Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.     

水青冈林中两种鹅耳枥种子萌发和幼苗的生存与生长（英文）

 千金榆(Carpinus cordata var．chinensis)和千筋树(Carpinus fargesiana)是四川省北部山地水青冈林中的两种主要落叶阔叶树。两种植物种子萌发和幼苗生存与生长的实验研究结果显示：千金榆种子秋季成熟落地后有一年多的休眠期,种子主要在落地后的第二和第三个生长季萌发;千筋树种子成熟落地后有的可在来年萌发,但大部分是在第二个生长季萌发;由于每2～3年两种树就有一个较大的结实年,每年都会有一些新的幼苗在林中出土;尽管两种树是耐荫的,但幼苗在较好光照条件下(如林窗)生存和生长明显好于在林冠遮荫的地方;施肥会加速幼苗的死亡,特别是在阴暗的林下,但在光照较好的条件下能促进幼苗的生长。结果说明两种鹅耳枥的幼苗可以在林下持续存在,但只有在较好的光照条件下,如稀疏的林冠下或林窗下才会有明显的净生长。     

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.     

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.     

Comparative demography of three coexisting Acer species in gaps and under closed canopy

Questions: 1. Is there a trade‐off between gap dependency and shade tolerance in each of the life‐history stages of three closely related, coexisting species, Acer amoenum (Aa), A. mono (Am) and A. rufinerve (Ar)? 2. If not, what differences in life‐history traits contribute to the coexistence of these non‐pioneer species? Location: Ogawa Forest Reserve, a remnant (98 ha), species‐rich, temperate deciduous forest in central Japan (36°56’ N, 140°35’ E, 600 ‐ 660 m a.s.l.). Methods: We estimated the demographic parameters (survival, growth rate and fecundity) by stage of each species growing in gaps and under closed canopy through observations of a 6‐ha permanent plot over 12 years. Population dynamics were analysed with stage‐based matrix models including gap dynamics. Results : All of the species showed high seedling and sapling survival rates under closed canopies. However, demographic parameters for each growth stage in gaps and under closed canopies revealed inter‐specific differences and ontogenetic shifts. The trade‐off between survival in the shade and growth in gaps was detected only at the small sapling stage (height < 30 cm), and Ar had the highest growth rate both in the shade and in the gaps at most life stages. Conclusions: Inter‐specific differences and ontogenetic shifts in light requirements with life‐form differences may contribute to the coexistence of the Acer species in old‐growth forests, with Aa considered a long‐lived sub‐canopy tree, Am a long‐lived canopy tree, and Ar a short‐lived,‘gap‐phase’ sub‐canopy tree.     

Spatial structure and regeneration of Tetramerista glabra in peat swamp rain forest in Indonesian Borneo

We examined the spatial structure and regeneration of Tetramerista glabra, a dominant canopy tree in peat swamp rain forest in Borneo (West Kalimantan, Indonesia). T. glabra has strong spatial structure that changes dramatically during the life cycle; seedlings were highly aggregated, saplings were random and trees were evenly distributed. Germination and seedling relative growth were highest within canopy gaps, but seedling densities were highest at gap edges. Concentration of seedlings in gap edges, rather than localized seed dispersal, was responsible for seedling patchiness. The slope of the relationship between relative growth rate and seedling height decreased from gap, to gap edge to understory habitats, suggesting that competition within the seedling layer is more important in gaps than in the understory. The processes that break down seedling aggregation, leading to over-dispersion of trees, must be density dependent, but remain unknown.     

Linking responses of native and invasive plants to hurricane disturbances: implications for coastal plant community structure

Hurricane disturbances produce significant changes in forest microclimates, creating opportunities for seedling regeneration of native and invasive plant species alike. However, there is limited information on how changes in microclimates and pre-existing forest conditions affect native and invasive plants responses to hurricane disturbances. In this manipulative study, we examined the responses of three common shrub/small stature tree species, two of which are native to the coastal region of the southeastern USA (Baccharis halimifolia and Morella cerifera) and one that is invasive (Triadica sebifera), to two key components of hurricane disturbance (canopy damage and saline storm surge). In a greenhouse, we grew seedlings of these species under a range of shade levels that mimicked pre-and post-hurricane canopy conditions for wet pine forest and mixed hardwood forest, two forest communities common in coastal areas of the southeastern USA. Seedlings were subjected to saline storm surges equivalent to full strength sea water for 3 days. Seedling responses (mortality and growth) to the treatments were monitored for 16 months. All species benefitted from higher canopy openness. Storm surge effects were short-lived and seedlings readily recovered under high light conditions. The storm surge had stronger negative effects on survival and growth of all species when coupled with high shade, suggesting storm surge has greater negative impacts on seedlings where hurricane winds cause minimal or no canopy damage. The invasive T. sebifera was by far more shade tolerant than the natives. Survival of T. sebifera seedlings under highly shaded conditions may provide it a competitive edge over native species during community reassembly following tropical storms. Differential responses of native and invasive species to hurricane disturbances will have profound consequences on community structure across coastal forest stands, and may be regulated by legacies of prior disturbances, community structure, extent of canopy damage, and species’ tolerance to specific microclimates.

     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.     

Arbuscular mycorrhizal fungi in the tree seedlings of two Australian rain forests: occurrence, colonization, and relationships with plant performance

The roots of rain forest plants are frequently colonized by arbuscular mycorrhizal fungi (AMF) that can promote plant growth in the nutrient poor soils characteristic of these forests. However, recent studies suggest that both the occurrence of AMF on rain forest plants and the dependence of rain forest plants on AMF can be highly variable. We examined the occurrence and levels of AMF colonization of some common seedling species in a tropical and a subtropical rain forest site in Queensland, Australia. We also used a long-term database to compare the growth and mortality rates of seedling species that rarely formed AMF with those that regularly formed AMF. In both forests, more than one-third of the seedling species rarely formed AMF associations, while 40% of species consistently formed AMF in the tropical site compared to 27% in the subtropical site. Consistent patterns of AMF occurrence were observed among plant families at the two sites. Variation among seedling species in AMF occurrence or colonization was not associated with differences in seed mass among species, variation in seedling size and putative age within a species, or lack of AMF inoculum in the soil. Comparisons of four seedling species growing both in the shaded understory and in small canopy gaps revealed an increase in AMF colonization in two of the four species in gaps, suggesting that light limitation partially explains the low occurrence of AMF. Seedling survival was significantly positively associated with seed biomass but not with AMF colonization. Furthermore, seedling species that regularly formed AMF and those that did not had similar rates of growth and survival, suggesting that mycorrhizal and nonmycorrhizal strategies were equivalent in these forests. Furthermore, the high numbers of seedlings that lacked AMF and the overall low rate of seedling growth (the average seedling required 6 years to double its height) suggest that most seedlings did not receive significant indirect benefits from AMF through connection to canopy trees via a common mycorrhizal network.     

光对热带雨林冠层树种绒毛番龙眼种子萌发及其幼苗早期建立的影响

绒毛番龙眼(Pometia tomentosa)为西双版纳热带季节雨林标志树种,被列为中国珍稀濒危植物。对其开展种子及幼苗生态学研究可为珍稀濒危树种的保护及当地森林恢复提供科学依据。试验采用遮荫试验和野外试验相结合的方法,研究光对绒毛番龙眼种子萌发及幼苗早期建立的影响。结果表明:在遮荫试验30%(林窗中心光)、10%(林窗边缘光)和3.5%(林下光)3个光处理及森林3种生境(林窗中心、林窗边缘和林下)条件下,种子萌发率差异无显著性差异,均达到95%以上,且平均萌发周期小于6 d。3个光处理下的幼苗生长、生物量分配模式及气体交换参数差异显著。30%光下绒毛番龙眼幼苗的根重比(RMR)和茎重比(SMR)最高,10%和3.5%光下幼苗的叶重比(LMR)最高,3.5%光下的叶面积比(LAR)显著高于30%光下。30%光下绒毛番龙眼幼苗的最大净光合速率(P_nmax)、暗呼吸速率(R_d)和光饱和点(I_sat)在3个光处理中均最大,光补偿点(I_comp)则无显著性差异。绒毛番龙眼幼苗具耐荫性,能够在低光环境下长期存活且能缓慢生长;同时具有适应林窗光环境生长的能力,如高的质量相对生长速率(RGR_M)和高度相对生长速率(RGR_H)。林窗的出现是绒毛番龙眼进行成功更新的必要条件,水分可能是限制其幼苗生境选择的另一环境因子。     

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.     

山地落叶阔叶林优势树种米心水青冈幼苗的定居

在郁闭的林冠下、模拟林窗和林间空旷地3种光照环境中,研究了施肥和未施肥的米心水青冈幼苗生存和生长发育过程．结果表明,幼苗在林下的发育受到光照不足的严重抑制,生长在林下的幼苗比生长在林窗和空旷地幼苗死亡快,死亡率高,幼苗在郁闭林下能够生存的时间不超过12周．施肥处理的幼苗比对照的幼苗死亡快,死亡率高．幼苗死亡很可能与土壤中的病原生物有关．虽然由林下光照强度增加到模拟林窗的光照强度后,幼苗发育有明显改善,但幼苗在模拟林窗环境和空旷地的生长没有显著差异．结果说明,米心水青冈新出土的幼苗在郁闭林冠下易受伤害,更新立苗阶段需要有比较好的光照条件,肥沃土壤可能也无助于改善郁闭林冠下幼苗的定居．     

The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance

Dipterocarps dominate the canopy of lowland tropical rain forest in South‐east Asia. Seedlings of these species form diverse assemblages on the forest floor where low irradiance severely limits their growth. Further growth depends largely upon the increased irradiance that can occur with the creation of canopy gaps. However, the response of dipterocarp seedlings to increased irradiance and their subsequent establishment in the canopy may be influenced by the availability of other resources, such as nutrient availability. We investigated the influence of nitrogen supply on aspects of the photosynthetic physiology and growth of seedlings of four dipterocarp species (Shorea leprosula, Shorea johorensis, Shorea oleosa and Dryobalanops lanceolata) growing under low irradiance, during transfer from low to high irradiance, and during subsequent growth at high irradiance. All four species increased growth and photosynthetic capacity in response to N‐supply at high irradiances but not at low irradiance approximating that which can be expected to occur in the forest understorey. When seedlings grown at low irradiances and varying N‐supply were exposed to a large increase in irradiance, all species showed some degree of initial photodamage (measured through chlorophyll fluorescence), the extent of which was similar between species but differed markedly depending on the pre‐exposure growth irradiance and N‐supply. Greater photodamage occurred in seedlings grown at lower compared with higher N‐supply and irradiance. Despite these initial difference in the extent of this photodamage, all seedlings demonstrated a similar capacity to recover from damage. However, the alterations in the photosynthetic physiology of leaves during this recovery differed between species and depended on N‐supply. Under high N‐supply all species apart from S. oleosa increased photosynthetic capacity per unit chlorophyll following exposure to high irradiance by increasing photosynthetic capacity per unit leaf area while, under low N‐supply, an increase in photosynthetic capacity per unit leaf only occurred in D. lanceolata. Our results suggest that variations in N‐availability may have a much greater impact on the relative competitiveness of dipterocarp seedlings during the regenerative phase following canopy gap formation than physiological differences between seedlings. Our results demonstrate a potentially significant role for N‐availability in the regeneration dynamics and distribution of canopy‐dominating dipterocarp species.     

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.     

Characterizing Tropical Tree Species Growth Strategies: Learning from Inter-Individual Variability and Scale Invariance

Understanding how tropical tree species differ in their growth strategies is critical to predict forest dynamics and assess species coexistence. Although tree growth is highly variable in tropical forests, species maximum growth is often considered as a major axis synthesizing species strategies, with fast-growing pioneer and slow-growing shade tolerant species as emblematic representatives. We used a hierarchical linear mixed model and 21-years long tree diameter increment series in a monsoon forest of the Western Ghats, India, to characterize species growth strategies and question whether maximum growth summarizes these strategies. We quantified both species responses to biotic and abiotic factors and individual tree effects unexplained by these factors. Growth responses to competition and tree size appeared highly variable among species which led to reversals in performance ranking along those two gradients. However, species-specific responses largely overlapped due to large unexplained variability resulting mostly from inter-individual growth differences consistent over time. On average one-third of the variability captured by our model was explained by covariates. This emphasizes the high dimensionality of the tree growth process, i.e. the fact that trees differ in many dimensions (genetics, life history) influencing their growth response to environmental gradients, some being unmeasured or unmeasurable. In addition, intraspecific variability increased as a power function of species maximum growth partly as a result of higher absolute responses of fast-growing species to competition and tree size. However, covariates explained on average the same proportion of intraspecific variability for slow- and fast-growing species, which showed the same range of relative responses to competition and tree size. These results reflect a scale invariance of the growth process, underlining that slow- and fast-growing species exhibit the same range of growth strategies.     

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.     

Cold tolerance of photosynthesis as a determinant of tree species regeneration patterns in an evergreen temperate forest

Niche partitioning of light among seedling species is a key mechanism supporting coexistence in forests. Species sort along light gradients through direct responses to light and through indirect responses mediated by other environmental factors. Canopy gaps in temperate evergreen rainforests experience sub-zero temperatures and thus gap-dependent species are vulnerable to cold photoinhibition from exposure to high light at low temperatures. We used a shadehouse experiment to test two hypotheses: (1) that gap-dependent species are resistant to cold photoinhibition; and (2) that gap-dependence observed in the field may be driven by the interaction between high light and low temperatures. Specifically, we predicted that some species restricted to shade in the field are excluded from gaps because of low resistance to cold photoinhibition. Gap dependence of angiosperm and conifer seedlings was estimated from expert opinion, and from experimental growth and survival responses to light treatments representing a forest understorey and forest gap. Additional seedlings were used to evaluate resistance to cold photoinhibition (sub-zero temperatures at dawn). Gap-dependent species were resistant to cold photoinhibition. Our second hypothesis was supported by Beilschmiedia tawa (Lauraceae), which had low resistance to photoinhibition, a strong positive growth response to the light treatments, and is restricted to shade in the field. Seedling regeneration niches in temperate rainforest are shaped in part by the interaction between light and low temperatures, and this interaction will be crucial for determining seedling responses to climate warming.     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

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。     

Photosynthetic responses of tree seedlings in grass and under shrubs in early-successional tropical old fields, Costa Rica

Only recently have studies addressed the effect of early-colonizing vegetation on tree seedling survival and growth during secondary succession in tropical old fields, and few studies have elucidated the physiological responses of tree seedlings to different vegetational communities. We compared growth and various photosynthetic parameters for seedlings of four rain-forest tree species, Cedrela tonduzii, Inga punctata, Ocotea whitei, and Tapirira mexicana, growing in areas of pasture grass and shrubs in early-successional abandoned pasture in Costa Rica; in addition, we made measurements for two species in forest gaps. We tested the general hypothesis that early-colonizing shrubs facilitate growth of forest tree seedlings. Specifically, we measured microclimate, growth, CO₂ assimilation, stomatal conductance, photosystem II quantum yield (KPSII), and xanthophyll pigment pools for all seedlings. Photosynthetic flux density (PFD) was higher under grass than shrubs or forest gaps, but was highly variable in each growth environment. For three of the four species, height growth was greatest in the grass compared to the shrubs and forest gaps; growth was similar below grass and shrubs for O. whitei. Photosynthetic capacity, apparent quantum yield, and stomatal conductance did not vary across habitats, but light compensation point and PFD at light saturation tended to be higher in the grass compared to forest and shrub growth environments. Water use efficiency differed across growth environments for three of the species. For plants in ambient PFD and dark-adapted plants, the efficiency of excitation energy transfer through PSII was lowest for plants in the grass compared to shrubs and forest gaps and also differed across species. Measurement of steady-state responses of KPSII to increasing PFD indicated a significant effect of growth environment at low PFD for all species and significant effects at high PFD only for I. punctata. All species exhibited a high degree of midday xanthophyll de-epoxidation in the different growth environments. Xanthophyll pigment pool size on an area basis was highest in the grass compared to shrubs and forest gaps for all four species. The results suggest that shrubs do not provide a facilitative effect for growth or photosynthesis for ~1.5-year-old seedlings of these four species. We conclude that site differences in success of tree seedlings during succession are a result of complex interactions of facilitation and competition and are not simply based on physiological responses to PFD.