Spatiotemporal pattern of specialization of sunbird-plant networks on Mt. Cameroon.

Differences in interaction specializations between nectarivorous birds and plants across continents serve as common examples of evolutionary trajectory specificity. While New World hummingbird-plant networks have been extensively studied and are considered highly specialized, knowledge on the network specialization of their Old World counterparts, sunbirds (Nectariniidae), remains limited. A few studies from tropical Africa indicate that sunbird-plant networks are rather generalized. Unfortunately, these studies are limited to dry seasons and high elevations at the tree line, environments where niche-based hypotheses also often predict lower resource partitioning. In our study, we explored the specialization of sunbird-plant networks and their spatiotemporal variability on Mt. Cameroon (Cameroon). Using a combination of automatic video recordings and personal observations, we constructed eight comprehensive sunbird-plant networks in four forest types at different elevations in both the dry and wet seasons. As reported in previous studies, the montane forest plants, birds and whole networks were highly generalized. Nevertheless, we observed a much higher specialization in forests at lower elevations. Except at the lowest altitude, the wet season was also characterized by higher specialization. While less specialized flowering trees dominated in the dry season networks, more specialized herbs and shrubs were visited by birds during the wet season. As our findings do not support the generally accepted assumption that Old World bird-plant networks are rather generalized, we need further studies to understand the differences in bird-plant specializations on individual continents.

Conservation and restoration of Central European fens by mowing: A consensus from 20 years of experimental work.

While the importance of conservation mowing for mesic grasslands is generally accepted, its use for fens and fen grasslands interspersed within agricultural land is still controversial. Although fens may persist naturally, ongoing environmental changes increase productivity and accelerate succession. These processes can be mitigated through conservation management with appropriate settings. However, long-term management experiments are challenging and provide only locally valid results. Here, we analysed vegetation data (bryophytes and vascular plants) from seven management experiments (spanning 3-20 years) conducted in Central European poor, moderately-rich, and calcareous spring fens (Czech Republic, Slovakia). Two of these experiments examined the effects of restoration of abandoned fens, while five experiments examined changes in mowing regimes in managed fens (cessation, intensification, delay to autumn, and litter removal). Data were analysed using unidimensional and multidimensional methods separately for the initial, extended, and entire period. Mowing had a statistically significant effect on species composition except for the shortest (3-year) experiment. Litter removal did not compensate for mowing. Mowing twice or delayed mowing significantly affected the species composition of calcareous fens. In all cases, cessation of mowing significantly reduced the richness of species, especially those of conservation importance. In contrast, any mowing of abandoned fens increased species richness. The effects of mowing intensification or cessation on species richness and composition of a restored calcareous fen were evident in the first 2-3 years. Other effects were initially weak or nonsignificant but later became stronger, such as mowing delay and restoration removal of litter, which became significant only after nearly 20 years. We found that cessation or restoration of mowing usually triggers a rapid vegetation change, whereas it can take decades to detect the response caused by changes in mowing timing. Importantly, mowing can stabilise or even restore vegetation of fen ecosystems that have been weakened by their fragmentation in the temperate agricultural landscapes.

Spatiotemporal variation in the role of floral traits in shaping tropical plant-pollinator interactions.

The pollination syndrome hypothesis predicts that plants pollinated by the same pollinator group bear convergent combinations of specific floral functional traits. Nevertheless, some studies have shown that these combinations predict pollinators with relatively low accuracy. This discrepancy may be caused by changes in the importance of specific floral traits for different pollinator groups and under different environmental conditions. To explore this, we studied pollination systems and floral traits along an elevational gradient on Mount Cameroon during wet and dry seasons. Using Random Forest (Machine Learning) models, allowing the ranking of traits by their relative importance, we demonstrated that some floral traits are more important than others for pollinators. However, the distribution and importance of traits vary under different environmental conditions. Our results imply the need to improve our trait-based understanding of plant-pollinator interactions to better inform the debate surrounding the pollination syndrome hypothesis.

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements.

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

Differences in Nectar Traits between Ornithophilous and Entomophilous Plants on Mount Cameroon.

Despite a growing number of studies, the role of pollinators as a selection agent for nectar traits remains unclear. Moreover, the lack of data from some biogeographic regions prohibits us from determining their general importance and global patterns. We analyzed nectar carbohydrate traits and determined the main pollinators of 66 plant species in the tropical forests of Mount Cameroon (tropical West Africa). The measured nectar traits included total sugar amounts and proportions of sucrose and hexoses (i.e., glucose and fructose). We report the nectar properties for plants visited by five pollinator groups (bees, butterflies, moths, hoverflies, and specialized birds). Our results indicate that, rather than specific evolution in each of the five plant groups, there was a unique nectar-trait evolution in plants pollinated by specialized birds. The ornithophilous plants had a higher proportion of sucrose and produced larger sugar amounts than the plants pollinated by insects. We also demonstrated a significant phylogenetic signal in the nectar properties in some lineages of the studied plants.

Elevational and seasonal patterns of butterflies and hawkmoths in plant-pollinator networks in tropical rainforests of Mount Cameroon.

Butterflies and moths are conspicuous flower visitors but their role in plant-pollinator interactions has rarely been quantified, especially in tropical rainforests. Moreover, we have virtually no knowledge of environmental factors affecting the role of lepidopterans in pollination networks. We videorecorded flower-visiting butterflies and hawkmoths on 212 plant species (> 26,000 recorded hrs) along the complete elevational gradient of rainforests on Mount Cameroon in dry and wet seasons. Altogether, we recorded 734 flower visits by 80 butterfly and 27 hawkmoth species, representing only ~ 4% of all flower visits. Although lepidopterans visited flowers of only a third of the plant species, they appeared to be key visitors for several plants. Lepidopterans visited flowers most frequently at mid-elevations and dry season, mirroring their local elevational patterns of diversity. Characteristics of interaction networks showed no apparent elevational or seasonal patterns, probably because of the high specialisation of all networks. Significant non-linear changes of proboscis and forewing lengths were found along elevation. A positive relationship between the lengths of proboscis of hesperiid butterflies and tube of visited flowers was detected. Differences in floral preferences were found between sphingids and butterflies, revealing the importance of nectar production, floral size and shape for sphingids, and floral colour for butterflies. The revealed trait-matching and floral preferences confirmed their potential to drive floral evolution in tropical ecosystems.

A geographical perspective on the relationship between Impatiens spur lengths and bill lengths of sunbirds in Afrotropical mountains.

Trait matching-a correlation between the morphology of plants and their pollinators-has been frequently observed in pollination interactions. Different intensities of natural selection in individual regions should cause such correlations to be observable across different local assemblages. In this study, we focused on matching between spur lengths of the genus Impatiens and bill lengths of sunbirds in tropical Africa. For 25 mountain and island locations, we compiled information about the composition and traits of local Impatiens and sunbird assemblages. We found that assemblage mean and maximum values of bill lengths were positively correlated with mean and maximum spur lengths across locations. Moreover, our results suggest that the positive correlations hold only for forest sunbird assemblages sharing the same habitat with Impatiens species. We further show that long-billed sunbirds seem to locally match the morphology of multiple Impatiens plant species, not vice versa. Our observation implies that trait matching significantly contributes to structuring of Impatiens-sunbird pollination systems. We suggest that special habitat preferences together with spatial isolation of mountain environment might play a role in this case.

Carbohydrate storage in herbs: the forgotten functional dimension of the plant economic spectrum.

BACKGROUND AND AIMS: Although the plant economic spectrum seeks to explain resource allocation strategies, carbohydrate storage is often omitted. Belowground storage organs are the centre of herb perennation, yet little is known about the role of their turnover, anatomy and carbohydrate storage in relation to the aboveground economic spectrum. METHODS: We collected aboveground traits associated with the economic spectrum, storage organ turnover traits, storage organ inner structure traits and storage carbohydrate concentrations for ~80 temperate meadow species. KEY RESULTS: The suites of belowground traits were largely independent of one another, but there was significant correlation of the aboveground traits with both inner structure and storage carbohydrates. Anatomical traits diverged according to leaf nitrogen concentration on the one hand and vessel area and dry matter content on the other; carbohydrates separated along gradients of leaf nitrogen concentration and plant height. CONCLUSIONS: Contrary to our expectations, aboveground traits and not storage organ turnover were correlated with anatomy and storage carbohydrates. Belowground traits associated with the aboveground economic spectrum also did not fall clearly within the fast-slow economic continuum, thus indicating the presence of a more complicated economic space. Our study implies that the generally overlooked role of storage within the plant economic spectrum represents an important dimension of plant strategy.

Are Reproductive Traits Related to Pollen Limitation in Plants? A Case Study from a Central European Meadow.

The deficiency of pollen grains for ovule fertilization can be the main factor limiting plant reproduction and fitness. Because of the ongoing global changes, such as biodiversity loss and landscape fragmentation, a better knowledge of the prevalence and predictability of pollen limitation is challenging within current ecological research. In our study we used pollen supplementation to evaluate pollen limitation (at the level of seed number and weight) in 22 plant species growing in a wet semi-natural meadow. We investigated the correlation between the pollen limitation index (PL) and floral traits associated with plant reproduction or pollinator foraging behavior. We recorded significant pollen limitation for approximately 41% of species (9 out of 22 surveyed). Seven species had a significant positive response in seed production and two species increased in seed weight after pollen supplementation. Considering traits, PL significantly decreased with the number of pollinator functional groups. The relationship of PL with other examined traits was not supported by our results. The causes of pollen limitation may vary among species with regard to (1) different reproductive strategies and life history, and/or (2) temporary changes in influence of biotic and abiotic factors at a site.

Ecological fitting is a sufficient driver of tight interactions between sunbirds and ornithophilous plants.

Plant-bird pollination interactions evolved independently on different continents. Specific adaptations can lead to their restriction when potential partners from distant evolutionary trajectories come into contact. Alternatively, these interactions can be enabled by convergent evolution and subsequent ecological fitting.We studied the interactions between New World plants from the genus Heliconia, Asian plants of genus Etlingera and African sunbirds on a local farm in Cameroon. Heliconia spp. evolved together with hummingbirds and Etlingera spp. with spiderhunters -an oriental subgroup of the sunbird family.Sunbirds fed on all studied plants and individual plant species were visited by a different sunbird spectrum. We experimentally documented a higher number of germinated pollen grains in sunbird-visited flowers of Etlingera spp. For Heliconia spp., this experiment was not successful and pollen tubes were rarely observed, even in hand-pollinated flowers, where enough pollen was deposited. The analyses of contacts with plant reproductive organs nevertheless confirmed that sunbirds are good pollen vectors for both Heliconia and Etlingera species.Our study demonstrated a high ecological fit between actors of distinct evolutionary history and the general validity of bird-pollination syndrome. We moreover show that trait matching and niche differentiation are important ecological processes also in semi-artificial plant-pollinator systems.

First records of 31 species of butterflies and moths (Lepidoptera) in Cameroon, with remarks on their elevational ranges.

BACKGROUND: The biodiversity of West and Central Africa is understudied, including butterflies and moths (Lepidoptera). Cameroon, through its position in between few biogeographic regions and diversity of habitats, is an important hotspot of lepidopteran diversity. However, the country also ranks low when it comes to local biodiversity knowledge. During our long-term ecological projects in the Cameroonian part of the Gulf of Guinea Highlands, we collected rich material of butterflies and moths, including a number of interesting faunistic records. NEW INFORMATION: In this study, we report 31 species of butterflies and moths which have not yet been recorded in Cameroon. These species comprised eight new genera records for the country. In many cases, our records represented an important extension of the species' known distribution, including ten species whose distribution ranges extended into the Guinean biogeographic region. We also comment on the species' elevational distribution ranges on Mount Cameroon where most of our records originated. Additionally, we confirm the presence of a butterfly Telchinia encedena, after more than a century since its first and so far its only record in Cameroon.

The role of ultraviolet reflectance and pattern in the pollination system of Hypoxis camerooniana (Hypoxidaceae).

Apart from floral morphology and colours perceived by the human eye, ultraviolet (UV) reflectance acts as an important visual advertisement of numerous flowering plant species for pollinators. However, the effect of UV signalling on attracting pollinators of particular plant species is still insufficiently studied, especially in the Afrotropics. Therefore, we studied the pollination system of Hypoxis camerooniana in montane grasslands of Mount Cameroon, West/Central Africa. We focused mainly on the effects of the flowers' UV reflectance on its visitors. We experimentally removed UV reflection from petals either completely or partially. Thereafter, flower visitors were recorded and pistils were collected post-flowering to quantify germinated pollen tubes per treatments. The most important visitors were bees, followed by flies. Due to their contacts with reproductive organs bees are considered as the primary pollinators. Visitation rates were lower when UV reflectance was completely removed, whereas the decrease of frequency on half-treated flowers did not differ significantly from control treatments. The complete removal of UV also affected bees' landing behaviour, but not that of flies. We showed that the presence of UV reflectance is more important than UV pattern for bees visiting flowers of H. camerooniana. We hypothesize that exploiting all flowers irrespective of their pattern can be more efficient for pollinators in the open grasslands of high altitudes to spot these relatively scarce flowers by their UV reflectance. Furthermore, we highlight the necessity of both experimental and natural controls in similar studies to control for additional effects of the used UV manipulations. Many plants advertise their flowers with UV reflectance visible to their insect visitors. By manipulating the UV reflectance and pattern of Hypoxis camerooniana in the Afromontane grasslands of Mount Cameroon, we have shown how crucial it is for the predominant visitor, bees. Both bees' preferences for flowers and their behaviour during visits are influenced by changes in UV reflectance. However, the presence of some UV signal is more important than the specific pattern. Especially in montane grasslands with higher UV irradiation, the UV floral colours are important for recognition of flowers by potential pollinators.

Experimental loss of generalist plants reveals alterations in plant-pollinator interactions and a constrained flexibility of foraging.

Species extinctions undermine ecosystem functioning, with the loss of a small subset of functionally important species having a disproportionate impact. However, little is known about the effects of species loss on plant-pollinator interactions. We addressed this issue in a field experiment by removing the plant species with the highest visitation frequency, then measuring the impact of plant removal on flower visitation, pollinator effectiveness and insect foraging in several sites. Our results show that total visitation decreased exponentially after removing 1-4 most visited plants, suggesting that these plants could benefit co-occurring ones by maintaining high flower visitor abundances. Although we found large variation among plant species, the redistribution of the pollinator guild affected mostly the other plants with high visitor richness. Also, the plant traits mediated the effect of removal on flower visitation; while visitation of plants which had smaller inflorescences and more sugar per flower increased after removal, flower visitors did not switch between flower shapes and visitation decreased mostly in plants visited by many morpho-species of flower visitors. Together, these results suggest that the potential adaptive foraging was constrained by flower traits. Moreover, pollinator effectiveness fluctuated but was not directly linked to changes of flower visitation. In conclusion, it seems that the loss of generalist plants alters plant-pollinator interactions by decreasing pollinator abundance with implications for pollination and insect foraging. Therefore, generalist plants have high conservation value because they sustain the complex pattern of plant-pollinator interactions.

Flying between raindrops: Strong seasonal turnover of several Lepidoptera groups in lowland rainforests of Mount Cameroon.

Although seasonality in the tropics is often less pronounced than in temperate areas, tropical ecosystems show seasonal dynamics as well. Nevertheless, individual tropical insects' phenological patterns are still poorly understood, especially in the Afrotropics. To fill this gap, we investigated biodiversity patterns of Lepidoptera communities at three rainforest localities in the foothills of Mount Cameroon, West Africa, one of the wettest places in the world. Our multitaxa approach covered six lepidopteran groups (fruit-feeding butterflies and moths, the families Sphingidae, Saturniidae, and Eupterotidae, and the subfamily Arctiinae of Erebidae) with diverse life strategies. We sampled adults of the focal groups in three distinct seasons. Our sampling included standardized bait trapping (80 traps exposed for 10 days per locality and season) and attraction by light (six full nights per locality and season). Altogether, our dataset comprised 20,576 specimens belonging to 559 (morpho)species of the focal groups. The biodiversity of Lepidoptera generally increased in the high-dry season, and either increased (fruit-feeding moths, Arctiinae, Saturniidae) or decreased (butterflies, Sphingidae) in the transition to the wet season in particular groups. Simultaneously, we revealed a strong species turnover of fruit-feeding Lepidoptera and Arctiinae among the seasons, indicating relatively high specialization of these communities for particular seasons. Such temporal specialization can make the local communities of butterflies and moths especially sensitive to the expected seasonal perturbations caused by the global change. Because of the key role of Lepidoptera across trophic levels, such changes in their communities could strengthen this impact on entire tropical ecosystems.

Is the scaling relationship between carbohydrate storage and leaf biomass in meadow plants affected by the disturbance regime?

BACKGROUND AND AIMS: Below-ground carbohydrate storage is considered an adaptation of plants aimed at regeneration after disturbance. A theoretical model by Iwasa and Kubo was empirically tested which predicted (1) that storage of carbohydrates scales allometrically with leaf biomass and (2) when the disturbance regime is relaxed, the ratio of storage to leaf biomass increases, as carbohydrates are not depleted by disturbance. METHODS: These ideas were tested on nine herbaceous species from a temperate meadow and the disturbance regime was manipulated to create recently abandoned and mown plots. Just before mowing in June and at the end of the season in October, plants with below-ground organs were sampled. The material was used to assess the pool of total non-structural carbohydrates and leaf biomass. KEY RESULTS: In half of the cases, a mostly isometric relationship between below-ground carbohydrate storage and leaf biomass in meadow plants was found. The ratio of below-ground carbohydrate storage to leaf biomass did not change when the disturbance regime was less intensive than that for which the plants were adapted. CONCLUSIONS: These findings (isometric scaling relationship between below-ground carbohydrate storage and leaf biomass; no effect of a relaxed disturbance regime) imply that storage in herbs is probably governed by factors other than just the disturbance regime applied once in a growing season.

The nectar spur is not only a simple specialization for long-proboscid pollinators.

Since the time of Darwin, biologists have considered the floral nectar spur to be an adaptation representing a high degree of plant specialization. Nevertheless, some researchers suggest that nature is more complex and that even morphologically specialized plants attract a wide spectrum of visitors. We observed visitors on Impatiens burtonii (Balsaminaceae) and measured the depth of the proboscis insertion into the spur, the distance of the nectar surface from the spur entrance and the visitor's effectiveness. The hoverfly Melanostoma sp., with the shortest proboscis, was most active early in the morning and fed on pollen and nectar near the spur entrance. The honeybee Apis mellifera and the hoverfly Rhingia mecyana were the most frequent visitors before and after noon, respectively. Although R. mecyana, the only visitor able to reach the end of the spur, was the most frequent, it did not deposit the largest number of pollen grains per visit. Nectar spurs may function as complex structures allowing pollination by both short- and long-proboscid visitors and separating their spatial and temporal niches. Spurred plants should be considered as more generalized and exposed to more diverse selection pressures than previously believed.

Asymmetric competition for nectar between a large nectar thief and a small pollinator: an energetic point of view.

There are two alternative hypotheses related to body size and competition for restricted food sources. The first one supposes that larger animals are superior competitors because of their increased feeding abilities, whereas the second one assumes superiority of smaller animals because of their lower food requirements. We examined the relationship between two unrelated species of different size, drinking technique, energy requirements and roles in plant pollination system, to reveal the features of their competitive interaction and mechanisms enabling their co-existence while utilising the same nectar source. We observed diurnal feeding behaviour of the main pollinator, the carpenter bee Xylocopa caffra and a nectar thief, the northern double-collared sunbird Cinnyris reichenowi on 19 clumps of Hypoestes aristata (Acanthaceae) in Bamenda Highlands, Cameroon. For comparative purpose, we established a simplistic model of daily energy expenditure and daily energy intake by both visitor species assuming that they spend all available daytime feeding on H. aristata. We revealed the energetic gain-expenditure balance of the studied visitor species in relation to diurnal changes in nectar quality and quantity. In general, smaller energy requirements and related ability to utilise smaller resources made the main pollinator X. caffra competitively superior to the larger nectar thief C. reichenowi. Nevertheless, sunbirds are endowed with several mechanisms to reduce asymmetry in exploitative competition, such as the use of nectar resources in times of the day when rivals are inactive, aggressive attacks on carpenter bees while defending the nectar plants, and higher speed of nectar consumption.

Effects of long- and short-term management on the functional structure of meadows through species turnover and intraspecific trait variability.

The functional structures of communities respond to environmental changes by both species replacement (turnover) and within-species variation (intraspecific trait variability; ITV). Evidence is lacking on the relative importance of these two components, particularly in response to both short- and long-term environmental disturbance. We hypothesized that such short- and long-term perturbations would induce changes in community functional structure primarily via ITV and turnover, respectively. To test this we applied an experimental design across long-term mown and abandoned meadows, with each plot containing a further level of short-term management treatments: mowing, grazing and abandonment. Within each plot, species composition and trait values [height, shoot biomass, and specific leaf area (SLA)] were recorded on up to five individuals per species. Positive covariations between the contribution of species turnover and ITV occurred for height and shoot biomass in response to both short- and long-term management, indicating that species turnover and intraspecific adjustments selected for similar trait values. Positive covariations also occurred for SLA, but only in response to long-term management. The contributions of turnover and ITV changed depending on both the trait and management trajectory. As expected, communities responded to short-term disturbances mostly through changes in intraspecific trait variability, particularly for height and biomass. Interestingly, for SLA they responded to long-term disturbances by both species turnover and intraspecific adjustments. These findings highlight the importance of both ITV and species turnover in adjusting grassland functional trait response to environmental perturbation, and show that the response is trait specific and affected by disturbance regime history.

Effects of disturbance regime on carbohydrate reserves in meadow plants.

Carbohydrate storage enables plants to tolerate both seasonally unfavourable conditions and recover from disturbance. Although short-term changes in storage levels due to disturbance are fairly well known, less is known about long-term changes in storage levels, especially in response to cessation of repeated disturbance. Additionally, whereas it is presumably the total amount (pool) of storage carbohydrate reserves that is of importance, typically carbohydrate concentrations are measured instead, as a proxy. We assessed changes in carbohydrate concentrations and pools in storage organs and changes in above- versus belowground biomass in response to mowing cessation in nine herbs from two meadows (dry and wet) at the (June) peak of vegetation development and the (October) growing season end 1 and 3 years after the change in the disturbance regime. We tested three hypotheses: (1) storage will increase with abandonment of mowing only in the first year after disturbance cessation, but not further increase subsequently, as high storage would hinder competitive ability; (2) storage will increase towards the end of the season in both disturbed and undisturbed plants; and (3) changes in carbohydrate concentrations are accurate predictors of changes in pools. Although species-specific changes in carbohydrate reserves occurred in the wet meadow, more general trends appeared in the dry meadow. There, plants accumulated higher carbohydrate reserves at the end of the season, especially in unmown plots. However, the reserves for plants in both disturbance regimes were the same at the growing season peak (June) in both examined years. The increase in storage of carbohydrates on unmown plots in October was manifested by increases of both storage organ biomass and carbohydrate concentration, whereas in mown plots, it was due only to increased carbohydrate concentration. Although concentrations and pools represent different aspects of plant carbohydrate economy, concentrations will represent short-term responses to changed disturbance regimes.