Contrasting species responses to continued nitrogen and phosphorus addition in tropical montane forest tree seedlings

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient‐limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth‐limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species‐specific and that successful adaptation to increased nutrient availability will depend on species‐specific morphological and physiological plasticity.     

Topographic and spatial controls of palm species distributions in a montane rain forest, southern Ecuador

The northern Andes harbour a flora that is as species-rich or even richer than the 18-times larger lowland Amazon basin. Gaining an understanding of how the high species richness of the Andean region is generated and maintained is therefore of particular interest. Environmental sorting due to elevational gradients in climate has been emphasized as a driver of vegetation distribution and plant community assembly in tropical mountain areas such as the Andes for two centuries, while alternative mechanisms have been little studied. Here, we investigated the importance of topography and spatial location as factors controlling species distributions in a palm community in a montane rain forest landscape in the Andes of southern Ecuador (1900–2150 m above sea level). Eleven species were present: Aiphanes verrucosa, Ceroxylon parvifrons, Chamaedorea pinnatifrons, Dictyocaryum lamarckianum, Euterpe precatoria, Geonoma densa, Geonoma orbignyana, Geonoma paradoxa, Prestoea acuminata and Wettinia aequatorialis. To study their spatial distribution, forty 250 m² (5 × 50 m²) plots were laid out perpendicular to four paths that were categorized into three areas and two topographic units (ridges and gullies). Mantel tests and indicator species analysis showed that both topography and spatial location imposed strong controls on palm species distributions at the study site. Our results suggest that species distributions in the studied montane forest landscape were partly determined by the species’ habitat requirements, but also by unknown spatial effects. Although a number of possible explanations exist for the latter, such as unmeasured environmental variables and historical disturbance events, we believe dispersal limitation is likely to be involved. Furthermore, although the gully- or ridge-association of some species corresponded to their general elevational ranges in southern Ecuador, this was not the case for other species. Based on such considerations, we conclude that elevational climatic gradients are likely to only form part of the explanation for the topographic effects on palm species distributions at the study site. Other factors must also be involved, notably wind-exposure and hydrology, as discussed for lowland palm communities. Our results show that to understand plant community assembly in the tropical montane forests of the Andes it is too simple to focus just on environmental sorting by elevational climatic gradients.     

Responses of legume versus nonlegume tropical tree seedlings to elevated CO2 concentration

We investigated responses of growth, leaf gas exchange, carbon-isotope discrimination, and whole-plant water-use efficiency (W(P)) to elevated CO(2) concentration ([CO(2)]) in seedlings of five leguminous and five nonleguminous tropical tree species. Plants were grown at CO(2) partial pressures of 40 and 70 Pa. As a group, legumes did not differ from nonlegumes in growth response to elevated [CO(2)]. The mean ratio of final plant dry mass at elevated to ambient [CO(2)] (M(E)/M(A)) was 1.32 and 1.24 for legumes and nonlegumes, respectively. However, there was large variation in M(E)/M(A) among legume species (0.92-2.35), whereas nonlegumes varied much less (1.21-1.29). Variation among legume species in M(E)/M(A) was closely correlated with their capacity for nodule formation, as expressed by nodule mass ratio, the dry mass of nodules for a given plant dry mass. W(P) increased markedly in response to elevated [CO(2)] in all species. The ratio of intercellular to ambient CO(2) partial pressures during photosynthesis remained approximately constant at ambient and elevated [CO(2)], as did carbon isotope discrimination, suggesting that W(P) should increase proportionally for a given increase in atmospheric [CO(2)]. These results suggest that tree legumes with a strong capacity for nodule formation could have a competitive advantage in tropical forests as atmospheric [CO(2)] rises and that the water-use efficiency of tropical tree species will increase under elevated [CO(2)].     

Nutrients limit photosynthesis in seedlings of a lowland tropical forest tree species

We investigated how photosynthesis by understory seedlings of the lowland tropical tree species Alseis blackiana responded to 10 years of soil nutrient fertilization with N, P and K. We ask whether nutrients are limiting to light and CO₂ acquisition in a low light understory environment. We measured foliar nutrient concentrations of N, P and K, isotopic composition of carbon (δ¹³C) and nitrogen (δ¹⁵N), and light response curves of photosynthesis and chlorophyll fluorescence. Canopy openness was measured above each study seedling and included in statistical analyses to account for variation in light availability. Foliar N concentration increased by 20% with N addition. Foliar P concentration increased by 78% with P addition and decreased by 14% with N addition. Foliar K increased by 8% with K addition. Foliar δ¹³C showed no significant responses, and foliar δ¹⁵N decreased strongly with N addition, matching the low δ¹⁵N values of applied fertilizer. Canopy openness ranged from 0.01 to 6.71% with a mean of 1.76 ± 0.14 (±1SE). Maximum photosynthetic CO₂ assimilation rate increased by 9% with N addition. Stomatal conductance increased with P addition and with P and K in combination. Chlorophyll fluorescence measurements revealed that quantum yield of photosystem II increased with K addition, maximum electron transport rate trended 9% greater with N addition (p = 0.07), and saturating photosynthetically active radiation increased with N addition. The results demonstrate that nutrient addition can enhance photosynthetic processes, even under low light availability.     

Water relations of indigenous versus exotic tree species,growing at the same site in a tropical montane forest in southern Ethiopia

The objective of the study was to compare the water relations of two indigenous [Podocarpus falcatus (Thunb.) Endl., Croton macrostachys Hochst. ex. Del.] and two exotic tree species (Eucalyptus globulus Labille., Cupressus lusitanica Miller) growing in the same location in the montane Munessa State Forest, southern Ethiopia. Stem flow was measured with Granier type thermal dissipation probes. Sap flux, normalized per unit sapwood area, and the total sapwood areas of the particular trees were used to estimate daily transpiration. Maximum daily transpiration values (60 kg water) were recorded for Croton when at full foliage. After shedding most of its leaves in the dry season transpiration was reduced to 8 kg per day. Eucalyptus had the next highest transpiration (55 kg), in this case at the peak of the dry season. It transpired 4–5 times more than Podocarpus and Cupressus trees of similar size. Maximum stem flux density was tree-size dependent only in Croton. Diurnal patterns of stem flux indicated that Croton, Eucalyptus and Podocarpus, in contrast to Cupressus, responded more directly to light than to atmospheric water pressure deficit. At high VPD (>1.0 kPa) stem flux reached a plateau in Croton and Podocarpus indicating stomatal limitation. Per unit leaf area Croton had the highest and Podocarpus and Cupressus the lowest daily transpiration rates. In summary, the pioneer tree Croton had the lowest and Podocarpus the highest water use efficiency. The contribution of the study to the understanding of the role of each tree species in the hydrology of the natural forest and the plantations is discussed.     

Vegetation response to removal of non-native feral pigs from Hawaiian tropical montane wet forest

Globally, non-native ungulates threaten native biodiversity, alter biotic and abiotic factors regulating ecological processes, and incur significant economic costs via herbivory, rooting, and trampling. Removal of non-native ungulates is an increasingly common and crucial first step in conserving and restoring native forests. However, removal is often controversial and there is currently little information on plant community responses to this management action. Here, we examine the response of native and non-native understory vegetation in paired sites inside and outside of exclosures across a 6.5–18.5 year chronosequence of feral pig (Sus scrofa) removal from canopy-intact Hawaiian tropical montane wet forest. Stem density and cover of native plants, species richness of ground-rooted native woody plants, and abundance of native plants of conservation interest were all significantly higher where feral pigs had been removed. Similarly, the area of exposed soil was substantially lower and cover of litter and bryophytes was greater with feral pig removal. Spatial patterns of recruitment were also strongly affected. Whereas epiphytic establishment was similar between treatments, the density of ground-rooted woody plants was four times higher with feral pig removal. Abundance of invasive non-native plants also increased at sites where they had established prior to feral pig removal. We found no patterns in any of the measured variables with time, suggesting that commonly occurring species recover within 6.5 years of feral pig removal. Recovery of species of conservation interest, however, was highly site specific and limited to areas that possessed remnant populations at the time of removal, indicating that some species take much longer (>18.5 years) to recover. Feral pig removal is the first and most crucial step for conservation of native forests in this area, but subsequent management should also include control of non-native invasive plants and outplanting native species of conservation interest that fail to recruit naturally.     

Responses of communities of tropical tree species to elevated CO2 in a forest clearing

Communities of ten species of tropical forest tree seedlings from three successional classes were grown at ambient and elevated CO₂ in large open-top chambers on the edge of a forest in Panamá. Communities grew from 20?cm to approximately 2?m in height in 6 months. No enhancements in plant biomass accumulation occurred under elevated CO₂ either in the whole communities or in growth of individual species. Reductions in leaf area index under elevated CO₂ were observed, as were decreases in leaf nitrogen concentrations and increases in the C:N ratio of leaf tissue. Species tended to respond individualistically to elevated CO₂, but some generalizations of how successional groupings responded could be made. Early and mid-successional species generally showed greater responses to elevated CO₂ than late-successional species, particularly with respect to increases in photosynthetic rates and leaf starch concentrations, and reductions in leaf area ratio. Late-successional species showed greater increases in C:N ratios in response to elevated CO₂ than did other species. Our results indicate that there may not be an increase in the growth of regenerating tropical forest under elevated CO₂, but that there could be changes in soil nutrient availability because of reductions in leaf tissue quality, particularly in late-successional species.     

Diverging temperature response of tree stem CO2 release under dry and wet season conditions in a tropical montane moist forest

It is commonly presumed that plant respiratory CO₂ release increases with increasing temperature. However, we report on very contrasting stem CO₂ release (R _S)–temperature relationships of trees in a species-rich tropical montane forest of southern Ecuador under dry and wet season conditions. Rates of R _S were low and completely uncoupled from the dial temperature regime during the humid season. In contrast, during the dry season, R _S was generally higher and temperature sensitivity of R _S differed greatly in degree and even in the direction of response, indicating that temperature might not be the only determinant of R _S. In order to explain the heterogeneity of R _S, we related R _S to vapour pressure deficit, wind speed and solar radiation as important abiotic drivers influencing transpiration and photosynthesis. Stepwise multiple regression analyses with these meteorological predictors either were biased by high collinearity of the independent variables or could not enhance the ability to explain the variability of R _S. We assume maintenance respiration to dominate under humid conditions unfavourable for energy acquisition of the tree, thus explaining the pronounced uncoupling of R _S from atmospheric parameters. In contrast, the drier and hotter climate of the dry season seems to favour R _S via enhanced assimilatory substrate delivery and stem respiratory activity as well as elevated xylem sap CO₂ imports with increased transpiration. In addition, tree individual differences in the temperature responses of R _S may mirror diverging climatic adaptations of co-existing moist forest tree species which have their distribution centre either at higher or lower elevations.     

Growth of tree seedlings in a tropical dry forest in relation to soil moisture and leaf traits

"AimsThe growth of plant species in tropical dry forest (TDF) is expected to be largely governed by the availability of soil moisture. In this study we attempt to identify mechanisms by which seedlings of dry tropical trees cope with water stress by adjusting their leaf characteristics to water availability and micro environments, and address following questions: How are leaf traits and relative growth rate (RGR) of the dominant seedling species of TDF affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional traits with each other? Can leaf traits singly or in combination predict the growth rate of seedling species of TDF? The study was conducted in situ on four sites (viz., Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India. Methods Five leaf traits viz., specific leaf area (SLA), leaf dry matter content (LDMC), concentrations of leaf nitrogen (leaf N), phosphorus (leaf P) and chlorophyll (Chl) and two physiological processes, viz., stomatal conductance (Gs net) and photosynthetic rate (A net), and RGR, of four dominant tree seedling species of a TDF (viz., Buchanania lanzan, Diospyros melanoxylon, Shorea robusta and Terminalia tomentosa) on four sites were analysed for species, site and season effects over a 2-year period. Step-wise multiple regression was performed to predict RGR from mean values of SMC, leaf traits and physiological processes. Principal component analysis (PCA) was performed to observe the extent of intra- vs. inter-specific variability in the leaf traits and physiological rates.Important findings All the traits and physiological rates were interrelated and showed significant positive relationship with RGR except for the correlation of LDMC with RGR which was not significant. Further, relationships of SMC with all leaf traits, physiological rates and RGR were significant, except for that between SMC and SLA for B. lanzan and D. melanoxylon. The slope of seedling trait:SMC relationship, a measure of phenotypic plasticity in response to soil moisture gradient, varied among species. Among the four species, T. tomentosa was the most plastic and S. robusta the least. In conclusion, leaf traits and physiological processes were strongly related to soil water availability on the one hand and seedling growth on the other. Gs net is the most important variable which accounted for the greatest amount of variability (62%) in RGR, emphasizing the role of stomatal conductance in shaping growth patterns across spatial and temporal gradients of soil water availability. Gs net and SMC together explained 64% variability in RGR, indicating that other traits/factors, not studied by us are also important in modulating the growth of tropical tree seedlings.     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

海南霸王岭山地原始林与伐后林中木质藤本对支持木的选择

通过对海南霸王岭热带山地原始林与伐后林中树木及其攀附木质藤本的调查,研究原始林与伐后林中木质藤本对支持木的选择性。结果表明:1)6科优势树木中附藤率最高的是野牡丹科(Melastomataceae),附藤率最低的科,原始林中是山矾科(Symplocaceae),伐后林中是茜草科(Rubiaceae)。2)原始林中,谷木(Memecylon ligustrifolium)与线枝蒲桃(Syzygiumaraiocladum)的附藤比率和每木藤本数均高于样地平均水平;三角瓣花(Prismatomeris tetrandra)和龟背灰木(Symplocosandenophylla)的附藤比率均低于样地平均水平,而每木藤本数与样地平均水平之间没有显著差异。伐后林中,谷木的附藤比率和每木藤本数高于样地平均水平;九节(Psychotria rubra)的附藤比率和每木藤本数低于样地平均水平。3)杜仲藤(Parabariummicranthum)的主要支持木是谷木,夜花藤(Hypserpa nitida)的主要支持木是线枝蒲桃。研究表明,木质藤本对支持木在科和种水平上都具有选择性,因此木质藤本会对树木造成不对称影响,进而影响森林的结构和动态。     

A comparison of alpha and beta diversity patterns of ferns,bryophytes and macrolichens in tropical montane forests of southern Ecuador

We present a first comparison of patterns of alpha and beta diversity of ferns, mosses, liverworts and macrolichens in neotropical montane rainforests, and explore the question whether specific taxa may be used as surrogates for others. In three localities in southern Ecuador, we surveyed terrestrial and epiphytic species assemblages in ridge and slope forests in 28 plots of 400 m2 each. The epiphytic habitat was significantly richer in ferns, liverworts, and macrolichens than the terrestrial habitat; mosses, however, were primarily terrestrial. Alpha diversity of ferns and of liverworts was congruent in both habitats. Mosses were similar to ferns and liverworts only in the epiphytic habitat. Macrolichens did not share patterns of alpha diversity with any other group. Beta diversity of ferns, mosses and liverworts (lichens excluded due to low species richness) was similar in the terrestrial habitat, but not in the epiphytic habitat. Our results demonstrate that patterns of alpha diversity of the studied taxa cannot be used to predict patterns of beta diversity. Moreover, diversity patterns observed in epiphytes are different from terrestrial plants. We noted a general coincidence in species patterns of liverworts and ferns. Diversity patterns of macrolichens, in contrast, were completely independent from any other taxonomic group studied.     

高CO2浓度对温带三种针叶树光合光响应特性的影响

将长白山地区阔叶红松林中主要针叶树种红松、红皮云杉和长白落叶松的幼苗 ,盆栽于模拟自然光照和人工调节CO2 浓度为 70 0和 40 0 μmol·mol-1的气室内两个生长季 ,在各自的生长环境条件下 ,利用CI 30 1PS便携式CO2 分析系统测定针叶的光合光响应曲线 .结果表明 ,不同树种及同一树种的不同CO2浓度处理间差异明显 .比较饱和净光合速率、暗呼吸、光补偿点、光饱和点、及光能利用率 (QUE)的变化可见 ,长白落叶松为阳性树种 ,其光合作用对高CO2 浓度的适应能力较好 ,红松树种次之 ,阴性树种红皮云杉光合作用对高CO2 浓度适应能力最差 .并初步探讨了供试树种光合生理特性及其演替状况间的联系     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Fern richness,tree species surrogacy,and fragment complementarity in a Mexican tropical montane cloud forest

We related pteridophytes versus tree species composition to identify surrogate measures of diversity, and complementarity of seven cloud forest fragments. Forest structure, and fern and tree composition were determined in 70 (2 × 50 m) transects. Fern density (10,150–25,080 individuals/ha) differed among sites. We recorded 83 fern species in the transects. Nonparametric richness estimators indicated that more sampling effort was needed to complete fern inventories (14 more species). However, ferns recorded outside of the transects increased richness to 103 species (six more species than predicted). Twenty-eight species were unique and rare due to special habitat requirements (Diplazium expansum, Hymenophyllum hirsutum, Melpomene leptostoma, Terpsichore asplenifolia), or were at a geographical distribution edge (Diplazium plantaginifolium, Lycopodium thyoides, Pecluma consimilis, Polypodium puberulum). Correlations between fern richness and tree richness and density were not significant, but were significant between fern richness and fern density, between epiphytic fern density and tree richness and density. Tree richness is not a good surrogate for fern diversity. Only three species were recorded in all fragments (Polypodium lepidotrichum, P. longepinnulatum, P. plebeium); thus fragments pteridophytes compositions are highly complementary, but more similar for ferns than for trees. A regional conservation approach which includes many small reserves needs to focus supplementarity on patterns of tree and fern species richness.