Leaf demography and leaf traits of temperate-deciduous and tropical evergreen-broadleaved trees in a Mexican montane cloud forest

The spatial variation in epilithic lichen community structure was investigated as part of a larger study of the vegetation and ecology of the tall limestone cliffs within the Niagara Escarpment Biosphere Reserve in southern Ontario, Canada. The cover of all lichen taxa was visually estimated for a total of 199 samples taken from the top, middle, or bottom of the cliff face at five sites. Twelve environmental variables were also measured. Twenty-seven lichen taxa were identified on the samples. Multivariate ordinations of species composition (DCA, CCA, PCCA) revealed variation in community structure on multiple scales, but no groupings of sites that would have suggested the presence of several distinct species assemblages. A gradient in species composition from north to south, most clearly reflected in the decreasing cover of foliose and fruticose species, may reflect a gradient in human disturbance. There was also intermediate-scale patchiness in species composition in a horizontal plane across cliffs, but despite earlier claims made in the literature, no evidence of vertical zonation of the lichens on cliffs was found. Species composition also responded to small-scale factors possibly related to exposure, light, or moisture. Unlike community composition, the total cover of all lichens was homogeneous over large spatial scales and varied only on a small scale, illustrating that scale as well as resolution of a study may influence the ecological patterns seen. More than half of the species found on the Niagara Escarpment are rare on rock substrates elsewhere in southern Ontario, and two are new for North America (Candelariella heidelbergensis (Nyl.) Poelt and Lecanora perpruinosa Fröberg). The result that cliffs support a distinct flora containing many rare species suggests that they are a reservoir for biodiversity not just for vascular plants, but also for lichens.     

Breeding systems in a montane tropical cloud forest in Venezuela

The breeding systems, reproductive efficacies and population densities of 75 species of trees, shrubs, perennial herbs and vines in a montane tropical cloud forest in Venezuela are investigated. 56.96% and 44.32% of the trees, versus the other life forms considered, respectively possess obligate outbreeding mechanisms. Two shrubs are non-pseudogamous apomicts. The percentage of dioecy among tree species (31%) is among the highest recorded in tropical forests. Reproductive efficacy is similar under all breeding systems in the forest interior. Obligately outbred taxa are slower recolonizers of a disturbed border as compared with non-obligate outbreeders. Explanations are advanced for the high incidence of dioecy combined with a low level of self-incompatibility among hermaphroditic species. It is concluded that the breeding system spectrum reflects an unpredictable pollination pattern, rather than insufficient pollinator servicing.     

Foliage dynamics, leaf traits, and growth of coexisting evergreen and deciduous trees in a tropical montane forest in Ethiopia

Foliage dynamics of three functional tree types representing major components of the tropical montane evergreen forest in southern part of Central Ethiopia were compared. The species were Podocarpus falcatus (evergreen gymnosperm), Prunus africana (evergreen broadleaf), and Croton macrostachyus (facultative deciduous). The hypothesis examined is that in such tropical trees, endogenous control of foliage dynamics by the leaf life-spans (LLS) is largely dominant over external signals. Crown foliage turnover, leafiness of twigs, LLS, photosynthetic performance, respiration rate, specific leaf area, and relative growth rates of the stems were investigated. Foliage dynamics and leafiness of the twigs were monitored over 2?years while leaf traits were followed over 3?months. The degree of inter and intra-individual synchronization of foliage phenophases was examined to get an estimate of the contributions of endogenous and external signals to the dynamics of the foliages. Autoregression analysis indicated significant influence of the moisture regime on leaf sprouting of Croton and Podocarpus. During pronounced dry periods, new leaves were not developed. Analysis of phenological data using circular statistics revealed that in spite of strong inter-individual synchronization of leaf flush and fall (Podocarpus and Croton), the dynamics of individual parts of the crowns were less synchronized. LLS was independent of climate factors and it had substantial contribution to the control of foliage turnover. Moreover, examination of ecophysiological traits of developing leaves of the studied functional types showed differing patterns with LLS corroborating the ecophysiological characteristics. Although overlaid by fungal infestation, both the foliage and ecophysiological properties of Prunus resemble that of Podocarpus but the former exhibited a shorter LLS and slightly higher metabolic rates. Nevertheless, all species reacted positively to high moisture with respect to stem growth. In spite of largely differing weather conditions of the 2?years, direct competitive advantage of one of the species over the others could not be detected.     

尖峰岭热带山地雨林林冠层乔木某些功能性状的系统发育信号、关联性及其演化模式

热带雨林林冠层具有丰富的物种、功能和系统发育(谱系)多样性,能够显著影响生态系统生物地球化学循环,调节大气水热平衡,缓解气候变化与人为干扰的负面作用。因此热带雨林林冠层功能和谱系生态学研究是目前群落生态学研究的热点之一。选取海南尖峰岭热带山地雨林3个1 hm2样地的林冠层为研究对象,利用样地林冠树种清查数据基于APG III系统发育结构重建样地林冠层系统发育树。利用Blomber's K对和Pagel'sλ指标结合系统发育独立性比较法对组成林冠层乔木树种的结构、化学计量和水力学功能性状(11个性状)的系统发育信号、关联性和演化模式进行研究,以探讨物种亲缘关系对尖峰岭热带山地雨林林冠层多维功能性状关联性及其动态演化的影响。结果表明,有8个功能性状(叶面积、叶厚度和潜在最大高度除外)具有显著的系统发育信号(通过P0.05的显著性检验),Blomber's K值和Pagel'sλ值分别介于0.202—0.392和0.277—0.847之间,表明尖峰岭热带山地雨林林冠层功能性状普遍存在系统发育保守性。在系统发育背景下,林冠层乔木树种结构、化学和水力学功能性状在物种水平普遍存在显著关联性(P0.05),表现出趋同或趋异进化;而且林冠层乔木随物种分化其功能性状分化模式大致呈水平"漏斗"状,姐妹类群功能性状间差异性在物种分化早期(大约1亿2千万年前至6000万年前)明显小于其在中后期(大约6000万年前至今)的差异性,导致了林冠层性状空间在中后期迅速膨大。然而林冠层主要功能性状在系统发育树内部每一节点上姊妹类群分化产生的系统发育独立性比较值绝大部分与零模型随机模拟值并无显著性差异。了解系统发育背景下林冠层功能性状的权衡关系及其随物种演化的分化模式与时间动态为进一步探究热带雨林林冠生态系统功能发挥奠定基础。     

Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Seasonal changes in stem diameter and leaf development in a tropical montane forest

Abstract. Seasonal patterns of stem diameter changes in evergreen and deciduous species of a tropical montane forest in the Central Himalayas (300–2250 m a.s.l.) were investigated in relation to leaf development. Ca. 75 % of the annual rainfall in this region occurs in a short period, from mid-June to mid-September and the remaining months are dry. It was assumed that changes in stem diameter are correlated with changes in water stress. Each evergreen species could be characterized by leaf longevity of about one year; each species showed pronounced summer leaf drop and simultaneous new leaf formation. Winter stem shrinkage was more pronounced in deciduous species than in evergreen ones. The deciduous species also showed a greater proportional loss of leaf mass (before abscission) than the evergreen species. Winter leaf fall in deciduous species was related to the pronounced stem shrinkage. The leaf fall enabled these species to control further water loss. Being more resistant to desiccation, the evergreen species retained their leaves throughout the winter but showed gradual loss of leaf mass, presumably in order to control water loss. In all species, leaf expansion was completed before the onset of the rainy season, when water stress was high. This strategy has definite advantages in a climate with a monsoon pattern of rainfall. Evergreen species, showing pronounced leaf drop in summer, have advantages over deciduous species; hence their preponderance in the region.     

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.     

Coordinated changes in photosynthesis,water relations and leaf nutritional traits of canopy trees along a precipitation gradient in lowland tropical forest

We investigated leaf physiological traits of dominant canopy trees in four lowland Panamanian forests with contrasting mean annual precipitation (1,800, 2,300, 3,100 and 3,500 mm). There was near complete turn-over of dominant canopy tree species among sites, resulting in greater dominance of evergreen species with long-lived leaves as precipitation increased. Mean structural and physiological traits changed along this gradient as predicted by cost–benefit theories of leaf life span. Nitrogen content per unit mass (N_mass) and light- and CO₂-saturated photosynthetic rates per unit mass (P_mass) of upper canopy leaves decreased with annual precipitation, and these changes were partially explained by increasing leaf thickness and decreasing specific leaf area (SLA). Comparison of 1,800 mm and 3,100 mm sites, where canopy access was available through the use of construction cranes, revealed an association among extended leaf longevity, greater structural defense, higher midday leaf water potential, and lower P_mass, N_mass, and SLA at wetter sites. Shorter leaf life spans and more enriched foliar ¹⁵N values in drier sites suggest greater resorption and re-metabolism of leaf N in drier forest. Greater dominance of short-lived leaves with relatively high P_mass in drier sites reflects a strategy to maximize photosynthesis when water is available and to minimize water loss and respiration costs during rainless periods. Overall, our study links coordinated change in leaf functional traits that affect productivity and nutrient cycling to seasonality in lowland tropical forests.     

Thermal acclimation of leaf respiration of tropical trees and lianas: response to experimental canopy warming,and consequences for tropical forest carbon balance

Climate warming is expected to increase respiration rates of tropical forest trees and lianas, which may negatively affect the carbon balance of tropical forests. Thermal acclimation could mitigate the expected respiration increase, but the thermal acclimation potential of tropical forests remains largely unknown. In a tropical forest in Panama, we experimentally increased nighttime temperatures of upper canopy leaves of three tree and two liana species by on average 3  ° C for 1 week, and quantified temperature responses of leaf dark respiration. Respiration at 25  ° C (R₂₅) decreased with increasing leaf temperature, but acclimation did not result in perfect homeostasis of respiration across temperatures. In contrast, Q₁₀ of treatment and control leaves exhibited similarly high values (range 2.5–3.0) without evidence of acclimation. The decrease in R₂₅ was not caused by respiratory substrate depletion, as warming did not reduce leaf carbohydrate concentration. To evaluate the wider implications of our experimental results, we simulated the carbon cycle of tropical latitudes (24 ° S–24 ° N) from 2000 to 2100 using a dynamic global vegetation model (LM3VN) modified to account for acclimation. Acclimation reduced the degree to which respiration increases with climate warming in the model relative to a no‐acclimation scenario, leading to 21% greater increase in net primary productivity and 18% greater increase in biomass carbon storage over the 21st century. We conclude that leaf respiration of tropical forest plants can acclimate to nighttime warming, thereby reducing the magnitude of the positive feedback between climate change and the carbon cycle.     

Understory spider diversity in two remnants of tropical montane cloud forest in Chiapas, Mexico

We evaluated the spider diversity of a tropical montane cloud forest understory in two nearby sites with different degree of human disturbance at the Biosphere Reserve Volcán Tacaná, Chiapas, Mexico. The study was conducted over a 24 days period distributed in 6 months in 2009, covering dry and rainy seasons. A total of 8,370 spiders (1,208 adults and 7,162 juveniles) were collected. Determined specimens (7,747) represented 112 species and morphospecies, 71 genera and 22 families. The results showed that human disturbance has an influence on spider communities: species richness was significantly higher in the preserved site as regards to the disturbed site. Despite their proximity, the composition of spider communities showed only a moderate similarity between the two sites. No differences in abundance were found among sites when considering the whole sample, but sites differed clearly when seasons were analyzed separately. The rainy season had a negative effect on the abundance of spiders in the preserved site. Although the spider community structure was very similar between sites, there was a trend towards a greater species evenness in the preserved site for the whole sampling period and for the dry season.     

Habitat associations with topography and canopy structure of tree species in a tropical montane forest on Mount Kinabalu,Borneo

Habitat associations with topography and canopy structure of 42 abundant tree species were studied in a 2.74-ha plot of tropical montane forest on Mount Kinabalu, Borneo. Many of these species belong to the same higher taxa including eight families and four genera. Analysis of intraspecific spatial distributions for stems ≥ 10 cm diameter revealed that 28 species (including all six species of Fagaceae) showed aggregated distributions at the 100-m² and/or 400-m² scales, and that 20 species showed habitat associations with topography by torus-translation tests; 17 species showed both characteristics. Species' associations with the local canopy structure were characterized by crown position index (CPI), which was defined relative to neighbour trees. The CPI differed greatly among individual stems at 10–40 cm diameter, and 19 species showed significantly different frequencies of crowns exposed vertically versus those shaded beneath the canopy. Mean growth rates at 10–40 cm diameter and size distributions of species were not related to topographic associations, but were explained by the associations with canopy structure; species with more exposed crowns grew faster and had less positively skewed distributions. Diversity in habitat associations was manifest between two genera (Syzygium and Tristaniopsis) in the family Myrtaceae and among species in these genera, but was less evident in other families and two genera (Garcinia and Lithocarpus). This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

Root biomass distribution in a moist tropical montane forest

Effects of N sources (ammonium, nitrate and ammonitrate) and VA mycorrhizae (Glomus intraradices) on rhizosphere soil characteristics (pH, exchangeable acidity, exchangeable cations, inorganic N concentrations) growth and nutrient acquisition of coffee seedlings (Coffea arabica L. cv guatemala) were investigated in a pot study with an acid soil (Red Bluff Loam) sterilized by autoclaving. Ammonium addition decreased rhizosphere pH while nitrate and ammonitrate additions both increased rhizosphere pH. Mycorrhizae induced a higher pH, a lower exchangeable acidity and higher values of exchangeable cations in the rhizosphere. Ammonium addition resulted in a lower mycorrhizal infection than the two other N sources. Mycorrhizal plants grew better and accumulated more N, Ca and Mg than non-mycorrhizal plants.     

Nutrient limitation restricts growth and reproductive output in a tropical montane cloud forest bromeliad: findings from a long-term forest fertilization experiment

From studies in seasonal lowland tropical forests, bromeliad epiphytes appear to be limited mainly by water, and to a lesser extent by nutrient supply, especially phosphorous. Less is understood about the mineral nutrition of tropical montane cloud forest (TMCF) epiphytes, even though their highest diversity is in this habitat. Nutrient limitation is known to be a key factor restricting forest productivity in TMCF, and if epiphytes are nutritionally linked to their host trees, as has been suggested, we would expect that they are also nutrient limited. We studied the effect of a higher nutrient input on reproduction and growth of the tank bromeliad Werauhia sintenisii in experimental plots located in a TMCF in Puerto Rico, where all macro- and micronutrients had been added quarterly starting in 1989 and continuing throughout the duration of this study. We found that bromeliads growing in fertilized plots were receiving litterfall with higher concentrations of N, P, and Zn and had higher concentrations of P, Zn, Fe, Al, and Na in their vegetative body. The N:P ratios found (fertilized = 27.5 and non-fertilized = 33.8) suggest that W. sintenisii may also be phosphorous limited as are lowland epiphytes. Fertilized plants had slightly longer inflorescences, and more flowers per inflorescence, than non-fertilized plants, but their flowers produced nectar in similar concentrations and quantities. Fertilized plants produced more seeds per fruit and per plant. Frequency of flowering in two consecutive years was higher for fertilized plants than for controls, suggesting that fertilized plants overcome the cost of reproduction more readily than non-fertilized plants. These results provide evidence that TMCF epiphytic bromeliads are nutrient limited like their lowland counterparts.     

Interactions between epiphytes during canopy soil formation: an experiment in a lower montane cloud forest of southeast Mexico

• In several montane forests around the world, epiphytes coexist in mats, sharing the rhizosphere and forming histosol-type soils rich in nutrients. The role of these epiphytes in the formation of canopy soil and the fitness costs that epiphytes face when cohabiting in these mats are unknown.
• In a lower montane cloud forest in central Veracruz, Mexico, a 2-year factorial experiment was carried out with the presence/absence of ramets of Phlebodium areolatum (Polypodiaceae), Tillandsia kirchhoffiana, T. multicaulis and T. punctulata (Bromeliaceae). We examined (i) which epiphyte species contribute to the formation of canopy soil, (ii) the role of epiphyte composition in the soil nutrient composition, and (iii) the fitness costs faced by epiphytes when cohabiting.
• Canopy soil formation highest when P. areolatum is present. Soil nutrient content does not change with epiphyte composition, is influenced by the microbiota, and P content decreases with the presence of epiphytes. The fitness costs show that the species compete, decreasing their survival and growth, but the competitive capacity differs between the species.
• We conclude that P. areolatum is an ecosystem engineer that promotes the creation of canopy soil but is a poor competitor. The results coincide with the model of succession by facilitation. Canopy soil is a slow-created component whose nutrient content does not depend on the epiphytic flora. In epiphyte mats, the dominant interactions are competitive, but there is also facilitation.