Have bird distributions shifted along an elevational gradient on a tropical mountain?

An upward shift in elevation is one of the most conspicuous species responses to climate change. Nevertheless, downward shifts and, apparently, the absences of response have also been recently reported. Given the growing evidence of multiple responses of species distributions due to climate change and the paucity of studies in the tropics, we evaluated the response of a montane bird community to climate change, without the confounding effects of land‐use change. To test for elevational shifts, we compared the distribution of 21 avian species in 1998 and 2015 using occupancy models. The historical data set was based on point counts, whereas the contemporary data set was based on acoustic monitoring. We detected a similar number of species in historical (36) and contemporary data sets (33). We show an overall pattern of no significant change in range limits for most species, although there was a significant shift in the range limit of eight species (38%). Elevation limits shifted mostly upward, and this pattern was more common for upper than lower limits. Our results highlight the variability of species responses to climate change and illustrate how acoustic monitoring provides an easy and powerful way to monitor animal populations along elevational gradients.     

Tree growth response along an elevational gradient: climate or genetics?

Environment and genetics combine to influence tree growth and should therefore be jointly considered when evaluating forest responses in a warming climate. Here, we combine dendroclimatology and population genetic approaches with the aim of attributing climatic influences on growth of European larch (Larix decidua) and Norway spruce (Picea abies). Increment cores and genomic DNA samples were collected from populations along a ~900-m elevational transect where the air temperature gradient encompasses a ~4 °C temperature difference. We found that low genetic differentiation among populations indicates gene flow is high, suggesting that migration rate is high enough to counteract the selective pressures of local environmental variation. We observed lower growth rates towards higher elevations and a transition from negative to positive correlations with growing season temperature upward along the elevational transect. With increasing elevation there was also a clear increase in the explained variance of growth due to summer temperatures. Comparisons between climate sensitivity patterns observed along this elevational transect with those from Larix and Picea sites distributed across the Alps reveal good agreement, and suggest that tree-ring width (TRW) variations are more climate-driven than genetics-driven at regional and larger scales. We conclude that elevational transects are an extremely valuable platform for understanding climatic-driven changes over time and can be especially powerful when working within an assessed genetic framework.     

Do differences in understory light contribute to species distributions along a tropical rainfall gradient?

In tropical forests, regional differences in annual rainfall correlate with differences in plant species composition. Although water availability is clearly one factor determining species distribution, other environmental variables that covary with rainfall may contribute to distributions. One such variable is light availability in the understory, which decreases towards wetter forests due to differences in canopy density and phenology. We established common garden experiments in three sites along a rainfall gradient across the Isthmus of Panama in order to measure the differences in understory light availability, and to evaluate their influence on the performance of 24 shade-tolerant species with contrasting distributions. Within sites, the effect of understory light availability on species performance depended strongly on water availability. When water was not limiting, either naturally in the wetter site or through water supplementation in drier sites, seedling performance improved at higher light. In contrast, when water was limiting at the drier sites, seedling performance was reduced at higher light, presumably due to an increase in water stress that affected mostly wet-distribution species. Although wetter forest understories were on average darker, wet-distribution species were not more shade-tolerant than dry-distribution species. Instead, wet-distribution species had higher absolute growth rates and, when water was not limiting, were better able to take advantage of small increases in light than dry-distribution species. Our results suggest that in wet forests the ability to grow fast during temporary increases in light may be a key trait for successful recruitment. The slower growth rates of the dry-distribution species, possibly due to trade-offs associated with greater drought tolerance, may exclude these species from wetter forests.     

Predation on artificial bird nests along an urban gradient: predatory risk or relaxation in urban environments?

Urbanization - the anthropogenic conversion of natural ecosystems into human-dominated ecosystems - has occurred on global scales. The human-dominated landscape presents particular challenges to researchers because the effects of urbanization on ecological processes are not well understood. We investigated the influence of urbanization on predation by conducting an artificial nest experiment along an urban gradient of six sites ranging from natural to urbanized ecosystems. Previous hypotheses suggest that predation pressures in urban environments will either 1) increase because of the high abundance of exotic species which act as predators or 2) decrease due to the lack of natural predators. To determine relative predation pressures among sites along the urban gradient, we monitored the fates of 16 artificial avian nests at each of the six sites for a total of 96 nests in each year (1996, 1997). We analyzed the dependency of nest fate (depredated or undisturbed) on intensity of urbanization (sites along the urban gradient), nest height (ground, above-ground), and year using loglinear models. The frequency of nest predation was strongly dependent on site along the urban gradient, indicating that urbanization intensity was an important determinant of nest fate. Predation pressure exhibited an overall decline from natural to urban sites in both years, suggesting that urban environments have low predation pressures relative to natural areas. The predatory relaxation in urban environments may partially explain the greater abundance of some species in urban environments, particularly urban exploiters such as european starlings Sturnis vulgaris. house sparrows Passer domesticus. and rock doves Columba livia.     

Maternal modulation of natal dispersal in a passerine bird: an adaptive strategy to cope with parasitism?

The decision of how far to disperse from the natal territory has profound and long-lasting consequences for young animals, yet the optimal dispersal behavior often depends on environmental factors that are difficult or impossible to assess by inexperienced juveniles. Natural selection thus favors mechanisms that allow the adaptive and flexible adjustment of the offspring's dispersal behavior by their parents via either paternal or maternal effects. Here we show that different dispersal strategies maximize the reproductive success of young great tits (Parus major) originating from a parasite-infested or a parasite-free nest and demonstrate that differential transfer of maternal yolk androgens in response to parasitism can result in a modification of the offspring's dispersal behavior that appears adaptive. It demonstrates that prenatal maternal effects are an important yet so far neglected determinant of natal dispersal and highlights the potential importance of maternal effects in mediating coevolutionary processes in host-parasite systems.     

Does coping style predict optimization? An experimental test in a wild passerine bird

A number of studies have suggested that avian brood size is individually optimized. Yet, optimal reproductive decisions likely vary owing to among-individual differences in environmental sensitivity. Specifically, ‘proactive’ individuals who do not track environmental changes may be less able to produce optimal brood sizes than ‘reactive’ individuals who have more precise local environmental knowledge. To test this, we quantified exploratory behaviour (a proxy for proactivity) in a great tit (Parus major) population, manipulated brood sizes (reduced, control, enlarged) and evaluated whether individuals of dissimilar coping style differed in their level of optimization. If reactive females behaved optimally, any deviation from their original brood size should lower fitness, whereas this should not be the case for proactive females. Reactive females indeed performed best at their natural brood size, whereas proactive females performed best when raising an enlarged brood. These findings imply that proactive females produced sub-optimal brood sizes. We speculate that proactive females might (i) take decisions based on biased perception of their environment, (ii) face energetic constraints in offspring production and/or (iii) be more willing to invest into current reproduction when given the option. Our findings provide experimental evidence for coping style-related differences in optimal reproductive decisions and life-history strategies.     

What drives the species richness patterns of non‐volant small mammals along a subtropical elevational gradient?

The biodiversity of non‐volant small mammals along an extensive subtropical elevational gradient was studied for the first time on Gongga Mountain, the highest mountain in Hengduan Mountain ranges in China, located in one of the 25 global biodiversity hotspots. Non‐volant small mammals were replicate sampled in two seasons at eight sampling sites between 1000 and 4200 m elevation on the eastern slope of Gongga Mountain. In all, 726 individual small mammals representing 25 species were documented in 28 800 trap nights. The species richness pattern for non‐volant small mammals along the elevational gradients was hump‐shaped with highest richness at mid‐elevations. However, different richness patterns emerged between endemic and non‐endemic species, between larger‐ranged and smaller‐ranged species and between rodents and insectivores. Temperature, precipitation, plant species richness and geometric constraints (mid‐ domain effect) were most significant in explaining species richness patterns. Based on the analysis of simple ordinary least squares (OLS) and stepwise multiple regressions, the overall richness pattern, as well as the pattern of insectivores, endemic species and larger‐ranged species showed strong correlation with geometric constraint predictions. However, non‐endemic species richness was more strongly correlated with temperature, while rodent richness was correlated with plant species richness. Our study shows that no single key factor can explain all richness patterns of non‐volant small mammals. We need to be cautious in summarizing a general richness pattern of large species groups (e.g. small mammals or mammals) from species in smaller groups having different ecological distributions and life histories. Elevational richness patterns and their driving factors for small mammals are more likely dependent on what kind of species we study.     

Facilitation consequences for reproduction of the benefactor cushion plant Laretia acaulis along an elevational gradient: costs or benefits?

Environmental stress may favour facilitative interactions among plants but whether these interactions are positive for the benefactor and how this depends on stress factors, remains to be determined. We studied the effect of beneficiary cover and biomass on reproduction of the benefactor cushion plant Laretia acaulis (Apiaceae) in the central Chilean Andes during three years. Study sites were situated along an elevational gradient at 2600, 2800, 3000 and 3150 m a.s.l. This range comprises a cold‐ and a drought‐stress gradient, with moisture increasing and temperature decreasing with elevation. We studied the effect of natural gradients in beneficiary cover and of experimental cover removal on cushion flower and fruit production. Beneficiary cover had a negative effect on flower production but not on infructescence and fruit densities or fruit weights. A positive effect of beneficiaries on the fraction of flowers converted into fruits was detected for hermaphrodite cushions. The effect of beneficiary cover on flowering was independent of elevation or cushion gender, although these latter factors explained most of the variation. Removing the aboveground parts of the beneficiaries positively affected flowering at 2800 m a.s.l. but not at the other elevations. Our results suggest negative effects of facilitation on L. acaulis flowering, but these are neutralized in fruit production. Surprisingly, this conclusion holds along the entire elevational or stress gradient. This suggests that this system of facilitation is evolutionarily stable and not very sensitive to environmental change. It remains to be tested, however, whether facilitation affects fitness via growth and long‐term survival in these slow‐growing alpine cushions.     

Patterns of grassland community composition and structure along an elevational gradient on the Qinghai–Tibet Plateau

青藏高原草地群落组成和结构的海拔梯度格局 青藏高原高寒草地是维持区域生态安全的天然屏障,也在一定程度上造就了该区域较高的生物多样性。然而,我们对青藏高原高寒草地植物群落组成和结构的海拔分布格局及其自身维持机制仍知之甚少。本研究在青藏高原东北部沿公路形成的海拔梯度设置了39个实验样地(海拔跨度为2800–5100m),每个样地设5个调查样方进行群落调查,包括物种组成、高度、盖度,评估青藏高原高寒草地植物群落的α和β多样性的海拔梯度格局及其影响因素。研究结果发现草地群落高度随着海拔的增加而显著降低,而群落盖度变化却不显著。随着海拔的增加,植物物种丰富度(α多样性)显著增加,而群落变异性(β多样性)显著降低。约束聚类分析表明,随海拔增加草地群落结构逐渐发生变化,基于此,在这种变化过程中,我们监测到3个渐变的海拔间断点,分别在海拔3640、4252和4333 m处。结构方程模型(SEM)表明,降水增加和温度降低对α多样性有显著的正向作用,但植物群落α多样性的变化显著改变群落变异性。以上结果表明,青藏高原的群落组成和结构沿海拔梯度发生了从量变到质变的过程。     

Does thermal history affect metabolic plasticity?: a study in three Phyllotis species along an altitudinal gradient

(1) The aim of this study was to understand the effects of thermal history in metabolic features such as maximum (MMR) and basal (BMR) metabolic rates, as well as in metabolic plasticity, considered as the total variation of MMR and BMR during the acclimation period. (2) We studied three species of the genus Phyllotis, from different thermal environments, in an altitudinal gradient from sea level to 3800m.a.s.l. Animals were acclimated to contrasting temperatures of 5 and 30 degrees C. To determine the metabolic flexibility, MMR was measured at intervals of 6 days during the acclimation period, while BMR values were obtained at the end of acclimations. Aerobic scope and the rates of change of MMR were estimated in all populations. (3) High- and low-altitude rodents did not show differences in BMR. However, both upper and lower limits of MMR, as well as aerobic scope, were significantly different between high- and low-altitude species, indicating similar ranges of metabolic plasticity. On the other hand, the rates of change of MMR were similar in all populations. (4) Our results indicate that thermal history has a profound effect on the individuals' thermogenic capacity, probably in both phylogenetic and ontogenetic levels. Low-altitude species could not increase MMR to the same levels as high-altitude species, while the later were unable to decrease MMR to achieve the values of the low-altitude species.     

Ecological character displacement in competing common green lacewings in Europe: a route to speciation?

Competitive interactions among the three common green lacewing species of the Chrysoperla complex in Europe are presented. Four morphological characters were registered as semi‐quantitative traits to assess variation between populations. Data were obtained from 325 adults originating from 14 locations. A statistical approach was performed in conjunction with observations of ecological traits and a survey of genetic relationships assessed by mitochondrial DNA cytochrome b analysis. The results indicate character displacement in Chrysoperla affinis when living in sympatry with Chrysoperla carnea. The morphological traits of Ch. affinis sympatric with Ch. carnea differ from those of allopatric Ch. affinis, although the genetic distance between those strains was weak. This was associated with a large difference in microhabitats and premating behaviour. Variation in courtship songs manifests a process of speciation within related populations. Thus, that ecological switching can also promote character displacement is supported. This suggests that character displacement acts through phenotypic variability and mediates negative interactions among species. These findings suggest that the highest rate of speciation occurs in groups of species in which competition is the strongest. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 292–300.     

Needle δ13C and mobile carbohydrates in Pinus koraiensis in relation to decreased temperature and increased moisture along an elevational gradient in NE China

A tree’s crown interacts with atmospheric variables such as CO₂, temperature, and humidity. Physioecology of leaves/needles (e.g. δ¹³C, mobile carbohydrates, and nitrogen) is, therefore, strongly affected by microclimate in and surrounding a tree crown. To understand the physiological responses of leaves to changes in air temperature and moisture, we measured δ¹³C, soluble sugars, starch, and total nitrogen (N) concentrations in current year and 1-yr-old needles of Pinus koraiensis trees, and compared the growing season air temperature and relative humidity within and outside P. koraiensis crowns along an elevational gradient from 760 to 1,420 m a.s.l. on Changbai Mountain, NE China. Our results indicated that needle N and mobile carbohydrates concentrations, as well as needle δ¹³C values changed continuously with increasing elevation, corresponding to a continuous decrease in air temperature and an increase in relative humidity. Needle carbon and nitrogen status is highly significantly negatively correlated with temperature, but positively correlated with relative humidity. These results indicate that increases in air temperature in combination with decreases in relative humidity may result in lower levels of N and mobile carbohydrates in P. koraiensis trees, suggesting that future climate changes such as global warming and changes in precipitation patterns will directly influence the N and carbon physiology at P. koraiensis individual level, and indirectly affect the competitive ability, species composition, productivity and functioning at the stand and ecosystem level in NE China. Due to the relatively limited range of the transect (760–1,420 m) studied, further research is needed to explain whether the present results are applicable to scales across large elevational gradients.     

Can changes in ant diversity along elevational gradients in tropical and subtropical Australian rainforests be used to detect a signal of past lowland biotic attrition?

Increasing temperatures are predicted to have profound effects on montane ecosystems. In tropical forests, biotic attrition may reduce lowland diversity if losses of species due to upslope range shifts are not matched by influxes of warmer‐adapted species, either because there are none or their dispersal is impeded. Australian rainforests consist of a north–south chain of patches, broken by dry corridors that are barriers to the dispersal of rainforest species. These rainforests have repeatedly contracted and expanded during Quaternary glacial cycles. Many lowland rainforests are expansions since the Last Glacial Maximum and may, therefore, show a signal of historical biotic attrition. We surveyed ants from replicated sites along three rainforest elevational transects in eastern Australia spanning 200 to 1200 m a.s.l. and nearly 14° of latitude. We examined elevational patterns of ant diversity and if there was possible evidence of lowland biotic attrition. Each transect was in a different biogeographic region; the Australian Wet Tropics (16.3°S), the central Queensland coast (21.1°S) and subtropical south‐eastern Queensland (28.1°S). We calculated ant species density (mean species per site) and species richness (estimated number of species by incorporating site‐to‐site species turnover) within elevational bands. Ant species density showed no signal of lowland attrition and was high at low and mid‐elevations and declined only at high elevations at all transects. Similarly, estimated species richness showed no evidence of lowland attrition in the Wet Tropics and subtropical south‐east Queensland; species richness peaked at low elevations and declined monotonically with increasing elevation. Persistence of lowland rainforest refugia in the Wet Tropics during the Last Glacial Maximum and latitudinal range shifts of ants in subtropical rainforests during the Holocene climatic optimum may have counteracted lowland biotic attrition. In central Queensland, however, estimated richness was similar in the lowlands and mid‐elevations, and few ant species were indicative of lower elevations. This may reflect historical biotic attrition due perhaps to a lack of lowland glacial refugia and the isolation of this region by a dry forest barrier to the north.     

Elevational speciation in action? Restricted gene flow associated with adaptive divergence across an altitudinal gradient

Evolutionary theory predicts that divergent selection pressures across elevational gradients could cause adaptive divergence and reproductive isolation in the process of ecological speciation. Although there is substantial evidence for adaptive divergence across elevation, there is less evidence that this restricts gene flow. Previous work in the boreal chorus frog (Pseudacris maculata) has demonstrated adaptive divergence in morphological, life history and physiological traits across an elevational gradient from approximately 1500–3000 m in the Colorado Front Range, USA. We tested whether this adaptive divergence is associated with restricted gene flow across elevation – as would be expected if incipient speciation were occurring – and, if so, whether behavioural isolation contributes to reproductive isolation. Our analysis of 12 microsatellite loci in 797 frogs from 53 populations revealed restricted gene flow across elevation, even after controlling for geographic distance and topography. Calls also varied significantly across elevation in dominant frequency, pulse number and pulse duration, which was partly, but not entirely, due to variation in body size and temperature across elevation. However, call variation did not result in strong behavioural isolation: in phonotaxis experiments, low‐elevation females tended to prefer an average low‐elevation call over a high‐elevation call, and vice versa for high‐elevation females, but this trend was not statistically significant. In summary, our results show that adaptive divergence across elevation restricts gene flow in P. maculata, but the mechanisms for this potential incipient speciation remain open.     

Can we predict butterfly diversity along an elevation gradient from space?

An important challenge in ecology is to predict patterns of biodiversity across eco‐geographical gradients. This is particularly relevant in areas that are inaccessible, but are of high research and conservation value, such as mountains. Potentially, remotely‐sensed vegetation indices derived from satellite images can help in predicting species diversity in vast and remote areas via their relationship with two of the major factors that are known to affect biodiversity: productivity and spatial heterogeneity in productivity. Here, we examined whether the Normalized Difference Vegetation Index (NDVI) can be used effectively to predict changes in butterfly richness, range size rarity and beta diversity along an elevation gradient. We examined the relationship between butterfly diversity and both the mean NDVI within elevation belts (a surrogate of productivity) and the variability in NDVI within and among elevation belts (surrogates for spatial heterogeneity in productivity). We calculated NDVI at three spatial extents, using a high spatial resolution QuickBird satellite image. We obtained data on butterfly richness, rarity and beta diversity by field sampling 100 m quadrats and transects between 500 and 2200 m in Mt Hermon, Israel. We found that the variability in NDVI, as measured both within and among adjacent elevation belts, was strongly and significantly correlated with butterfly richness. Butterfly range size rarity was strongly correlated with the mean and the standard deviation of NDVI within belts. In our system it appears that it is spatial heterogeneity in productivity rather than productivity per se that explained butterfly richness. These results suggest that remotely‐sensed data can provide a useful tool for assessing spatial patterns of butterfly richness in inaccessible areas. The results further indicate the importance of considering spatial heterogeneity in productivity along elevation gradients, which has no lesser importance than productivity in shaping richness and rarity, especially at the local scale.     

Qualitative variation in roadside weed vegetation along an urban–rural road gradient

Information on mechanisms of differentiation or homogenization of urban floras is deficient, despite their importance for urban nature conservation and urban land management. Roads, as a major human promoter of urbanization, can be an initial habitat for plants dispersed by transportations. We assumed that variation in weed vegetation along urban–rural roadside gradients is small, particularly in curbside cracks. We classified vegetation occurring in curbside cracks along the National Route 3 in southern Japan and compared the characteristics of the vegetation types recognized. Species in curbside cracks were recorded on 40 plots. Three vegetation types were classified, in part related to surrounding land-use types. Although the Shannon–Wiener diversity index and the number of native species clearly differed among the vegetation types in the curbside cracks, no significant differences between the three floristic vegetation types were found in the number of non-natives and invasive alien species. This may result from the small specificity and complexity of landscape structures, due to the continuity and connectivity of paved-road networks. Of the 122 species, 44 were non-natives or invasive alien species. The vegetation types generally involved the same plant families, with large numbers of species from the Asteraceae and Poaceae, regardless of vegetation types, but frequency of occurrence of the two families clearly differed between natives and non-natives and invasive alien species. Ephemeral non-natives and invasive alien species, particularly Asteraceae and Poaceae, seem to have advantageous dispersal strategies or low habitat dependency facilitating their occurrence in curbside cracks, regardless of adjacent land uses and the urban–rural landscape gradient. Expansion of these species may cause a homogenization of regional floras along roads.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

δ¹³C values of C₃ plants are indicators of plant carbon–water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant δ¹³C at the landscape scale. We determined variation in leaf δ¹³C, leaf nitrogen per leaf area (N_area), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 × 12-m belt transect spanning an upland–lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The δ¹³C values of Prosopis glandulosa (deciduous N₂-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1–4‰ with decreasing elevation, with the δ¹³C value of P. glandulosa leaves being 1–3‰ higher than those of the two shrub species. Contrary to theory and results from previous studies, δ¹³C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N_area was positively correlated with leaf δ¹³C of all species (p < 0.01) and appeared to exert the strongest control over δ¹³C along this topoedaphic gradient. Since leaf N_area is positively related to photosynthetic capacity, plants with high leaf N_area are likely to have low p _I/p _a ratios and therefore higher δ¹³C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf δ¹³C. Following a progressive growing season drought in July/August, leaf δ¹³C decreased. The lower δ¹³C in August may reflect the accumulation of ¹³C-depleted epicuticular leaf wax. We suggest control of leaf δ¹³C along this topoedaphic gradient is mediated by leaf N_area rather than by stomatal conductance limitations associated with water availability.