a single species, Acacia karroo. When these trees die, they create large canopy gaps. If this promotes the persistence of pioneer species to the detriment of other forest species, then the end goal of a restored coastal dune forest may be unobtainable. We wished to ascertain whether tree species composition and richness differed significantly between canopy gaps and intact canopy, and across a gradient of gap sizes. In three known‐age regenerating coastal dune forest sites, we measured 146 gaps, the species responsible for gap creation, the species most likely to reach the canopy and the composition of adults, seedlings and saplings. We paired each gap with an adjacent plot of the same area that was entirely under intact canopy and sampled in the same way. Most species (15 of 23) had higher abundance in canopy gaps. The probability of self‐replacement was low for A. karroo even in the largest gaps. Despite this predominance of shade‐intolerant species, regenerating dune forest appears to be in the first phase of succession with ‘forest pioneers’ replacing the dominant canopy species. The nature of these species should lead to successful regeneration of dune forest.     

浙江天童国家森林公园常绿阔叶林主要组成种的种群结构及更新类型

 以浙江天童国家森林公园常绿阔叶林为对象,应用永久样方法和每木调查法调查了群落种类组成和结构,并对主要组成种的种群结构进行了分析。依据胸径级频率分布的形状,将各树种的种群结构归纳为5种类型,并结合生物学、生态学特征,分别讨论了其更新类型,同时对调查群落所处的演替阶段进行了诊断。结果表明：1）群落为6种共优势群落,出现木本植物69种。2）种群结构为单峰型的马尾松（Pinus massoniana）、枫香（Liquidambar formosana）和檫木（Sassafras tzumu）等为阳性高大乔木,无正常更新能力,更新类型是先锋群落优势种或是顶极性先锋种;间歇型的木荷（Schima superba）、黑山山矾（Symplocos heishanensis）和红楠（Machilus thunbergii）等为具有耐荫性的常绿阔叶树种,更新具有波动性和机会性,是演替系列或顶极群落优势种;逆-J字型的米槠（Castanopsis carlesii）、栲树（Castanopsis fargesii）和长叶石栎（Lithocarpus harlandii）等为强耐荫性的常绿阔叶树种,通过幼苗库和根萌生枝进行更新,是顶极群落的优势种;浙江新木姜子（Neolitsea aurata var. chekiangensis）、厚皮香（Ternstroemia gymnanthera）等为L字型;连蕊茶（Camellia fraternna）、老鼠矢（Symplocos stellaris）等为单柱型,此二型是群落亚乔木层和灌木层的主要组成种。3）调查群落已进入演替的顶极阶段,但仍处于其前期,将进一步向以米槠、栲树为主要优势种的方向发展。     

Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments

 Light saturated photosynthesis (A) in field saplings of shade tolerant, intermediate, and intolerant tree species was analyzed for stomatal and nonstomatal limitations to test differences between species and sun and shade phenotypes during drought. Throughout the study, photosynthesis was highest and mesophyll limitations of A (L_m) lowest in the intolerant species in both open and understory habitats. The shade tolerant species exhibited the only drought-related decreased A and increased L_m in the open, and the greatest drought-related decreased A and increased L_m in the understory. Few species exhibited significant habitat or drought-related differences in stomatal conductance to CO₂ (g_c), but even slight decreases in g_c during drought were associated with large increases in stomatal limitations to A (L_g). Combined changes in L_m and L_g resulted in increased relative stomatal limitation to A (l _g) in several species during drought. Nevertheless, the overall lack of stomatal closure allowed for nonstomatal limitations to play a major role in reduced A during drought. Higher leaf N was associated with shallower slope of the l _g versus g_c relationship, an indication of greater A capacity. Photosynthetic capacity tended to be greater in the intolerant species than the tolerant species, and it tended to decrease during drought primarily in the shade tolerant species in the understory. Findings in the literature suggest that carbon reduction reactions may be more susceptible to drought than photosynthetic light reactions. If so, reduced carbon reduction capacity of shade tolerant species or shade phenotypes may predispose them to drought conditions, which suggests a mechanism behind the well-recognized tradeoff between drought tolerance and shade tolerance of temperate tree species. Received: 20 October 1995 / Accepted: 20 February 1996     

Small gap dynamics in the southern boreal forest of eastern Canada: Do canopy gaps influence stand development?

Question: To what extent do small‐scale disturbances in the forest canopy, created by natural disturbance agents, affect stand development? Doubts exist as to whether small canopy openings have any real effect on the understory tree recruitment, especially in boreal forests. Location: Conifer and mixed stands in the Gaspesian region in eastern Québec. The main natural disturbance agents are recurring outbreaks of Choristoneura fumiferana (eastern spruce budworm) and winds. Methods: Linear transects in 27 sites were used to describe the gap (< 0.1 ha) regime parameters, including gap fraction, gap size and change in disturbance severity through time. Three stand types were distinguished, based on a gradient of abundance of tree host species for the eastern spruce budworm. The impact of gaps was evaluated on the basis of changes in the number, the period of recruitment, and the composition of tree saplings present within gap areas. Changes were measured along the gap size gradient, and according to the pattern of recent budworm epidemics. Results: The gap fraction is highly variable (18%‐64%) and is on average relatively high (42%). Gap sizes have a positively skewed distribution. In most cases the growth rate among gap filling saplings increased sufficiently to date disturbance events. The composition and the structure of understory trees were affected by gap formation. The number of shade‐intolerant tree species did increase during or following periods of particularly severe canopy disturbances. However, the establishment or survival of shade intolerant species was not restricted to larger gaps or more intensely disturbed periods. Conclusions: In sub‐boreal forests of Eastern Canada, small scale disturbances in the tree canopy influence stand regeneration dynamics, but not to the extent that parameters such as sapling composition and recruitment patterns depend on gap regime characteristics.     

Are variations of direct and indirect plant interactions along a climatic gradient dependent on species’ strategies? An experiment on tree seedlings

Investigating how interactions among plants depend on environmental conditions is key to understand and predict plant communities’ response to climate change. However, while many studies have shown how direct interactions change along climatic gradients, indirect interactions have received far less attention. In this study, we aim at contributing to a more complete understanding of how biotic interactions are modulated by climatic conditions. We investigated both direct and indirect effects of adult tree canopy and ground vegetation on seedling growth and survival in five tree species in the French Alps. To explore the effect of environmental conditions, the experiment was carried out at 10 sites along a climatic gradient closely related to temperature. While seedling growth was little affected by direct and indirect interactions, seedling survival showed significant patterns across multiple species. Ground vegetation had a strong direct competitive effect on seedling survival under warmer conditions. This effect decreased or shifted to facilitation at lower temperatures. While the confidence intervals were wider for the effect of adult canopy, it displayed the same pattern. The monitoring of micro‐environmental conditions revealed that competition by ground vegetation in warmer sites could be related to reduced water availability; and weak facilitation by adult canopy in colder sites to protection against frost. For a cold‐intolerant and shade‐tolerant species (Fagus sylvatica), adult canopy indirectly facilitated seedling survival by suppressing ground vegetation at high temperature sites. The other more cold tolerant species did not show this indirect effect (Pinus uncinata, Larix decidua and Abies alba). Our results support the widely observed pattern of stronger direct competition in more productive climates. However, for shade tolerant species, the effect of direct competition may be buffered by tree canopies reducing the competition of ground vegetation, resulting in an opposite trend for indirect interactions across the climatic gradient.     

Differences in Survival and Growth Among Tropical Rain Forest Pioneer Tree Seedlings in Relation to Canopy Openness and Herbivory

Although differences in canopy openness, herbivory and their interaction may promote species coexistence, how these factors affect pioneer tree species and potentially limit growth, and survival has been poorly studied, particularly in tropical South Asia. We monitored the effect of canopy openness and herbivore damage on seedling survival and growth of 960 individuals of six pioneer tree species: Dillenia triquetra, Macaranga indica, Macaranga peltata, Schumacheria castaneifolia, Trema orientalis, and Wendlandia bicuspidata. Seedlings were placed in four gap‐understory positions—center, outer gap edge, inner forest edge, and understory—in four large, natural gaps within the Sinharaja World Heritage Reserve, Sri Lanka. Canopy openness positively affected survival probability beyond the 550‐d experiment, while herbivory decreased survival and was highest in understory conditions. The relative order of species survival stayed fairly consistent between gap‐understory positions and followed their known shade tolerance rankings. When averaged across all experimental conditions, T. orientalis had the lowest survival probability estimate beyond the 550‐d experiment (0.05), but the greatest capacity for growth where it successfully established, while the species with highest averaged survival probability (0.79), D. triquetra, showed the lowest growth. One species, W. bicuspidata, responded positively to herbivory by re‐sprouting. Coexistence of D. triquetra, T. orientalis, and W. bicuspidata can be explained by a trade‐off among species in survival, growth, and response to herbivory. In addition to variation in canopy light environment, herbivory may be important in determining pioneer species distribution through fine‐scale niche partitioning and should be carefully considered in reforestation efforts.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata