Infrared heater system for warming tropical forest understory plants and soils

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.     

Nitrogen addition reduces soil respiration in a mature tropical forest in southern China

Response of soil respiration (CO₂ emission) to simulated nitrogen (N) deposition in a mature tropical forest in southern China was studied from October 2005 to September 2006. The objective was to test the hypothesis that N addition would reduce soil respiration in N saturated tropical forests. Static chamber and gas chromatography techniques were used to quantify the soil respiration, following four‐levels of N treatments (Control, no N addition; Low‐N, 5 g N m^?2 yr^?1; Medium‐N, 10 g N m^?2 yr^?1; and High‐N, 15 g N m^?2 yr^?1 experimental inputs), which had been applied for 26 months before and continued throughout the respiration measurement period. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates found in the warm and wet growing season (April–September) and the lowest rates in the dry dormant season (December–February). Soil respiration rates showed a significant positive exponential relationship with soil temperature, whereas soil moisture only affect soil respiration at dry conditions in the dormant season. Annual accumulative soil respiration was 601±30 g CO₂‐C m^?2 yr^?1 in the Controls. Annual mean soil respiration rate in the Control, Low‐N and Medium‐N treatments (69±3, 72±3 and 63±1 mg CO₂‐C m^?2 h^?1, respectively) did not differ significantly, whereas it was 14% lower in the High‐N treatment (58±3 mg CO₂‐C m^?2 h^?1) compared with the Control treatment, also the temperature sensitivity of respiration, Q₁₀ was reduced from 2.6 in the Control with 2.2 in the High‐N treatment. The decrease in soil respiration occurred in the warm and wet growing season and were correlated with a decrease in soil microbial activities and in fine root biomass in the N‐treated plots. Our results suggest that response of soil respiration to atmospheric N deposition in tropical forests is a decline, but it may vary depending on the rate of N deposition.     

Leaf nonstructural carbohydrate concentrations of understory woody species regulated by soil phosphorus availability in a tropical forest

Leaf soluble sugars and starch are important components of nonstructural carbohydrates (NSCs), which are crucial for plant growth, development, and reproduction. Although there is a large body of research focusing on the regulation of plant NSC (soluble sugars and starch) concentrations, the response of foliar NSC concentrations to continuous nitrogen (N) and phosphorus (P) addition is still unclear, especially in tropical forests. Here, we used a long‐term manipulative field experiment to investigate the response of leaf NSC concentrations to continuous N and P addition (3‐, 5‐, and 8‐year fertilization) in a tropical forest in southern China. We found significant species‐specific variation in leaf NSC concentrations in this tropical forest. Phosphorus addition dramatically decreased both leaf soluble sugar and starch concentrations, while N addition had no significant effects on leaf soluble sugar and starch concentrations. These results suggest that, in plants growing in P‐limiting tropical soil, leaf NSC concentrations are regulated by soil P availability rather than N availability. Moreover, the negative relationships between NSC concentrations and leaf mass per area (LMA) revealed that NSCs could supply excess carbon (C) for leaf expansion under P addition. This was further supported by the increased structural P fraction after P fertilization in our previous study at the same site. We conclude that soil P availability strongly regulates leaf starch and soluble sugar concentrations in the tropical tree species included in this study. The response of leaf NSC concentrations to long‐term N and P addition can reflect the close relationships between plant C dynamics and soil nutrient availability in tropical forests. Maintaining relatively higher leaf NSC concentrations in tropical plants can be a potential mechanism for adapting to P‐deficient conditions.     

High species diversity in fleshy-fruited tropical understory plants

Key innovations may increase the number of taxa in a clade that possesses the proposed innovation in comparison to its sister group that lacks the trait through either increased speciation or reduced extinction rates. Comparing sister clades across several independent lineages provides statistical support that the trait has increased species diversity. Previous studies have indicated that there may not be a relationship between biotic dispersal and higher species diversity, but only a few of these studies specified habit, habitat, or type of disperser. No previous study has specified all of the above parameters and used a phylogenetic approach. This article examines species diversity in numerous lineages of tropical understory plants with small, fleshy, bird-dispersed fruits for which a reliable estimate of phylogenetic relationships is available. Clades with fleshy fruits are significantly more diverse than sister clades with dry fruits.     

Nitrogen conservation in a tropical rain forest

Summary A series of plots in an Amazonian Rain Forest were trenched and treated with calcium to determine the effects of perturbation on numbers and activity of nitrifying bacteria. Although treatment resulted in between 2 and 26% of the nitrogen being lost from the humus layer, virtually all of it was in the NH ₄ ⁺ form. Numbers of nitrifying bacteria in the plots were relatively low. The low numbers and low activity of nitrifying bacteria is attributed to the low pH and high concentration of tannins in the root mat. It is hypothesized that the suppression of nitrifying bacteria results in nitrogen conservation in the rain forest.     

The effects of nitrogen addition on the growth of two exotic and two native forest understory plants

Many studies have linked the spread of exotic, invasive species to high nitrogen supply, but most of this work does not distinguish between various inorganic forms and different concentrations of available nitrogen. Previous research has suggested that exotic, invasive species common in eastern deciduous forests may preferentially utilize nitrate in contrast to native species that preferentially make use of ammonium. We tested this hypothesis by comparing the growth response of two common forest invaders, Berberis thunbergii and Microstegium vimineum, and two co-occurring native species (Vaccinium pallidum and Hamamelis virginiana) to different forms of nitrogen under varying concentrations in a greenhouse experiment. Two forms of nitrogen (nitrate or ammonium) were added at different concentrations (22, 106, and 212 mg N l⁻¹) to all species. Growth response variables included survival, stem length, stem diameter, above and belowground biomass, and estimated seed production (Microstegium only). Unexpectedly the exotic species did not respond preferentially to nitrate addition. Microstegium responded most strongly to both nitrogen forms depending on the response variable. Berberis and Hamamelis surprisingly reacted similarly to nitrogen additions. As expected, Vaccinium fared poorly under most treatment conditions, but did show some growth in NH₄ ⁺ treatments. Our findings suggest species response to nitrogen addition is complex, and that exotic species do not all respond similarly to nitrogen inputs. In this study, the response of exotic and native species to available nutrients does not provide a general mechanism of invasion success.     

东北天然次生林下木树种的生物量器官分配规律

以帽儿山天然次生林主要下木树种为研究对象,探讨了植物生物量器官(叶、新枝、多年枝、细根、粗根)分配特征及其与物种和个体大小的关系。结果表明:1)林下植物器官生物量的相对生长遵循异速生长理论,相对生长关系并不唯一。叶与新枝(0.924～1.055)、多年枝与粗根(0.917～1.024)和地上部分与地下部分(1.064～1.125)近于等速生长,新枝与多年枝(0.585～0.700),叶与总枝(0.742～0.795),叶与总根(0.853～0.918),以及细根与粗根(0.658～0.750)的生物量相对生长表现为异速生长。2)林下植物生物量器官分配遵循异速生长分配理论,叶、新枝、多年枝、细根和粗根等的生物量比例依次是5.83%～20.60%、0.83%～7.42%、36.25%～68.24%、1.32%～6.75%和16.38%～42.88%、根冠比是0.272～0.866。3)器官生物量比例与物种和植物大小等有关,各植物的器官生物量比例有差异,随植物生长,各植物生物量的叶分配、新枝分配、细根分配和根冠比明显减小、多年枝分配明显增加(P0.05),仅少数植物的粗根分配无明显变化(P0.05)。     

Convergence in light capture efficiencies among tropical forest understory plants with contrasting crown architectures: a case of morphological compensation

Leaf and crown characteristics were examined for 24 tree and herbaceous species of contrasting architectures from the understory of a lowland rainforest. Light-capture efficiency was estimated for the crowns of the different species with a three-dimensional geometric modeling program. Causal relationships among traits affecting light absorption at two hierarchical levels (leaf and whole crown) were quantified using path analysis. Light-capture and foliage display efficiency were found to be very similar among the 24 species studied, with most converging on a narrow range of light absorption efficiencies (ratio of absorbed vs. available light of 0.60-0.75). Exceptionally low values were found for the climber vines and, to a lesser extent, for the Bromeliad Aechmea magdalenae. Differences in photosynthetic photon flux density (PFD) absorbed per unit leaf area by individual plants were mostly determined by site to site variation in PFD and not by the differences in crown architecture among individuals or species. Leaf angle, and to a lesser extent also supporting biomass, specific leaf area, and internode length, had a significant effect on foliage display efficiency. Potential constraints on light capture such as the phyllotactic pattern were generally offset by other compensatory adjustments of crown structure such as internode length, arching stems, and plagiotropy. The variety of shoot morphologies capable of efficiently capturing light in tropical forest understories is greater than initially thought, extending over species with very different phyllotactic patterns, crown architectures, leaf sizes, and morphologies.     

Xanthophyll-cycle pigments and photosynthetic capacity in tropical forest species: a comparative field study on canopy,gap and understory plants

Xanthophyll-cycle pigments and photosynthetic capacity (PS_max) were analyzed in 25 species from different light environments (canopy, gap, understory) within a Panamanian tropical forest. (1) Sun-exposed leaves of canopy tree species showed the highest photosynthetic capacities and largest xanthophyll-cycle pools (violaxanthin, antheraxanthin, zeaxanthin) of about 87 mmol mol^-1 chlorophyll with only small amounts of -carotene [about 7 mmol mol^-1 chlorophyll = 8% of total (+) carotene pool]. Under high natural photon flux densities (PFDs) canopy leaves rapidly converted up to 96% of the xanthophyll-cycle pool into zeaxanthin. The back reaction to violaxanthin occurred much faster in low light than in complete darkness. At the end of the night, zeaxanthin still accounted for, on average, 14% of the total xanthophyll-cycle pigments. (2) Leaves of gap plants had intermediate values of PS_max and a 43% lower total carotenoid content than canopy leaves. The average size of the xanthophyll-cycle pool was 35 mmol mol^-1 chlorophyll, and -carotene accounted for up to 66% of the total (+) carotene pool. Under high light conditions gap plants converted, on average, 86% of the xanthophyll-cycle pigments into zeaxanthin. The back reaction, following a decrease in ambient PFD, was slower than the forward reaction. At the end of the night, zeaxanthin accounted for, on average, 7% of the xanthophyll-cycle pigments in gap plants. (3) Understory plants showed the lowest values of PS_max and the smallest xanthophyll-cycle pool of about 22 mmol mol^-1 chlorophyll. -Carotene accounted for up to 70% of total carotene. The conversion of xanthophyll-cycle pigments into zeaxanthin was negligible during short sunflecks of 1–2 min duration and PFDs up to about 400 mol m^-2 s^-1. At predawn, leaves of understory plants rarely contained any detectable zeaxanthin. Aechmea magdalenae, an understory CAM plant, showed exceptionally high rates of PS_max per unit leaf area compared to sympatric C₃ understory species.     

Nitrogen addition promotes terrestrial plants to allocate more biomass to aboveground organs: A global meta-analysis

A significant increase in reactive nitrogen (N) added to terrestrial ecosystems through agricultural fertilization or atmospheric deposition is considered to be one of the most widespread drivers of global change. Modifying biomass allocation is one primary strategy for maximizing plant growth rate, survival, and adaptability to various biotic and abiotic stresses. However, there is much uncertainty as to whether and how plant biomass allocation strategies change in response to increased N inputs in terrestrial ecosystems. Here, we synthesized 3516 paired observations of plant biomass and their components related to N additions across terrestrial ecosystems worldwide. Our meta-analysis reveals that N addition (ranging from 1.08 to 113.81 g m⁻² year⁻¹) increased terrestrial plant biomass by 55.6% on average. N addition has increased plant stem mass fraction, shoot mass fraction, and leaf mass fraction by 13.8%, 12.9%, and 13.4%, respectively, but with an associated decrease in plant reproductive mass (including flower and fruit biomass) fraction by 3.4%. We further documented a reduction in plant root-shoot ratio and root mass fraction by 27% (21.8%–32.1%) and 14.7% (11.6%–17.8%), respectively, in response to N addition. Meta-regression results showed that N addition effects on plant biomass were positively correlated with mean annual temperature, soil available phosphorus, soil total potassium, specific leaf area, and leaf area per plant. Nevertheless, they were negatively correlated with soil total N, leaf carbon/N ratio, leaf carbon and N content per leaf area, as well as the amount and duration of N addition. In summary, our meta-analysis suggests that N addition may alter terrestrial plant biomass allocation strategies, leading to more biomass being allocated to aboveground organs than belowground organs and growth versus reproductive trade-offs. At the global scale, leaf functional traits may dictate how plant species change their biomass allocation pattern in response to N addition.     

Treefall gaps versus forest understory as environments for a defoliating moth on a tropical forest shrub

Summary The moth Zunacetha annulata (Dioptidae) is a specialist on the understory shrub Hybanthus prunifolius (Violaceae) in the forest of Barro Colorado Island (BCI), Panama. The larvae, which are capable of defoliating entire shrubs, concentrate their attack upon the small minority of H. prunifolius individuals that grow in treefall gaps. Field experiments demonstrated that larval growth rates were 37% higher, and weights at pupation 25% higher, on shrubs in gaps than on shrubs in the understory. In a common environment in the laboratory, growth rates of larvae were 23% higher on foliage taken from shrubs in gaps than on foliage from shrubs in the understory.However, larvae grown in a temperature regime simulating that of gaps did not grow faster than larvae in an understory regime, when the two groups were reared in growth chambers on foliage taken from the same shrubs. In the field, predation appeared higher in gaps: experimental groups of larvae survived at rates of 65% per day in gaps and 78% per day in the understory. Quality of foliage, and not direct effects of the environment, appears to be responsible for the observed pattern of defoliation by this moth.     

Functional characteristics of corticolous lichens in the understory of a tropical lowland rain forest

In tropical lowland forests, corticolous crustose green algal lichens are abundant and highly diverse. This may be related to adaptation to prevailing microenvironmental conditions including, for example, high precipitation and low light intensities. In the understory of a tropical lowland rain forest in French Guiana, we studied the morphology of crustose green algal lichens and measured gas exchange and chlorophyll a fluorescence. We found that (i) periods of thallus suprasaturation with water were reduced by the presence of water-repelling surface structures of mycobiont hyphae at the thallus surface and the medulla; (ii) photosynthesis was adapted to the low light intensities present in the understory; and (iii) photosynthesis was rapidly activated in fluctuating light. The combination of these three mechanisms enables corticolous lichens to implement specific morphological and physiological strategies, which may favour growth in the limiting understory habitat of tropical lowland rain forests.     

Phosphorus limitation on photosynthesis of two dominant understory species in a lowland tropical forest

Aims Elevated nitrogen (N) deposition in tropical regions may accelerate ecosystem phosphorus (P) limitation. However, it is not explicitly addressed that how changes in soil N and P availability affect foliar nutrients and photosynthesis of plants in tropical forests. In this study, we examined the effects of N and P additions on foliar nutrients and net photosynthesis of two dominant understory species, Randia canthioides (R. canthioides) and Cryptocarya concinna (C. concinna) in an N-saturated old-growth tropical forest (>400-year-old) in southern China.Methods A full factorial NP addition experiment (2×2) was established in 2007 and continued through August 2010. Four treatments, including control, N addition (150kg N ha^-1 year^-1), P addition (150kg P ha^-1 year^-1) and NP addition (150kg N ha^-1 year^-1 plus 150kg P ha^-1 year^-1) were set up in this experiment. Photosynthetic traits (maximum photosynthetic CO₂ assimilation (A max), stomatal conductance (g s), leaf transpiration (E), light saturating point, concentrations of chlorophyll a/b and foliar nutrients (N and P) of the two species were measured with standard methods.Important findings Three years of N addition had no significant effects on any measured photosynthetic parameter of either species. However, N addition significantly elevated foliar N and P concentrations of one species (R. canthioides), resulting in lower photosynthetic nitrogen use efficiency (PNUE). N treatments decreased foliar P concentration of the other (C. concinna), resulting in increased photosynthetic phosphorus use efficiency, which was potentially related to N-induced P shortage. In contrast, positive effects of P treatments on g s of R. canthioides, A max and chlorophyll a+b of C. concinna were observed. P treatments also elevated foliar P and PNUE of both species, implying P induced more efficient use of N. Our results suggested a more important role of P than N on influencing photosynthetic traits of these two understory species. Alleviation of P shortage through P addition may enhance photosynthetic performances of some understory species in N-rich tropical forests.     

Nitrogen uptake and preference in a forest understory following invasion by an exotic grass

Plant–soil interactions have been proposed as a causative mechanism explaining how invasive plant species impact ecosystem processes. We evaluate whether an invasive plant influences plant and soil-microbe acquisition of nitrogen to elucidate the mechanistic pathways by which invaders might alter N availability. Using a ¹⁵N tracer, we quantify differences in nitrogen uptake and allocation in communities with and without Microstegium vimineum, a shade-tolerant, C₄ grass that is rapidly invading the understories of eastern US deciduous forests. We further investigate if plants or the microbial biomass exhibit preferences for certain nitrogen forms (glycine, nitrate, and ammonium) to gain insight into nitrogen partitioning in invaded communities. Understory native plants and M. vimineum took up similar amounts of added nitrogen but allocated it differently, with native plants allocating primarily to roots and M. vimineum allocating most nitrogen to shoots. Plant nitrogen uptake was higher in invaded communities due primarily to the increase in understory biomass when M. vimineum was present, but for the microbial biomass, nitrogen uptake did not vary with invasion status. This translated to a significant reduction (P < 0.001) in the ratio of microbial biomass to plant biomass nitrogen uptake, which suggests that, although the demand for nitrogen has intensified, microbes continue to be effective nitrogen competitors. The microbial biomass exhibited a strong preference for ammonium over glycine and nitrate, regardless of invasion status. By comparison, native plants showed no nitrogen preferences and M. vimineum preferred inorganic nitrogen species. We interpret our findings as evidence that invasion by M. vimineum leads to changes in the partitioning of nitrogen above and belowground in forest understories, and to decreases in the microbial biomass, but it does not affect the outcome of plant–microbe–nitrogen interactions, possibly due to functional shifts in the microbial community as a result of invasion.     

Diversity and seasonal variation of ground and understory spiders from a tropical mountain cloud forest

We made intensive samplings to study the seasonal response of spiders across different forest strata(ground and understory)in a tropical mountain cloud forest from Mexico.We sampled spiders from ten plots in six sampling events during the dry and rainy season,to analyze their abundance,structure(distribution of abundance among species),diversity and the response of the five dominant species at each stratum.Results demonstrated that seasonal patterns of spider communities differed among strata,revealing a complex spatiotemporal dynamic.Abundance,structure,diversity of ground spiders,as well as the responses of four dominant species at this stratum,showed low seasonal variations.In contrast,a strong seasonal variation was observed for the understory assemblage,with lowest abundance and highest diversity in the rainy season,and different assemblage structures for each season.Seasonal patterns of each assemblage seem linked to the responses of their dominant species.We found high co-occurrence among most of the ground dominant species with similar habitat use and with multivoltine patterns,contrasting with low co-occurrence among most of the understory dominant species with similar habitat use and univoltine patterns.Our results showed that the spiders’assemblages of tropical mountain cloud forest(opposed to what is found in temperate and boreal forests)increase their species richness with the height,and that their responses to seasonal change differ between strata.Management programs of these habitats should consider the spatial and temporal variations found here,as a better understanding of their ecological dynamics is required to support their sustainable management.     

Spatial distribution of understory fruit-eating birds and fruiting plants in a neotropical lowland wet forest

Spatial distribution of fruit-eating birds and fruiting shrubs of the Melastomataceae and Rubiaceae were examined on a 10 ha plot in tropical lowland wet forest of Costa Rica. Many plant species and most birds exhibited considerable spatial variation in their occurrence on the plot, as indicated by the distribution patterns of shrubs with ripe fruits and captures in mist nets, respectively. In many cases, captures of fruit-eating birds were correlated with abundance of fruiting plants, particularly for species that rely heavily on fruits. In general, fruit-eating birds concentrated their use of the plot to areas rich in fruiting shrubs. This differential use of certain areas likely results in differential visitation to fruiting plants located in these areas and in a heterogeneous dissemination of seeds into the habitat.     

Effectiveness of acoustic lures for increasing tropical forest understory bat captures

Bats are the most diverse mammalian order second to rodents, with 1400+ species globally. In the tropics, it is possible to find more than 60 bat species at a single site. However, monitoring bats is challenging due to their small size, ability to fly, cryptic nature, and nocturnal activity. Recently, bioacoustic techniques have been incorporated into survey methods, either through passive acoustic monitoring or acoustic bat lures. Lures have been developed on the premise that broadcasting acoustic stimuli increases the number of captures in harp traps or mist nets. However, this is a relatively new, niche method. This study tested the efficacy of two commonly used acoustic bat lure devices, broadcasting two different acoustic stimuli, to increase forest understory bat captures in the tropics. This is the first time an acoustic bat lure has been systematically tested in a tropical rainforest, and the first study to compare two lure devices (Sussex AutoBat and Apodemus BatLure). Using a paired experimental design, two synthesized acoustic stimuli were broadcasted, a feeding call and a social call, to understand the importance of the call type used on capture rates and genus‐specific responses. Using an acoustic lure significantly increased capture rates, while the type of device did not impact capture rates. The two acoustic stimuli had an almost even distribution of captures, suggesting that the type of call may be less important than previously thought. Results indicate a possible deterrent effect on Rhinolophous sp., while being particularly effective for attracting bats in the genera Murina and Kerivoula. This study highlights the effectiveness of lures, however, also indicates that lure effects can vary across genera. Therefore, lures may bias survey results by altering the species composition of bats caught. Future research should focus on a single species or genus, using synthesized calls of conspecifics, to fully understand the effect of lures.