Convergent elevation trends in canopy chemical traits of tropical forests

The functional biogeography of tropical forests is expressed in foliar chemicals that are key physiologically based predictors of plant adaptation to changing environmental conditions including climate. However, understanding the degree to which environmental filters sort the canopy chemical characteristics of forest canopies remains a challenge. Here, we report on the elevation and soil‐type dependence of forest canopy chemistry among 75 compositionally and environmentally distinct forests in nine regions, with a total of 7819 individual trees representing 3246 species collected, identified and assayed for foliar traits. We assessed whether there are consistent relationships between canopy chemical traits and both elevation and soil type, and evaluated the general role of phylogeny in mediating patterns of canopy traits within and across communities. Chemical trait variation and partitioning suggested a general model based on four interconnected findings. First, geographic variation at the soil‐Order level, expressing broad changes in fertility, underpins major shifts in foliar phosphorus (P) and calcium (Ca). Second, elevation‐dependent shifts in average community leaf dry mass per area (LMA), chlorophyll, and carbon allocation (including nonstructural carbohydrates) are most strongly correlated with changes in foliar Ca. Third, chemical diversity within communities is driven by differences between species rather than by plasticity within species. Finally, elevation‐ and soil‐dependent changes in N, LMA and leaf carbon allocation are mediated by canopy compositional turnover, whereas foliar P and Ca are driven more by changes in site conditions than by phylogeny. Our findings have broad implications for understanding the global ecology of humid tropical forests, and their functional responses to changing climate.     

Responses of belowground communities to large aboveground herbivores: Meta‐analysis reveals biome‐dependent patterns and critical research gaps

The importance of herbivore–plant and soil biota–plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta‐analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root‐feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.     

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.     

More is not necessarily better: the impact of limiting and excessive nutrients on herbivore population growth rates

Abstract 1. The body tissues of insect herbivores contain higher concentrations of nitrogen and phosphorus than do their host plants, leading to an elemental mismatch that can limit herbivore growth, fecundity and ultimately influence population dynamics. While low nutrient availability can limit herbivore growth and reproduction, nutrient levels that exceed an organism’s nutritional requirements, i.e. an organisms’ threshold elemental ratio, can also decrease performance. 2. We conducted a laboratory experiment to examine the impacts of nitrogen and phosphorus additions on population growth rates of a phloem‐feeding insect herbivore. 3. Herbivore per capita population growth rates were highest at intermediate foliar nitrogen concentrations, indicating a performance cost on the highest nitrogen foliage. While there was no direct effect of foliar phosphorus concentration on insect performance, there was a strong and unexpected indirect effect. High soil phosphorus availability increased both foliar nitrogen concentrations and aphid tissue nitrogen, resulting in low population growth rates when both soil nitrogen and phosphorus availabilities were high. 4. In this study, experimental increases in foliar nitrogen levels led to a decrease in herbivore performance suggesting that excessive nutrient levels can limit herbivore population growth rates.     

Phytochemical diversity impacts herbivory in a tropical rainforest tree community

Metabolomics provides an unprecedented window into diverse plant secondary metabolites that represent a potentially critical niche dimension in tropical forests underlying species coexistence. Here, we used untargeted metabolomics to evaluate chemical composition of 358 tree species and its relationship with phylogeny and variation in light environment, soil nutrients, and insect herbivore leaf damage in a tropical rainforest plot. We report no phylogenetic signal in most compound classes, indicating rapid diversification in tree metabolomes. We found that locally co-occurring species were more chemically dissimilar than random and that local chemical dispersion and metabolite diversity were associated with lower herbivory, especially that of specialist insect herbivores. Our results highlight the role of secondary metabolites in mediating plant–herbivore interactions and their potential to facilitate niche differentiation in a manner that contributes to species coexistence. Furthermore, our findings suggest that specialist herbivore pressure is an important mechanism promoting phytochemical diversity in tropical forests.     

Herbivore attack in Casearia nitida influenced by plant ontogenetic variation in foliage quality and plant architecture

Traits influencing plant quality as food and/or shelter for herbivores may change during plant ontogeny, and as a consequence, influence the amount of herbivory that plants receive as they develop. In this study, differences in herbivore density and herbivory were evaluated for two ontogenetic stages of the tropical tree Casearia nitida. To assess plant ontogenetic differences in foliage quality as food for herbivores, nutritional and defensive traits were evaluated in saplings and reproductive trees. Predatory arthropods were quantified and the foraging preferences of a parasitoid wasp of the genus Zacremnops were assessed. In addition, survival rates of lepidopteran herbivores (Geometridae) were evaluated experimentally. Herbivore density was three times higher and herbivory was 66% greater in saplings than in reproductive trees. Accordingly, concentrations of total foliar phenolics were higher in reproductive trees than in saplings, whereas leaf toughness, water and nitrogen concentration did not vary between ontogenetic stages. Survival rates of lepidopteran larvae exposed to natural enemies were equivalent in reproductive trees and saplings. Given the greater herbivore density on saplings, equal survival rates implied a greater foraging effort of predators on reproductive trees. Furthermore, observed foraging of parasitoid wasps was restricted to reproductive trees. I propose that herbivore density, and as a consequence, leaf damage were lower in reproductive trees than in saplings due to both traits influencing food quality, and architectural or unmeasured indirect defensive traits influencing foraging preference of natural enemies of herbivores.     

A Meta-analysis of Climatic and Chemical Controls on Leaf Litter Decay Rates in Tropical Forests

Although tropical forests occupy a small fraction of the earth’s total land area, they play a disproportionately large role in regulating the global carbon cycle. Yet controls on both primary productivity and decomposition in tropical forests are not well-studied in comparison with temperate forests and grasslands, despite their extreme biogeochemical heterogeneity. To evaluate the relative importance of climate and foliar chemical variables in driving decomposition in tropical forests, I performed a meta-analysis of reported leaf litter decay rates throughout tropical forest ecosystems. Using a model selection procedure based on Akaike’s Information Criterion, I found that temperature and precipitation played little direct role in regulating decomposition rates, except in montane forests where cool temperatures slowed decay. Foliar concentrations of calcium, magnesium, nitrogen, phosphorus, and potassium were important predictors of mass loss rates, although each of these factors explained a very small amount of variance when considered in isolation. The large amount of unexplained variation in decomposition rates observed both within and across tropical forest sites may be due to other factors not explored here, such as soil biota or complex plant secondary chemistry. Carbon cycling in tropical forests seems to be modulated by the availability of multiple nutrients, underscoring the need for additional manipulative experiments to explore patterns of belowground nutrient limitation across the biome. Because models of decomposition developed in temperate ecosystems do not appear to be generalizable to wet tropical forests, new biogeochemical paradigms should be developed to accommodate their unique combination of climatic, edaphic, and biotic factors.     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Expansion of gallery forests into central Brazilian savannas

Upland tropical forests have expanded and contracted in response to past climates, but it is not clear whether similar dynamics were exhibited by gallery (riparian) forests within savanna biomes. Because such forests generally have access to ample water, their extent may be buffered against changing climates. We tested the long‐term stability of gallery forest boundaries by characterizing the border between gallery forests and savannas and tracing the presence of gallery forest through isotopic analysis of organic carbon in the soil profile. We measured leaf area index, grass vs. shrub or tree coverage, the organic carbon, phosphorus, nitrogen and calcium concentrations in soils and the carbon isotope ratios of soil organic matter in two transitions spanning gallery forests and savanna in a Cerrado ecosystem. Gallery forests without grasses typically show a greater leaf area index in contrast to savannas, which show dense grass coverage. Soils of gallery forests have significantly greater concentrations of organic carbon, phosphorus, nitrogen and calcium than those of savannas. Soil organic carbon of savannas is significantly more enriched in ¹³C compared with that of gallery forests. This difference in enrichment is in part caused by the presence of C₄ grasses in savanna ecosystem and its absence in gallery forests. Using the ¹³C abundance as a signature for savanna and gallery forest ecosystems in 1 m soil cores, we show that the borders of gallery forests have expanded into the savanna and that this process initiated at least 3000–4000 bp based on ¹⁴C analysis. Gallery forests, however, may be still expanding as we found more recent transitions according to ¹⁴C activity measurements. We discuss the possible mechanisms of gallery forest expansion and the means by which nutrients required for the expansion of gallery forest might accumulate.     

Herbivory and Stoichiometric Feedbacks to Primary Production

Established theory addresses the idea that herbivory can have positive feedbacks on nutrient flow to plants. Positive feedbacks likely emerge from a greater availability of organic carbon that primes the soil by supporting nutrient turnover through consumer and especially microbially-mediated metabolism in the detrital pool. We developed an entirely novel stoichiometric model that demonstrates the mechanism of a positive feedback. In particular, we show that sloppy or partial feeding by herbivores increases detrital carbon and nitrogen allowing for greater nitrogen mineralization and nutritive feedback to plants. The model consists of differential equations coupling flows among pools of: plants, herbivores, detrital carbon and nitrogen, and inorganic nitrogen. We test the effects of different levels of herbivore grazing completion and of the stoichiometric quality (carbon to nitrogen ratio, C:N) of the host plant. Our model analyses show that partial feeding and plant C:N interact because when herbivores are sloppy and plant biomass is diverted to the detrital pool, more mineral nitrogen is available to plants because of the stoichiometric difference between the organisms in the detrital pool and the herbivore. This model helps to identify how herbivory may feedback positively on primary production, and it mechanistically connects direct and indirect feedbacks from soil to plant production.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.     

Effects of Local Tree Diversity on Herbivore Communities Diminish with Increasing Forest Fragmentation on the Landscape Scale

Forest fragmentation and plant diversity have been shown to play a crucial role for herbivorous insects (herbivores, hereafter). In turn, herbivory-induced leaf area loss is known to have direct implications for plant growth and reproduction as well as long-term consequences for ecosystem functioning and forest regeneration. So far, previous studies determined diverging responses of herbivores to forest fragmentation and plant diversity. Those inconsistent results may be owed to complex interactive effects of both co-occurring environmental factors albeit they act on different spatial scales. In this study, we investigated whether forest fragmentation on the landscape scale and tree diversity on the local habitat scale show interactive effects on the herbivore community and leaf area loss in subtropical forests in South Africa. We applied standardized beating samples and a community-based approach to estimate changes in herbivore community composition, herbivore abundance, and the effective number of herbivore species on the tree species-level. We further monitored leaf area loss to link changes in the herbivore community to the associated process of herbivory. Forest fragmentation and tree diversity interactively affected the herbivore community composition, mainly by a species turnover within the family of Curculionidae. Furthermore, herbivore abundance increased and the number of herbivore species decreased with increasing tree diversity in slightly fragmented forests whereas the effects diminished with increasing forest fragmentation. Surprisingly, leaf area loss was neither affected by forest fragmentation or tree diversity, nor by changes in the herbivore community. Our study highlights the need to consider interactive effects of environmental changes across spatial scales in order to draw reliable conclusions for community and interaction patterns. Moreover, forest fragmentation seems to alter the effect of tree diversity on the herbivore community, and thus, has the potential to jeopardize ecosystem functioning and forest regeneration.     

滇中退化山地不同植被恢复下土壤碳氮磷储量与生态化学计量特征

植被恢复对土壤营养元素的存赋及其生态化学计量特征的影响广受关注,为了深入了解不同植被恢复类型下土壤碳、氮、磷储量与生态化学计量特征,选择滇中地区退化山地飒马场流域具有代表性的4种不同修复阶段的典型植被(荒坡灌草丛、云南松林、针阔混交林和次生常绿阔叶林)为研究对象,分析了不同植被类型下不同深度土壤中有机碳(SOC)、全氮(TN)、全磷(TP)储量和化学计量变化特征。结果表明,退化山地的植被恢复显著改变土壤碳氮磷储存能力和化学计量比,这种改变作用整体上随土壤深度增加而降低。其中,在0—60 cm土层上,SOC储量在次生常绿阔叶林最高,达123.41 t/hm~2,其次是针阔混交林(115.69 t/hm~2)和云南松林(93.08 t/hm~2),荒坡灌草丛(89.56 t/hm~2)最低;TN储量针阔混交林(4.91 t/hm~2)次生常绿阔叶林(4.58 t/hm~2)云南松林(4.43 t/hm~2)荒坡灌草丛(3.98 t/hm~2),4种植被类型间差异显著;TP储量云南松林最高(2.57 t/hm~2),次生常绿阔叶林(2.2 t/hm~2)最低;4种植被类型下土壤C/N介于15.77—30.18,C/P介于29.24—65.33,N/P介于1.28—2.68之间,在0—60 cm土层上均以次生常绿阔叶林最高。植被类型和土壤深度及其交互作用显著影响研究区的SOC、TN和TP储量和化学计量比。分析认为,退化山地不同植被类型对土壤碳氮磷储量和化学计量的影响过程复杂,修复演替进入到次生常绿阔叶林阶段土壤理化性质显著提升,该地区植被修复主要受到氮的限制。研究表征了滇中退化环境植被恢复过程中土壤主要元素变化特征,为揭示植被恢复与土壤生态功能演变关系提供数据支持。     

The effect of nitrogen additions on bracken fern and its insect herbivores at sites with high and low atmospheric pollution

The impact of atmospheric pollution, including nitrogen deposition, on bracken fern herbivores has never been studied. Bracken fern is globally distributed and has a high potential to accumulate nitrogen in plant tissue. We examined the response of bracken fern and its herbivores to N fertilization at a high and low pollution site in forests downwind of Los Angeles, California. Foliage from the high pollution site had higher total N and nitrate than the low pollution site. Bracken fern biomass, foliar N and herbivore abundance were all affected by fertilization treatments. Biomass and herbivore responses were greatest during a year of high precipitation. N additions at the low pollution site were primarily associated with decreased fern biomass, and with transient impacts on herbivore abundance. N additions significantly affected bracken fern and its herbivores at the high pollution site where foliar N and nitrate decreased in response to N addition treatments, while biomass and herbivore abundance increased. High atmospheric deposition and fertilization were both associated with increased herbivore richness. Herbivore abundance was most impacted by fertilization during the early expansion of fern fronds. The most abundant chewing herbivore, a sawfly, was positively associated with plant nitrogen at the low pollution site, but negatively associated with plant nitrogen at the high pollution site, where concentrations of both total N and nitrate were high. While overall growth and herbivore impacts in this xeric location were limited, the variable response we observed associated with rainfall, may indicate impacts could be larger in more mesic environments.     

Host specificity of insect herbivores in tropical forests

Studies of host specificity in tropical insect herbivores are evolving from a focus on insect distribution data obtained by canopy fogging and other mass collecting methods, to a focus on obtaining data on insect rearing and experimentally verified feeding patterns. We review this transition and identify persisting methodological problems. Replicated quantitative surveys of plant-herbivore food webs, based on sampling efforts of an order of magnitude greater than is customary at present, may be cost-effectively achieved by small research teams supported by local assistants. Survey designs that separate historical and ecological determinants of host specificity by studying herbivores feeding on the same plant species exposed to different environmental or experimental conditions are rare. Further, we advocate the use of host-specificity measures based on plant phylogeny. Existing data suggest that a minority of species in herbivore communities feed on a single plant species when alternative congeneric hosts are available. Thus, host plant range limits tend to coincide with those of plant genera, rather than species or suprageneric taxa. Host specificity among tropical herbivore guilds decreases in the sequence: granivores > leaf-miners > fructivore > leaf-chewers = sap-suckers > xylophages > root-feeders, thus paralleling patterns observed in temperate forests. Differences in host specificity between temperate and tropical forests are difficult to assess since data on tropical herbivores originate from recent field studies, whereas their temperate counterparts derive from regional host species lists, assembled over many years. No major increase in host specificity from temperate to tropical communities is evident. This conclusion, together with the recent downward revisions of extremely high estimates of tropical species richness, suggest that tropical ecosystems may not be as biodiverse as previously thought.     

Leaf damage and functional traits along a successional gradient in Brazilian tropical dry forests

Herbivory has significant impacts on individual plants and plant communities, both at ecological and evolutionary time scales. In this context, this study aims to evaluate herbivore damage and its relationship with leaf chemical and structural traits, nutritional status, and forest structural complexity along a successional gradient. We predicted that trees in early successional stages support conservative traits related to drought tolerance (high specific leaf mass and phenolics), whereas trees in light-limited, late successional stages tend to enhance light acquisition strategies (high nitrogen content). We sampled 261 trees from 26 species in 15 plots (50 × 20 m; five per successional stage). From each tree, twenty leaves were collected for leaf trait measures. Phenolic content increased whereas specific leaf mass and nitrogen content decreased from early to late stages. However, leaf damage did not differ among successional stages. Our results partially corroborate the hypothesis that early successional plants in tropical dry forests exhibit leaf traits involved in the conservative use of water. The unexpected decrease in nitrogen content along the chronosequence is likely related to the fact that thinner leaves with low specific leaf mass could have less nitrogen-containing mesophyll per unit area. Mechanisms affecting herbivory intensity varied across scales: at the species level, leaf damage was negatively correlated with tannin concentration and specific leaf mass; at the plot level, leaf damage was positively affected by forest structural complexity. Herbivory patterns in tropical forests are difficult to detect because abiotic factors and multiple top-down and bottom-up forces directly and indirectly affect herbivores.     

Nutrient Addition Differentially Affects Soil Carbon Sequestration in Secondary Tropical Dry Forests: Early‐ versus Late‐Succession Stages

There is considerable interest in the potential use of soils to sequester carbon for climate change mitigation. As such, there is a need to evaluate the potential for carbon accumulation in tropical regions. We compared the effects of three annual additions of nitrogen and/or phosphorus on soil carbon and nitrogen contents and pools (bulk soil, macro‐, meso‐, and microaggregates) of two regenerating secondary tropical dry forest differing in nutrient status and succession stage (10‐year‐old early‐succession stage and approximately 60‐year‐old late‐succession stage). The selected forest sites were located on a shallow calcareous soil in the Yucatán Peninsula (Mexico). The primary production is limited by nitrogen and phosphorus in early‐succession stage and by phosphorus in late‐succession stage. In each forest site, four independent plots (12 × 12 m²) were established, the treatments being: controls and plots fertilized during three consecutive years with nitrogen, phosphorus, or nitrogen plus phosphorus. In both forests, soil carbon and nitrogen contents were consistently high, with soil carbon:nitrogen ratios generally greater than 10. Results indicate that usually there are no significant increases of soil carbon stock associated to late succession but can be increased to 3.7 Mg·ha^?1·yr^?1 with adoption of fertilizer practices. The potential soil carbon sequestration in early‐succession forest was estimated to be 2.7 Mg·ha^?1·yr^?1, and there is no indication that fertilization improves carbon sequestration. In short, results suggest that the soil potential for carbon sequestration in these ecosystems is high and depends on the specific nutrient status of the site.     

Tight Coupling Between Shoot Level Foliar N and P,Leaf Area,and Shoot Growth in Arctic Dwarf Shrubs Under Simulated Climate Change

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.     

Distinct impacts of different mammalian herbivore assemblages on arctic tundra CO2 exchange during the peak of the growing season

Herbivores play a key role in the carbon (C) cycle of arctic ecosystems, but these effects are currently poorly represented within models predicting land–atmosphere interactions under future climate change. Although some studies have examined the influence of various individual species of herbivores on tundra C sequestration, few studies have directly compared the effects of different herbivore assemblages. We measured peak growing season instantaneous ecosystem carbon dioxide (CO₂) exchange (photosynthesis, respiration and net ecosystem exchange) on replicated plots in arctic tundra which, for 14 years, have excluded different portions of the herbivore population (grazed controls, large mammals excluded, both small and large mammals excluded). Herbivory suppressed photosynthetic CO₂ uptake, but caused little change in ecosystem respiration. Despite evidence that small mammals consume a greater portion of plant biomass in these ecosystems, the effect of excluding only large herbivores was indistinguishable from that of excluding both large and small mammals. The herbivory‐induced decline in photosynthesis was not entirely attributable to a decline in leaf area but also likely reflects shifts in plant community composition and/or species physiology. One shrub species – Betula nana – accounted for only around 13% of total aboveground vascular plant biomass but played a central role in controlling ecosystem CO₂ uptake and release, and was suppressed by herbivory. We conclude that herbivores can have large effects on ecosystem C cycling due to shifts in plant aboveground biomass and community composition. An improved understanding of the mechanisms underlying the distinct ecosystem impacts of different herbivore groups will help to more accurately predict the net impacts of diverse herbivore communities on arctic C fluxes.     

Insect Herbivory and Leaf Disease in Natural and Human Disturbed Habitats: Lessons from Early‐Successional Heliconia Herbs

The interaction of plants with insect herbivores and fungal pathogens can affect community dynamics, but there is little information on how this antagonistic interaction may be altered in human‐disturbed tropical systems. We examined whether the amount and quality of foliar damage on the pioneer herbs Heliconia latispatha and Heliconia collinsiana are distinct on road edges and secondary riparian vegetation compared with natural gaps in continuous forest (controls) in Mexico. We also investigated some physical and biological mechanisms that may jointly explain such differences. The overall insect damage in H. latispatha was similar between road edges and natural forest gaps (8.0% vs. 7.2% of leaf area). Damage by caterpillars, however, decreased from 4.2 percent in forest gaps to 0.5 percent on road edges, whereas damage by leaf‐cutting ants increased from 0 to 5.8 percent. In secondary riparian vegetation, where none of the leaves sampled were attacked by ants, overall herbivore damage in H. collinsiana was less than half that observed in forest gaps (3.0% vs. 6.7%), and driven mainly by differences in caterpillar damage (2.5% vs. 6.2%). By contrast, attack by leaf fungal pathogens was two to three times greater in both human‐disturbed habitats than in gaps (8.2–9.6% vs. 3.7–4.2%). Potential mechanisms underlying these differences involved human‐induced shifts in air and soil temperature driven by greater light availability, as well as changes in relative humidity, leaf toughness, foliar condensed tannins, and local abundance of herbivores. Our results indicate that human disturbance alters insect herbivory and may increase proliferation of leaf disease. Abstract in Spanish is available in the online version of this article.