Mean annual temperature influences local fine root proliferation and arbuscular mycorrhizal colonization in a tropical wet forest

Mean annual temperature (MAT) is an influential climate factor affecting the bioavailability of growth‐limiting nutrients nitrogen (N) and phosphorus (P). In tropical montane wet forests, warmer MAT drives higher N bioavailability, while patterns of P availability are inconsistent across MAT. Two important nutrient acquisition strategies, fine root proliferation into bulk soil and root association with arbuscular mycorrhizal fungi, are dependent on C availability to the plant via primary production. The case study presented here tests whether variation in bulk soil N bioavailability across a tropical montane wet forest elevation gradient (5.2°C MAT range) influences (a) morphology fine root proliferation into soil patches with elevated N, P, and N+P relative to background soil and (b) arbuscular mycorrhizal fungal (AMF) colonization of fine roots in patches. We created a fully factorial fertilized root ingrowth core design (N, P, N+P, unfertilized control) representing soil patches with elevated N and P bioavailability relative to background bulk soil. Our results show that percent AMF colonization of roots increased with MAT (r² = .19, p = .004), but did not respond to fertilization treatments. Fine root length (FRL), a proxy for root foraging, increased with MAT in N+P‐fertilized patches only (p = .02), while other fine root morphological parameters did not respond to the gradient or fertilized patches. We conclude that in N‐rich, fine root elongation into areas with elevated N and P declines while AMF abundance increases with MAT. These results indicate a tradeoff between P acquisition strategies occurring with changing N bioavailability, which may be influenced by higher C availability with warmer MAT.     

海南霸王岭热带雨林常见植物丛枝菌根真菌调查

对海南霸王岭热带雨林的12科16种常见植物的丛枝菌根状况进行了调查,用碱解离-酸性品红染色法进行了真菌鉴定。结果表明,13种植物形成典型的丛枝菌根,占所调查植物的81%;3种植物没有形成丛枝菌根,占所调查植物的19%。用湿筛沉淀法从这些植物根际土壤中共分离鉴定出了3属11种丛枝菌根真菌(AMF),即无梗囊霉属(Acaulos-pora)3种,球囊霉属(Glomus)7种,巨孢囊霉属(Gigaspora)1种;其中,球囊霉属是样地的优势属。在AMF中,孔窝无梗囊霉(A.foveata)分离频率最高,在14种植物的根际土中都有发现;此外,大果球囊霉(G.macrocarpum)的相对多度最大,为59%,具有最强的产孢能力。同时,在11种植物的根中发现了深色有隔内生真菌(DSE),占调查植物的69%;其中,11种植物同时被DSE和AMF感染。     

西双版纳热带次生林中的丛枝菌根调查

对西双版纳热带次生林中13个科的26种植物根系的丛枝菌根真菌（Arbuscular mycorrhizal fongii,AMF）侵染情况进行了研究,并从这些植物的根围土壤中分离鉴定了隶属于球囊霉属（Clomus）、巨孢囊霉属（Gigaspora）、盾巨孢囊霉属（Scutellospora）和无梗囊霉属（Acaulospora）的11种丛枝菌根真菌。该地次生林中AMF的孢子密度为13—29个／100g土壤,平均为19个;种的丰富度在4．9之间（平均为6）;平均频度为53．8％;相对多度为3．2％-26．5％;物种多样性指数和均匀度指数分别为0．94和0．93。丛枝菌根的侵染率达到44．8％．57．2％（平均为50．9％）;球囊霉属（Glomus）和无梗囊霉属（Acattlospora）是热带次生林根围土壤中菌根真菌的优势类群。     

Changes in seasonal climate outpace compensatory density‐dependence in eastern brook trout

Understanding how multiple extrinsic (density‐independent) factors and intrinsic (density‐dependent) mechanisms influence population dynamics has become increasingly urgent in the face of rapidly changing climates. It is particularly unclear how multiple extrinsic factors with contrasting effects among seasons are related to declines in population numbers and changes in mean body size and whether there is a strong role for density‐dependence. The primary goal of this study was to identify the roles of seasonal variation in climate driven environmental direct effects (mean stream flow and temperature) vs. density‐dependence on population size and mean body size in eastern brook trout (Salvelinus fontinalis). We use data from a 10‐year capture‐mark‐recapture study of eastern brook trout in four streams in Western Massachusetts, USA to parameterize a discrete‐time population projection model. The model integrates matrix modeling techniques used to characterize discrete population structures (age, habitat type, and season) with integral projection models (IPMs) that characterize demographic rates as continuous functions of organismal traits (in this case body size). Using both stochastic and deterministic analyses we show that decreases in population size are due to changes in stream flow and temperature and that these changes are larger than what can be compensated for through density‐dependent responses. We also show that the declines are due mostly to increasing mean stream temperatures decreasing the survival of the youngest age class. In contrast, increases in mean body size over the same period are the result of indirect changes in density with a lesser direct role of climate‐driven environmental change.     

A pre‐adaptive approach for tropical forest restoration during climate change using naturally occurring genetic variation in response to water limitation

Effective reforestation of degraded tropical forests depends on selecting planting material suited to the stressful environments typical at restoration sites that can be exacerbated by increased duration and intensity of dry spells expected with climate change. While reforestation efforts in nontropical systems are incorporating drought‐adapted genotypes into restoration programs to cope with drier conditions, such approaches have not been tested or implemented in tropical forests. As the first effort to examine genetic variation in plant response to drought in a tropical wet forest, we established a watering experiment using five replicated maternal lines (i.e. seedlings from different maternal trees) of five dipterocarp species native to Borneo. Apart from the expected species level variation in growth and herbivory (3‐fold variation in both cases), we also found intraspecific variation so that growth in some cases varied 2‐fold, and herbivory 3‐fold, among genetically different maternal lines. In two species we found that among‐maternal line variation in growth rate was negatively correlated with tolerance to water limitation, that is, the maternal lines that performed the best in the high water treatment lost proportionally more of their growth during water limitation. We argue that selection for tolerance to future drier conditions is not only likely to impact population genetics of entire forests, but likely extends from forest trees to the communities of canopy arthropods associated with these trees. In tropical reforestation efforts where increased drought is predicted from climate change, including plant material resilient to drier conditions may improve restoration effectiveness.     

Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest

Density-dependent seedling mortality could increase with a species relative abundance, thereby promoting species coexistence. Differences among species in light-dependent mortality also could enhance coexistence via resource partitioning. These compatible ideas rarely have been considered simultaneously. We developed models of mortality as functions of irradiance and local conspecific density (LCD) for seedlings of 53 tropical woody species. Species varied in mortality responses to these factors, but mortality consistently increased with shading and LCD. Across species, density-dependent mortality on a per-neighbour basis was inversely related to species community abundance, but higher LCD in more common species resulted in a weak relationship between species abundance and density-dependent mortality scaled to species maximum LCD. Species mortality responses to shading and maximum LCD were strongly and positively correlated. Our results suggest that species differences in density-dependent mortality are more strongly related to physiologically based life-history traits than biotic feedbacks related to community abundance.     

Ecosystem carbon storage does not vary with mean annual temperature in Hawaiian tropical montane wet forests

Theory and experiment agree that climate warming will increase carbon fluxes between terrestrial ecosystems and the atmosphere. The effect of this increased exchange on terrestrial carbon storage is less predictable, with important implications for potential feedbacks to the climate system. We quantified how increased mean annual temperature (MAT) affects ecosystem carbon storage in above‐ and belowground live biomass and detritus across a well‐constrained 5.2 °C MAT gradient in tropical montane wet forests on the Island of Hawaii. This gradient does not systematically vary in biotic or abiotic factors other than MAT (i.e. dominant vegetation, substrate type and age, soil water balance, and disturbance history), allowing us to isolate the impact of MAT on ecosystem carbon storage. Live biomass carbon did not vary predictably as a function of MAT, while detrital carbon declined by ~14 Mg of carbon ha^?1 for each 1 °C rise in temperature – a trend driven entirely by coarse woody debris and litter. The largest detrital pool, soil organic carbon, was the most stable with MAT and averaged 48% of total ecosystem carbon across the MAT gradient. Total ecosystem carbon did not vary significantly with MAT, and the distribution of ecosystem carbon between live biomass and detritus remained relatively constant across the MAT gradient at ~44% and ~56%, respectively. These findings suggest that in the absence of alterations to precipitation or disturbance regimes, the size and distribution of carbon pools in tropical montane wet forests will be less sensitive to rising MAT than predicted by ecosystem models. This article also provides needed detail on how individual carbon pools and ecosystem‐level carbon storage will respond to future warming.     

Symbiotic soil fungi enhance ecosystem resilience to climate change

Substantial amounts of nutrients are lost from soils through leaching. These losses can be environmentally damaging, causing groundwater eutrophication and also comprise an economic burden in terms of lost agricultural production. More intense precipitation events caused by climate change will likely aggravate this problem. So far it is unresolved to which extent soil biota can make ecosystems more resilient to climate change and reduce nutrient leaching losses when rainfall intensity increases. In this study, we focused on arbuscular mycorrhizal (AM) fungi, common soil fungi that form symbiotic associations with most land plants and which increase plant nutrient uptake. We hypothesized that AM fungi mitigate nutrient losses following intensive precipitation events (higher amount of precipitation and rain events frequency). To test this, we manipulated the presence of AM fungi in model grassland communities subjected to two rainfall scenarios: moderate and high rainfall intensity. The total amount of nutrients lost through leaching increased substantially with higher rainfall intensity. The presence of AM fungi reduced phosphorus losses by 50% under both rainfall scenarios and nitrogen losses by 40% under high rainfall intensity. Thus, the presence of AM fungi enhanced the nutrient interception ability of soils, and AM fungi reduced the nutrient leaching risk when rainfall intensity increases. These findings are especially relevant in areas with high rainfall intensity (e.g., such as the tropics) and for ecosystems that will experience increased rainfall due to climate change. Overall, this work demonstrates that soil biota such as AM fungi can enhance ecosystem resilience and reduce the negative impact of increased precipitation on nutrient losses.     

Nitrogen and phosphorus additions impact arbuscular mycorrhizal abundance and molecular diversity in a tropical montane forest

Increased nitrogen (N) depositions expected in the future endanger the diversity and stability of ecosystems primarily limited by N, but also often co‐limited by other nutrients like phosphorus (P). In this context a nutrient manipulation experiment (NUMEX) was set up in a tropical montane rainforest in southern Ecuador, an area identified as biodiversity hotspot. We examined impacts of elevated N and P availability on arbuscular mycorrhizal fungi (AMF), a group of obligate biotrophic plant symbionts with an important role in soil nutrient cycles. We tested the hypothesis that increased nutrient availability will reduce AMF abundance, reduce species richness and shift the AMF community toward lineages previously shown to be favored by fertilized conditions. NUMEX was designed as a full factorial randomized block design. Soil cores were taken after 2 years of nutrient additions in plots located at 2000 m above sea level. Roots were extracted and intraradical AMF abundance determined microscopically; the AMF community was analyzed by 454‐pyrosequencing targeting the large subunit rDNA. We identified 74 operational taxonomic units (OTUs) with a large proportion of Diversisporales. N additions provoked a significant decrease in intraradical abundance, whereas AMF richness was reduced significantly by N and P additions, with the strongest effect in the combined treatment (39% fewer OTUs), mainly influencing rare species. We identified a differential effect on phylogenetic groups, with Diversisporales richness mainly reduced by N additions in contrast to Glomerales highly significantly affected solely by P. Regarding AMF community structure, we observed a compositional shift when analyzing presence/absence data following P additions. In conclusion, N and P additions in this ecosystem affect AMF abundance, but especially AMF species richness; these changes might influence plant community composition and productivity and by that various ecosystem processes.     

Mechanisms driving plant functional trait variation in a tropical forest

Plant functional trait variation in tropical forests results from taxonomic differences in phylogeny and associated genetic differences, as well as, phenotypic plastic responses to the environment. Accounting for the underlying mechanisms driving plant functional trait variation is important for understanding the potential rate of change of ecosystems since trait acclimation via phenotypic plasticity is very fast compared to shifts in community composition and genetic adaptation. We here applied a statistical technique to decompose the relative roles of phenotypic plasticity, genetic adaptation, and phylogenetic constraints. We examined typically obtained plant functional traits, such as wood density, plant height, specific leaf area, leaf area, leaf thickness, leaf dry mass content, leaf nitrogen content, and leaf phosphorus content. We assumed that genetic differences in plant functional traits between species and genotypes increase with environmental heterogeneity and geographic distance, whereas trait variation due to plastic acclimation to the local environment is independent of spatial distance between sampling sites. Results suggest that most of the observed trait variation could not be explained by the measured environmental variables, thus indicating a limited potential to predict individual plant traits from commonly assessed parameters. However, we found a difference in the response of plant functional traits, such that leaf traits varied in response to canopy‐light regime and nutrient availability, whereas wood traits were related to topoedaphic factors and water availability. Our analysis furthermore revealed differences in the functional response of coexisting neotropical tree species, which suggests that endemic species with conservative ecological strategies might be especially prone to competitive exclusion under projected climate change.     

Environmental controls on net ecosystem-level carbon exchange and productivity in a Central American tropical wet forest

Difficulty in balancing the global carbon budget has lead to increased attention on tropical forests, which have been estimated to account for up to one third of global gross primary production. Whether tropical forests are sources, sinks, or neutral with respect to their carbon balance with the atmosphere remains unclear. To address this issue, estimates of net ecosystem exchange of carbon (NEE) were made for 3 years (1998–2000) using the eddy‐covariance technique in a tropical wet forest in Costa Rica. Measurements were made from a 42 m tower centred in an old‐growth forest. Under unstable conditions, the measurement height was at least twice the estimated zeroplane height from the ground. The canopy at the site is extremely rough; under unstable conditions the median aerodynamic roughness length ranged from 2.4 to 3.6 m. No relationship between NEE and friction velocity (u*) was found using all of the 30‐min averages. However, there was a linear relationship between the nighttime NEE and averaged u* (R² = 0.98). The diurnal pattern of flux was similar to that found in other tropical forests, with mean daytime NEE ca. ? 18 μ mol CO₂ m^?2 s^?1 and mean nighttime NEE 4.6 μ mol CO₂ m^?2 s^?1. However, because ～ 80% of the nighttime data in this forest were collected during low u* conditions ( < 0.2 m s^?1), nighttime NEE was likely underestimated. Using an alternative analysis, mean nighttime NEE increased to 7.05 μ mol CO₂ m^?2 s^?1. There were interannual differences in NEE, but seasonal differences were not apparent. Irradiance accounted for ～ 51% of the variation in the daytime fluxes, with temperature and vapour pressure deficit together accounting for another ～ 20%. Light compensation points ranged from 100 to 207 μ mol PPFD m^?2 s^?1. No was relationship was found between 30‐min nighttime NEE and tower‐top air temperature. A weak relationship was found between hourly nighttime NEE and canopy air temperature using data averaged hourly over the entire sampling period (Q₁₀ = 1.79, R² = 0.17). The contribution of below‐sensor storage was fairly constant from day to day. Our data indicate that this forest was a slight carbon source in 1998 (0.05 to ?1.33 t C ha^?1 yr^?1), a moderate sink in 1999 (?1.53 to ?3.14 t C ha^?1 yr^?1), and a strong sink in 2000 (?5.97 to ?7.92 t C ha^?1 yr^?1). This trend is interpreted as relating to the dissipation of warm‐phase El Niño effects over the course of this study.     

西双版纳四种壳斗科植物丛枝菌根状况的初步研究

研究了西双版纳热带次生林中杯丝锥(Castanopsis calathiformis)、红锥(C.hys-trix)、印度锥(C.indica)和截果柯(Lithocarpus truncatus)4种壳斗科植物的丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)的侵染状况,并从这些植物的根际土壤中分离鉴定了隶属于球囊霉属(Glomus)、巨孢囊霉属(Gigaspora)、盾巨孢囊霉属(Scutellospora)和无梗囊霉属(Acaulospora)的10种丛枝菌根真菌。这4种壳斗科植物根际AMF的孢子密度为14~22个.100g-1土壤,种的丰富度在4~7,平均频度为60.00%,相对多度为4.41%~22.06%,丛枝菌根真菌的定居水平达46.26%~51.40%。     

The impact of global climate change on tropical forest biodiversity in Amazonia

Aim To model long‐term trends in plant species distributions in response to predicted changes in global climate. Location Amazonia. Methods The impacts of expected global climate change on the potential and realized distributions of a representative sample of 69 individual Angiosperm species in Amazonia were simulated from 1990 to 2095. The climate trend followed the HADCM2GSa1 scenario, which assumes an annual 1% increase of atmospheric CO₂ content with effects mitigated by sulphate forcing. Potential distributions of species in one‐degree grid cells were modelled using a suitability index and rectilinear envelope based on bioclimate variables. Realized distributions were additionally limited by spatial contiguity with, and proximity to, known record sites. A size‐structured population model was simulated for each cell in the realized distributions to allow for lags in response to climate change, but dispersal was not included. Results In the resulting simulations, 43% of all species became non‐viable by 2095 because their potential distributions had changed drastically, but there was little change in the realized distributions of most species, owing to delays in population responses. Widely distributed species with high tolerance to environmental variation exhibited the least response to climate change, and species with narrow ranges and short generation times the greatest. Climate changed most in north‐east Amazonia while the best remaining conditions for lowland moist forest species were in western Amazonia. Main conclusions To maintain the greatest resilience of Amazonian biodiversity to climate change as modelled by HADCM2GSa1, highest priority should be given to strengthening and extending protected areas in western Amazonia that encompass lowland and montane forests.     

Changes in seasonal precipitation distribution but not annual amount affect litter decomposition in a secondary tropical forest

In the tropics of South China, climate change induced more rainfall events in the wet season in the last decades. Moreover, there will be more frequently spring drought in the future. However, knowledge on how litter decomposition rate would respond to these seasonal precipitation changes is still limited. In the present study, we conducted a precipitation manipulation experiment in a tropical forest. First, we applied a 60% rainfall exclusion in April and May to defer the onset of wet season and added the same amount of water in October and November to mimic a deferred wet season (DW); second, we increased as much as 25% mean annual precipitation into plots in July and August to simulate a wetter wet season (WW). Five single‐species litters, with their carbon to nitrogen ratio ranged from 27 to 49, and a mixed litter were used to explore how the precipitation change treatments would affect litter decomposition rate. The interaction between precipitation changes and litter species was not significant. The DW treatment marginally accelerated litter decomposition across six litter types. Detailed analysis showed that DW increased litter decomposition rate in the periods of January to March and October to December, when soil moisture was increased by the water addition in the dry season. In contrast, WW did not significantly affect litter decomposition rate, which was consistent with the unchanged soil moisture pattern. In conclusion, the study indicated that regardless of litter types or litter quality, the projected deferred wet season would increase litter decomposition rate, whereas the wetter wet season would not affect litter decomposition rate in the tropical forests. This study improves our knowledge of how tropical forest carbon cycling in response to precipitation change.     

The role of tropical dry forest as a long-term barrier to dispersal: a comparative phylogeographical analysis of dry forest tolerant and intolerant frogs

We used a comparative phylogeographical approach to investigate the origins of the disjunct wet forest biota of the Golfo Dulce region along the Pacific slope of Costa Rica. This region is isolated by Pacific dry forests north and south and isolated from Caribbean wet forests by mountains. We studied three sympatric lowland frog species in the Craugastor fitzingeri species group that prefer wet forest but differ in their response to dry habitats. In dry forest, C. fitzingeri can survive along streams while C. crassidigitus and C. talamancae are entirely absent. We collected samples from across the ranges of all three species, and obtained mitochondrial DNA sequence data from the COI and cytochrome b genes. We observed significant phylogeographical structure in C. crassidigitus and C. talamancae, but much less in C. fitzingeri, demonstrating that mountain barriers and dry forest habitat have reduced mitochondrial gene flow in the strictly wet-forest species. Additionally, we discovered that the Golfo Dulce and Central Panama populations of C. crassidigitus appear to have diverged in the Pliocene or earlier, suggesting that the dry forest separating these populations is old. Our phylogenetic analysis of 12 of approximately 16 species of the C. fitzingeri species group suggests that the three lowland species are each other's closest relatives. Because of this shared phylogenetic history, we attribute the striking differences in phylogeographical structure to the different ecologies of the frogs. In summary, we find that what appear to be minor differences in the natural history of these three closely related species may profoundly impact the potential for dispersal, range size, and cladogenesis.     

Disentangling the long‐term effects of disturbance on soil biogeochemistry in a wet tropical forest ecosystem

Climate change is increasing the intensity of severe tropical storms and cyclones (also referred to as hurricanes or typhoons), with major implications for tropical forest structure and function. These changes in disturbance regime are likely to play an important role in regulating ecosystem carbon (C) and nutrient dynamics in tropical and subtropical forests. Canopy opening and debris deposition resulting from severe storms have complex and interacting effects on ecosystem biogeochemistry. Disentangling these complex effects will be critical to better understand the long‐term implications of climate change on ecosystem C and nutrient dynamics. In this study, we used a well‐replicated, long‐term (10 years) canopy and debris manipulation experiment in a wet tropical forest to determine the separate and combined effects of canopy opening and debris deposition on soil C and nutrients throughout the soil profile (1 m). Debris deposition alone resulted in higher soil C and N concentrations, both at the surface (0–10 cm) and at depth (50–80 cm). Concentrations of NaOH‐organic P also increased significantly in the debris deposition only treatment (20–90 cm depth), as did NaOH‐total P (20–50 cm depth). Canopy opening, both with and without debris deposition, significantly increased NaOH‐inorganic P concentrations from 70 to 90 cm depth. Soil iron concentrations were a strong predictor of both C and P patterns throughout the soil profile. Our results demonstrate that both surface‐ and subsoils have the potential to significantly increase C and nutrient storage a decade after the sudden deposition of disturbance‐related organic debris. Our results also show that these effects may be partially offset by rapid decomposition and decreases in litterfall associated with canopy opening. The significant effects of debris deposition on soil C and nutrient concentrations at depth (>50 cm), suggest that deep soils are more dynamic than previously believed, and can serve as sinks of C and nutrients derived from disturbance‐induced pulses of organic matter inputs.     

Improving predictions of tropical forest response to climate change through integration of field studies and ecosystem modeling

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species‐specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model‐data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate.