Genetic variability of an endangered Bromeliaceae species (Pitcairnia albiflos) from the Brazilian Atlantic rainforest

Pitcairnia albiflos is a Bromeliaceae species endemic to Brazil that has been included as data-deficient in the extinction risk list of Brazilian flora. We analyzed genetic variability in P. albiflos populations using RAPD markers to investigate population structure and reproductive mechanisms and also to evaluate the actual extinction risk level of this species. Leaves of 56 individuals of P. albiflos from three populations were collected: Urca Hill (UH, 20 individuals), Chacrinha State Park (CSP, 24 individuals) and Tijuca National Park (TNP, 12 individuals). The RAPD technique was effective in characterizing the genetic diversity in the P. albiflos populations since it was possible to differentiate the populations and to identify exclusive bands for at least two of them. Even if there is low genetic diversity among them (CSP-UH = 0.463; CSP-TNP = 0.440; UH-TNP = 0.524), the populations seem to be isolated according to the low genetic diversity observed within them (H(pop) CSP = 0.060; H(pop) UH = 0.042; H(pop) TNP = 0.130). This fact might be the result of clonal and self-reproduction predominance and also from environmental degradation around the collection areas. Consequently, it would be important to protect all populations both in situ and ex situ to prevent the decrease of genetic variability. The low genetic variability among individuals of the same population confirms the inclusion of this species as critically endangered in the risk list for Brazilian flora.     

Interactions between aroids and arboreal mammals in the Brazilian Atlantic rainforest

The Brazilian Atlantic rainforest harbours at least 129 non-volant mammal species. There is also a diverse and numerous epiphytic flora associated with this Neotropical rainforest, including several species of Araceae. Nevertheless, knowledge of the ecology of this group of plants and its interaction with animals is poor. In this study we investigated the interactions between epiphytic aroids and arboreal mammals in the Atlantic forest. We carried out the field work between November 1995 and June 1997 in the Parque Estadual Intervales, southeastern Brazil, a still well preserved forest area of approximately 490 km². We analysed faecal samples from primates (Cebus apella and Brachyteles arachnoides), collected during observations in the field, and marsupials (Didelphis aurita, Micoureus demerarae, Gracilinanus microtarsus), collected from animals captured monthly or bi-monthly in live-traps. We recorded 17 species of Araceae in the study area. The seeds of eight species occurred in the faecal samples: Anthurium harrissi, Monstera adansonii, Philodendron corcovadense, P. appendiculatum, P. exymium, P. crassinervium, P. obliquifolium, and Heteropsis oblongifolia. Aroids were present in faecal samples of the primates during most of the year, with a mean monthly percentage of occurrence of about 56.2%. For marsupials, aroids also were a seasonally important food source, as between November and June the mean monthly percentage of occurrence in faecal samples was about 26.5%. Our data indicate that aroids are an important food source for arboreal and semi-arboreal mammals and that these animals may play an important role as seed dispersers of this group. Aroids and arboreal mammals appear to interact more strongly in Atlantic forest than in other Neotropical forests.     

Population density of primates in a large fragment of the Brazilian Atlantic rainforest

We performed a line transect survey (352.4 km) of primates in the Serra de Paranapiacaba, at one of the largest relatively undisturbed fragments of the Atlantic rainforest of Southeastern Brazil (ca. 1400 km²), in August 1998. The brown capuchin, Cebus apella nigritus, was the most common species found in the area (20 groups, density estimate: 5.31 ± 2.05 individuals per km², mean ± SE). Nine groups of the brown howler monkey, Alouatta guariba clamitans, and eight of the woolly spider monkey, Brachyteles arachnoides arachnoides, were also recorded, with preliminary density estimates of 0.79 ± 0.40 and 2.33 ± 1.37 individuals per km², respectively. Density estimates for these species in other fragments of Atlantic rainforest are reviewed, showing that densities in Paranapiacaba are among the lowest reported. It is suggested that the higher densities reported for isolated populations in small forest patches (<50 km²) is related to the absence of main primate predators, the density compensation phenomenon and the ecological plasticity of some primate species. In contrast, local extinction in many small patches is probably related to hunting pressure. Given the important primate populations found in the Paranapiacaba fragment, conservation strategies for the studied species should give priority to effective protection of the largest remnant fragments from illegal hunting and deforestation, rather than translocation of individuals or captive breeding programs to introduce monkeys in small forest fragments vulnerable to hunting and of uncertain future.     

Assessing future habitat availability for coastal lowland anurans in the Brazilian Atlantic rainforest

Global warming is expected to cause several modifications to physical environments, and sea level rise is a certain outcome. However, assessment of the potential impacts caused by sea level rise on biodiversity is still emerging. Therefore, we assessed the combined impact of global warming and sea level rise on the potential distribution of 19 coastal lowland anurans in the biodiversity hotspot Atlantic Forest. We applied a correlative species distribution model (SDM) (BIOCLIM) and GIS-based spatial analyses. We evaluated the extent of changes of potential distributions under extreme and moderate global warming scenarios as well as two extreme sea level rise scenarios. Our results suggest wide areas of suitable habitat for most species in the future. However, for 15% of these species the SDMs predict massive losses of range extent as a result of a combination of global warming and sea level rise. Such observations highlight an immediate need to consider the potential effects of sea level rise in conservation action plans. Since the current potential distribution of these anuran species is likely underestimated, we also analyzed their environmental niche under current conditions in order to provide a baseline for further field surveys. Considering this current state of knowledge for such species, species distribution modeling to help gather further information on unknown species is desirable.     

Distribution of Leguminosae tree species in different altitudinal levels along the Atlantic Rain Forest in the Brazilian coast

The Atlantic Forest is one of the most threatened tropical forests in the world. Leguminosae, by its great richness and dominance among arboreal stratum elements, is of major importance in the floristic composition and structure of this forest. We investigated the distribution of legume species on an altitudinal gradient to find out the altitudinal zones with higher richness of species; the altitudinal zones with greater floristic similarity; the possible presence of species that may be exclusive to certain vegetation types and the altitudinal amplitudes of those species, as well as the occurrence of species substitution along the altitudinal gradient. Therefore, thirty one studies conducted in different altitudinal levels between 5° S and 29° S were analyzed. A matrix with 142 tree species distributed in altitudinal zones (every 100 m) from sea level to 2100 m was built. The greatest species richness was observed in the Submontane Forest (50–500 m) with 92 species. The cluster analysis revealed a strong dissimilarity of the 1400–2100 m (Upper Montane) and 0–10 m zones (Restinga Forest). The Submontane and the Montane Forest share the highest number of species (38 ssp.). Forty species are unique to Submontane. Substitution of species was verified. Some species have their preferred habitat located at a specific altitudinal amplitude, as is the case of Inga laurina and I. subnuda (0–10 m), I. lanceifolia and Machaerium scleroxylon (800–1200 m). The Leguminosae, although well adapted to the first colonization and establishment of diverse environment, was poorly represented above 1500 m altitude.     

广西弄岗北热带喀斯特季节性雨林监测样地种群空间点格局分析

种群的空间分布格局是由多种机制的交互作用而形成,是探究生物多样性维持机制的基础。北热带喀斯特季节性雨林是我国北热带石灰岩山地具有地带性特征的植被,其生境的典型特征在于土层浅薄、岩石裸露率高、贮水能力低和周期性水淹,以及富钙强碱性环境等。本研究基于广西弄岗北热带喀斯特季节性雨林15 ha动态监测样地的第一次调查资料,采用双关联g(r)函数点格局方法,分析了雨林中出现个体数≥15株的160个种的种群空间分布格局以及不同类群间种群分布的差异。研究表明:160种木本植物中有146种在0–10 m的尺度上呈聚集分布,且随着空间尺度的增大,聚集度呈下降趋势;物种的种群聚集度与其物种多度、平均胸径和最大胸径成负相关;常绿物种的种群聚集度与落叶物种间差异不显著;不同生活型间的种群聚集度差异显著,总体表现为亚乔木层物种高于乔木层,灌木层物种显著高于亚乔木层;剔除生境异质性后大部分物种表现为随机分布,仅少部分物种的种群表现为聚集分布。这表明物种的功能属性如物种多度、生活型等可较好地预测物种的分布格局。此外,物种种群的分布格局还受生境异质性的影响,且不同物种受生境异质性影响的程度不同。     

Diversity and ecology of saxicolous vegetation mats on inselbergs in the Brazilian Atlantic rainforest

Inselbergs occur as mostly dome-shaped rock outcrops in all climatic and vegetational zones of the tropics. Consisting of Precambrian granites and gneisses, they form ancient and stable landscape elements. Due to harsh edaphic and microclimatic conditions, the vegetation of inselbergs differs markedly from those of the surroundings. Monocotyledonous mats form one of the most characteristic communities of this ecosystem. The floristic composition of this community was studied on six inselbergs located in the Brazilian Atlantic rainforest. Dominating are Bromeliaceae ( Alcantarea , Encholirium ), Velloziaceae ( Vellozia ), Cactaceae ( Coleocephalocereus ) and the Cyperaceae Trilepis . The alpha diversity of the mats was relatively uniform among the six outcrops. However, beta diversity varied considerably between the different sites. Beta diversity was highest at the most speciose locality indicating a high degree of stochasticity in colonization. In contrast to the low diversity mats on West African inselbergs, the Brazilian rock outcrops bear a floristically more diverse mat community rich in endemics. Thus the edaphically controlled inselberg vegetation reflects the outstanding diversity of the Mata Atlaântica. Possibly the high species richness of mats on East Brazilian inselbergs is a consequence of a large species-pool. The processes that regulate regional and local diversity in the Mata Atlaântica are not fully clear. It is assumed that historical (i.e. long-term stability) and biotic (i.e. evolutionary interchange of taxa between canopies and rock outcrops followed by differentiation of local populations) conditions have promoted high rates of speciation and their coexistence in isolated habitats. A danger to the unique vegetation of East Brazilian inselbergs is the establishment of invasive weeds.     

Intraspecific and interspecific trait variability in tadpole meta‐communities from the Brazilian Atlantic rainforest

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Effect of species rarity on the accuracy of species distribution models for reptiles and amphibians in southern California

Aim Several studies have found that more accurate predictive models of species’ occurrences can be developed for rarer species; however, one recent study found the relationship between range size and model performance to be an artefact of sample prevalence, that is, the proportion of presence versus absence observations in the data used to train the model. We examined the effect of model type, species rarity class, species’ survey frequency, detectability and manipulated sample prevalence on the accuracy of distribution models developed for 30 reptile and amphibian species. Location Coastal southern California, USA. Methods Classification trees, generalized additive models and generalized linear models were developed using species presence and absence data from 420 locations. Model performance was measured using sensitivity, specificity and the area under the curve (AUC) of the receiver‐operating characteristic (ROC) plot based on twofold cross‐validation, or on bootstrapping. Predictors included climate, terrain, soil and vegetation variables. Species were assigned to rarity classes by experts. The data were sampled to generate subsets with varying ratios of presences and absences to test for the effect of sample prevalence. Join count statistics were used to characterize spatial dependence in the prediction errors. Results Species in classes with higher rarity were more accurately predicted than common species, and this effect was independent of sample prevalence. Although positive spatial autocorrelation remained in the prediction errors, it was weaker than was observed in the species occurrence data. The differences in accuracy among model types were slight. Main conclusions Using a variety of modelling methods, more accurate species distribution models were developed for rarer than for more common species. This was presumably because it is difficult to discriminate suitable from unsuitable habitat for habitat generalists, and not as an artefact of the effect of sample prevalence on model estimation.     

Habitat selection by large mammals in a southern Brazilian Atlantic Forest

Habitat selection, which is mainly a consequence of competition and predation, allows species to coexist. The present study was conducted in two reserves in an Atlantic Forest area in Santa Catarina State, southern Brazil, and provided information on several large mammal species through photographic records. Records were related to certain environmental parameters, such as width of passages (trails and roads), vegetation density and proximity to water, in order to assess the relationship between each mammal species and its microhabitat. Thirty-two camera trap stations were placed during 17 months for 150.8 (±62.2) days on average. Terrestrial mammals tended to use different habitats: Puma concolor used mainly dirt roads and open areas; Leopardus pardalis, Cerdocyon thous and Nasua nasua used more large trails and intermediate-forested sites; and Cuniculus paca, Dasypus novemcinctus, Leopardus tigrinus, Eira barbara and Leopardus wiedii were recorded more often on narrow trails and in densely forested sites. Some of these forest species, such as D. novemcinctus, C. paca and L. pardalis, also showed relationships with watercourses. Information on habitat selection allows more effort to be addressed to the habitat associated with focal species, and indicates the significance of environmental heterogeneity, which makes it possible for species to coexist.     

Genetic diversity of reintroduced tree populations in restoration plantations of the Brazilian Atlantic Forest

Long‐term ecological success of large‐scale restoration programs planned for the next decades will rely on genetic diversity (GD) of reintroduced or colonizing species, a limiting factor in highly fragmented landscapes. In small and isolated natural remnants or restoration areas, substantial reduction in population's size or connectivity may lead to local extinctions due to the accumulation of deleterious recessive alleles and ongoing reduction of fecundity, plant vigor, recruitment success, and adaptive potential. Despite the paramount role of GD for species persistence, its levels in restoration programs are poorly known. We assessed the GD of four model tree species (different succession stages, dispersal, and pollination syndromes) from the Brazilian Atlantic Forest, comparing two high‐diversity restoration plantations, one forest fragment and one conserved remnant. Contrary to the expectation that the plantation strategies adopted in the restoration programs could result in genetic composition homogenization, we found that restoration areas established heterogeneous genetic groups with similar levels of neutral GD and inbreeding to those observed in natural forest remnants. This pattern was consistent across the four functionally different tree species, despite some species idiosyncrasies. For instance, we observed lower allelic richness in early successional species in restoration sites, suggesting that some species may be more prone to reintroduction with lower GD. Thus, we advocate the use of high GD levels in restoration to support biodiversity conservation in human‐modified landscapes, thus reinforcing the role of ecological restoration for recovering the diversity of genes—the basic constituent of biodiversity.     

Fine-scale genetic dynamics of a dominant neotropical tree in the threatened Brazilian Atlantic Rainforest

We present a case study of the relationship between spatial genetic structure (SGS) and age structure in Protium spruceanum (Burseraceae), an insect-pollinated, mass-fruiting, and secondary bird-dispersed tree, as determined through variation in allozyme loci. Using ten polymorphic loci, we investigated spatial and temporal patterns of a genetic structure within a 40?m?×?60?m plot in a small (1.0?ha) fragment of Atlantic Rainforest to investigate the processes shaping the distribution of genetic diversity. Individuals (n?=?345) from seedlings to adults were grouped and analyzed in four diameter classes. The results showed a high average level of genetic diversity (H _e?=?0.438), but genetic diversity parameters did not vary significantly among cohorts. The spatial distribution pattern analysis of individuals showed significant levels of aggregation among small- and medium-diameter classes and random distribution among the highest diameter class, likely due to processes of competitive thinning. There was an association between demographic and SGS at short distances (less than 10?m) which is likely the consequence of restricted seed dispersal. The degree of SGS decreased across small- to large-diameter classes. We inferred that limited seed dispersal and subsequent density-dependent mortality from the family clusters are responsible for the observed changes in fine-scale SGS across different demographic classes.     

Interactions between terrestrial mammals and the fruits of two neotropical rainforest tree species

Mammalian frugivory is a distinctive biotic interaction of tropical forests; however, most efforts in the Neotropics have focused on cases of animals foraging in the forest canopy, in particular primates and bats. In contrast much less is known about this interaction when it involves fruits deposited on the forest floor and terrestrial mammals. We conducted a camera-trapping survey to analyze the characteristics of the mammalian ensembles visiting fruits of Licania platypus and Pouteria sapota deposited on the forest floor in a well preserved tropical rainforest of Mexico. Both tree species produce large fruits but contrast in their population densities and fruit chemical composition. In particular, we expected that more species of terrestrial mammals would consume P. sapota fruits due to its higher pulp:seed ratio, lower availability and greater carbohydrate content. We monitored fruits at the base of 13 trees (P. sapota, n = 4 and L. platypus, n = 9) using camera-traps. We recorded 13 mammal species from which we had evidence of 8 consuming or removing fruits. These eight species accounted for 70% of the species of mammalian frugivores active in the forest floor of our study area. The ensemble of frugivores associated with L. platypus (6 spp.) was a subset of that associated with P. sapota (8 spp). Large body-sized species such as Tapirus bairdii, Pecari tajacu and Cuniculus paca were the mammals more frequently interacting with fruits of the focal species. Our results further our understanding of the characteristics of the interaction between terrestrial mammalian frugivores and large-sized fruits, helping to gain a more balanced view of its importance across different tropical forests and providing a baseline to compare against defaunated forests.     

Tibouchina pulchra (Melastomataceae): reproductive biology of a tree species at two sites of an elevational gradient in the Atlantic rainforest in Brazil

Low-temperature environments interfere with plant reproduction by reducing the frequency of pollinators, and this may favour reproductive strategies such as self-pollination and apomixis. Tibouchina pulchra is a common tree species that occurs at high and low sites of the Brazilian Atlantic rainforest. This study focussed on the pollination biology and breeding system of this species, describing the pollinators and the reproductive success at the two sites of an elevational gradient. Observations were made to determine extent of flowering and fruiting, to identify the richness and abundance of pollinators, and to record data on the floral and reproductive biology at these two sites. Despite more dense flowering at the high site, five visits of bees (two species) were recorded during the observation time (60?h), whereas at the low site there were 948 visits (seven species) during the same period. In contrast with the low site, the flowers of the high site released and received few pollen grains on the stigma. At the high site less fruit was set with fewer seeds as a result of open pollination than at the low site; at that site, however, more seeds were obtained from cross-pollination than at the low site. Tibouchina pulchra is self-compatible; however it is not apomictic and needs pollinators for seed set at both sites. Life-history traits other than the breeding system, for example more dense flowering, advantage of greater fertility in cross-pollination, and multiple reproductive events during the lifetime of the tree may reduce inbreeding depression, increase the hybrid vigour, and balance the lack of pollinators at the high site.     

Photosynthetic induction responses of two rainforest tree species in relation to light environment

Photosynthetic induction of in situ saplings of two Costa Rican rainforest tree species wre compared in relation to their light environment, using infrared gas analysis and hemispherical photography. The species studied were Dipteryx panamensis, a climax species found in bright microsites, and Cecropia obtusifolia, a pioneer species. In the morning, when leaves were most responsive, induction time necessary to reach 90% of the lightsaturated rate of photosynthesis was on average 16 min for Dipteryx and 10 min for Cecropia. However, induction times for both species increased in the afternoon resulting in shorter daily average induction times for Dipteryx than for Cecropia. Dipteryx also maintained higher levels of induction for a longer period under low light conditions than did Cecropia. The two species differed in the way they adjusted to light availability. Dipteryx saplings growing in shady sites had faster rates of induction than saplings growing in bright sites, with no difference in light-saturated photosynthetic rate. In contrast, Cecropia saplings growing in bright sites had higher light-saturated photosynthetic rates than saplings growing in shady sites, with no difference in rates of induction. Dipteryx appears to exploit temporal variation in light availability by refining the quickness of the induction response to the light environment, while Cecropia adjusts its scale of exploitation by realizing a higher lightsaturated photosynthetic rate in sites of higher light.     

Spatio-temporal microhabitat use by two co-occurring species of scorpions in Atlantic rainforest in Brazil

With the increasing devastation of the tropical rain forest, there is a critical need to understand how animal forest communities are structured and how habitat degradation will affect these communities. We conducted a field survey to investigate the microhabitat preferences of two co-occurring species of scorpions (Tityus pusillus and Ananteris mauryi) in a fragment of Atlantic rainforest, as well as their abundance and their ecological niche, during both the dry and rainy seasons. Behavioural aspects related to the use of the environment and the proportions of juveniles and adults are also described. The occurrence of intra- and interspecific coexistence was assessed by active search. In addition, pitfall catches were used to assess the structure of the population in the dry and rainy seasons. The differential patterns of spatial distribution in the litter layers provided evidence of partial niche partitioning between the two coexisting scorpion species depending on age and climatic conditions. Abundance, foraging behaviour and age structure (juveniles and adults) were seasonally influenced. We conclude that the diverse and subtle behaviours involved in interaction and habitat use may facilitate species coexistence. Resource partitioning and refuge sharing on a temporal and/or spatial scale, as well as predation pressure, may drive the dynamics and spatial distribution of scorpion species in the rain forest environment.     

Structure,function and floristic relationships of plant communities in stressful habitats marginal to the Brazilian Atlantic rainforest

The Brazilian Atlantic rainforest consists of a typical tropical rainforest on mountain slopes, and stands out as a biodiversity hotspot for its high species richness and high level of species endemism. This forest is bordered by plant communities with lower species diversity, due mostly to more extreme environmental conditions than those found in the mesic rainforest. Between the mountain slopes and the sea, the coastal plains have swamp forests, dry semi-deciduous forests and open thicket vegetation on marine sand deposits. At the other extreme, on top of the mountains (>2000 m a.s.l.), the rainforest is substituted by high altitude fields and open thicket vegetation on rocky outcrops. Thus, the plant communities that are marginal to the rainforest are subjected either to flooding, drought, oceanicity or cold winter temperatures. It was found that positive interactions among plants play an important role in the structuring and functioning of a swamp forest, a coastal sandy vegetation and a cold, high altitude vegetation in the state of Rio de Janeiro. Moreover, only a few species seem to adopt this positive role and, therefore, the functioning of these entire systems may rely on them. Curiously, these nurse plants are often epiphytes in the rainforest, and at the study sites are typically terrestrial. Many exhibit crassulacean acid metabolism. Conservation initiatives must treat the Atlantic coastal vegetation as a complex rather than a rainforest alone.     

Growth rates of selected Australian tropical rainforest tree species under controlled conditions

Controlled environment treatments were applied to assess the effects of temperature on the seedling mortality and growth rates of Toona australis and Flindersia brayleyana, two tropical rainforest tree species from northeast Queensland, Australia. Past workers have assigned these two species to the same ecological niche in terms of their response to canopy disturbance and gap-phase regeneration; however, their geographic ranges are very different. The hypothesis was that the species confined to the warm tropics (F. brayleyana) would have higher seedling mortality and a slower growth rate at lower temperatures than the species that occurs over a wide latitudinal range from the warm tropics to cooler temperate environments (T. australis). Significant differences were found in the growth rates of these two species in the warm (29/22° C) and cool (22/10°C), but not the intermediate (24/16° C), day/night temperature regimes. Their growth rates both decreased with decreasing temperature, but the decrease was significantly less for F. brayleyana which had the faster growth rate and lower seedling mortality in the cool regime. These results led to the rejection of the hypothesis and a test of the assignment of these two species to the same ecological niche. The test involved monitoring their growth to sapling-size in the intermediate temperature regime together with four other co-occurring tropical rainforest tree species belonging to different ecological niches. The growth rates and proportions of above-ground biomass allocated to woody tissue distinguished T. australis and a fast-growing pioneer species from F. brayleyana and three primary forest species. The stem heights and aboveground biomass of T. australis and the pioneer species exceeded the other four species by factors ranging from two to five. It is concluded that T. australis does not belong to the same ecological niche as F. brayleyana, and it is recommended that more research be conducted on the ecotypic temperature responses of the taxon T. australis.     

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence

We measured the response of dark respiration (R_d) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communites in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R_d at a reference temperature (R₁₀) and the extent to which R_d increased with temperature (described by E_o, a parameter related to the energy of activation, or the change in R_d over a 10°C range, Q₁₀). Site averaged E_o ranged from 44.4 kJ mol^–1 K^–1 at the 60-year-old site to 26.0 kJ mol^–1 K^–1 at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E_o or Q₁₀) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A_max/R_d) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.