Adaptive strategies to tolerate prolonged flooding in seedlings of floodplain and upland populations of Himatanthus sucuuba, a Central Amazon tree

Long-term flooding imposes a strong selection pressure on plants for the development of protective mechanisms to alleviate the harmful effects of hypoxic and anoxic conditions. This is particularly critical in the Amazonian floodplains where plants withstand annual periods of flooding lasting 7 months and mean flooding amplitude reaching 10 m or more. Himatanthus sucuuba (Apocynaceae) is a tree that is found in the varzea (VZ) floodplains and non-flooded terra firme (TF) forests. It was examined whether individuals from these two contrasting habitats respond differently when subjected to extreme flooding conditions. TF and VZ seedlings were experimentally well-watered, waterlogged (roots and parts of the stems flooded), or submerged (whole plant flooded) during a 4-month period. Anaerobic respiration, evaluated by measuring alcohol dehydrogenase (ADH) activity, and root carbohydrate reserves were quantified, given that the availability of readily fermentable carbohydrates is essential to sustain an active fermentative metabolism. We also assessed changes in morphoanatomy, seedling survival, biomass accumulation and distribution. VZ seedlings had greater root concentrations of soluble sugars and starch, larger seedling mass and accumulated more biomass in roots and stems while TF seedlings allocated more towards stem and leaves. ADH activity was low in seedlings of both populations before exposure to flooding. Waterlogging induced an increase in ADH activity that reached a maximum value in 15 days. Thereafter activity decreased slowly, meanwhile a rapid formation of lenticels, adventitious roots and aerenchyma was observed. Submergence induced leaf shedding and the development of aerenchyma in the root cortex. While VZ seedlings maintained high levels of ADH activity throughout the whole 4-month period, ADH activity in TF seedlings peaked about 15 days after submersion followed by a continuous decrease and death of all the plants. Thus, VZ and TF seedlings differed considerably in terms of tolerating long-term exposure to flooding, especially under total submersion. These results suggest that the predictability and long-term duration of flooding in Central Amazon rivers can impose a selective pressure that is strong enough to result in large phenotypic differences between the two populations of H. sucuuba in the two habitat types.     

2000-2015年安徽省植被净初级生产力时空分布特征及其驱动因素

安徽省是我国的农业大省,其生态系统的动态变化直接关系粮食安全。植被NPP的变化可以有效反映生态系统的变化。基于MOD17A3 NPP数据、气象数据和土地利用类型数据,采用偏差分析法、变异系数、趋势分析法和相关分析法对安徽省2000-2015年植被NPP的时空格局、变化趋势及驱动因子进行研究。结果表明：（1）2000-2015年安徽省植被NPP平均值为476.6gC/m²;波动范围为396.6-531.8gC/m²;植被NPP具有较强的空间分异性,整体上呈现南高北低趋势;（2）不同土地覆盖类型的年均NPP差异明显,其中林地最高,为535.5gC/m²,而且不同地类的NPP年际变化幅度不同,主要表现在林地和草地的变化幅度相对较大;（3）植被NPP受气候、环境变化以及人类活动等多种因素共同影响,其中受气候因素中降雨影响较大,但是随着人类活动日益频繁,城市化逐渐成为NPP变化的主要驱动力。     

A general review of tropical South American floodplains

High rainfall and its seasonal distribution cause periodic flooding of large areas in tropical South America. Floods result from lateral overflow of streams and rivers, or from sheet-flooding by rains as a consequence of poor drainage. Depending upon the size of the catchment area, flooding can occur with one peak (e.g., in the Amazon River and its large affluents) or in many peaks (e.g., in streams and small rivers).Vegetation cover of floodplains varies from different types of savannas and aquatic macrophyte communities to forests depending upon the hydrologic regime and local rainfall. Large differences exist in primary and secondary production due to large differences in nutrient levels in water and soils.An attempt is made to characterize the floodplains according to their hydrologic regimes, vegetation cover and nutrient status. The areal extent of different types of floodplains is estimated. The human impact is also evaluated.From a paper presented at the Third International Wetlands Conference, 19–23 September, 1988, University of Rennes, France.     

Environmental factors exert predominant effects on testate amoeba metacommunities during droughts in floodplains

Metacommunities have been evaluated as models for the relative importance of environmental and spatial processes in assembling ecological communities. Here, we tested the hypothesis that different hydrological periods (drought and flooding) influence the environmental associations of planktonic testate amoeba metacommunities. We predicted that environmental factors would exert the strongest effects on species dispersal under drought, but that they would be less significant during flooding. Testate amoebae were sampled during drought and flooding, from 72 lakes in four Brazilian floodplains (Amazonian, Araguaia, Pantanal and Paraná). Partial redundancy analysis indicated that only environmental factors were significant; they were significant in all floodplain lakes during the drought. Only the Paraná floodplain had significant results for environmental factors during both hydrological periods. Spatial factors did not contribute significantly to any of the metacommunities. The depth, pH and variables related to environmental productivity were identified as major predictors in the assembly of testate amoeba communities. Our results highlight that different hydrological periods vary in their relative importance in environmental and spatial processes. The species‐sorting model was predominant during drought, while stochastic processes prevailed during flooding in all but the Paraná floodplain. In the Paraná floodplain, the construction of dams could potentially alter the effects of environmental and spatial factors on the dispersal of planktonic testate amoebae. The pH and the environmental productivity factors were largely responsible for species selection and the structuring of the planktonic testate amoebae metacommunities in Brazilian floodplains.     

Species richness and abundance of epiphytic Araceae on adjacent floodplain and upland forest in Amazonian Ecuador

Terrestrial plant communities of adjacent upland and floodplain forest of the Amazonian lowland differ from each other in species richness and composition. Epiphytes are generally not considered as being affected by flooding, but we found considerable variation in the communities of epiphytic Araceae of flooded and unflooded forest. Contrary to findings from tree or ground herb communities, no depletion in overall species richness was observed among epiphytic aroids of the floodplains. Abundance and number of epiphytic aroid species per phorophyte were significantly higher than in upland forest, and the species composition varied conspicuously between the two forest types. We suggest that these differences are due to elevated humidity and better soil quality on the floodplains and reject the assumption that flooding has no effect on the epiphytic community.     

Species richness,composition, and spatial distribution of vascular epiphytes in Amazonian black-water floodplain forests

This study examines the occurrence of vascular epiphytic species in Central Amazonian black-water floodplain forests (igapó) and considers whether their horizontal and vertical distribution is influenced by the flood pulse, as is the case with tree species (phorophytes). Research was conducted in sixteen forest plots the Jaú National Park. In these, epiphytes on all phorophytes with DBH ≥ 10 cm were identified. We measured flood height using the watermark left by the last high-water period, then estimated the height relative to the ground of every epiphytic individual. We recorded 653 individuals in 37 species, distributed on 109 phorophytes. Igapó floodplain forests have much lower richness and abundance of vascular epiphyte species than do other Amazonian forests. This may reflect the limitation of available sites for colonization (only 24.9% of studied trees were occupied by epiphytes). Holoepiphytes predominated, and the combined presence of a flood-pulse, linked to the nutrient-poor soil poor seems to limit the occurrence of nomadic vines. Horizontal distribution of epiphytes followed the distribution of phorophytes, which in turn followed the flood-level gradient. Also flooding interacted strongly with vertical zonation to determine species richness. As already well-reported for trees, and unlike reports of epiphytes in other floodplains, flooding strongly influenced richness and distribution of vascular epiphytes in the studied igapó forests.     

Responses to flooding and drying in seedlings of a common Australian desert floodplain shrub: Muehlenbeckia florulenta Meisn. (tangled lignum)

We investigated the effects of flooding and drying over 6 months on growth and biomass allocation in seedlings of Muehlenbeckia florulenta Meisn. (tangled lignum), a common and widely distributed shrub of Australia's desert floodplains. We sought to determine if lignum seedlings respond to flooding or drying by altering traits or allocation patterns or instead display fixed patterns of development. Since desert floodplains are highly unpredictable and heterogeneous environments, we hypothesised that adaptive phenotypic plasticity is unlikely to have developed or be advantageous in seedlings of this species as environmental state changes are highly variable in their timing and duration and plants risk being caught out of kilter with environmental conditions. To test this, we conducted a glasshouse experiment in which lignum seedlings, grown in both clay and sandy sediments, were subjected to a range of hydrological conditions over a period of 6 months. Lignum seedlings exhibited considerable tolerance of both flooding and drying in our experiment and no mortality was observed. Growth was significantly reduced by flooding, however, and seedlings displayed extremely delayed development rather than plasticity in overall biomass allocation or any of the specific morphological variables we measured. Lignum seedlings were considerably more tolerant of drying than flooding and responded plastically by reducing leaf area ratios through reductions in specific leaf areas and leaf production and expansion. Sediment type had little effect on seedling development. Our results indicate that surface water hydrology is likely to be a major determinant of recruitment patterns in this ecologically significant species.     

The contribution of epiphyton to the primary production of tropical floodplain wetlands

Tropical floodplains are one of the most productive ecosystems on earth. Studies on floodplain productivity have primarily focused on trees and macrophytes, rather than algae, due to their greater biomass. However, epiphyton—algae and bacteria attached to the submerged portion of aquatic macrophytes—is a major source of energy in many tropical floodplains. Epiphyton productivity rates are unknown for most tropical floodplain wetlands, and spatial variability is not well understood. In this study, we measured primary productivity of epiphyton in Kakadu National Park in northern Australia. We estimated the relative contribution of epiphyton to aquatic production (epiphyton, + phytoplankton + macrophytes). We sampled sites dominated by different macrophyte structural types: vertical emerging grasses, horizontal emerging grasses, submerged macrophytes, and macrophytes with floating leaves. Epiphyton productivity was highly influenced by the structural type of the macrophyte. Highest potential productivity per weight was measured from epiphyton growing on macrophytes with floating leaves and horizontal grasses (1.52 ± 0.53 and 1.82 ± 0.61 mgC/dw g epiphyton/h, respectively) and lowest in submerged macrophytes and vertical grasses (0.57 ± 0.26 and 0.66 ± 0.47 mgC/dw g epiphyton/h, respectively). When considering the areal biomass of the macrophyte and the amount of epiphyton attached, epiphyton on horizontal grasses and submerged macrophytes had productivity values approximately ten times higher (45–219 mgC/m²/d) compared to those on vertical grasses and macrophytes with floating leaves (2–18 mgC/m²/d). Epiphyton contributed between 2 to 13 percent to the aquatic production of these tropical floodplain wetlands.     

水热波动和土地覆盖变化对东北地区植被NPP的相对影响

研究水热波动和土地覆盖变化对植被净初级生产力（Net Primary Productivity,NPP）的影响对于估算陆地碳循环及其驱动机制具有重要意义。利用MODIS遥感影像获得的时间序列NPP和土地覆盖产品,结合气象观测数据（气温和降水）,采用相关分析、回归分析和空间分析相结合的方法,研究2000-2015年东北地区植被NPP的时空变化特征,并定量评估水热波动和土地覆盖变化对该地区植被NPP的相对影响。研究结果表明,2000-2015年东北地区植被NPP呈波动上升趋势,从2000年的369.24 g C m^-2 a^-1增加到2015年的453.84 g C m^-2 a^-1,平均值是412.10 g C m^-2 a^-1,年际增加速率为4.54 g C m^-2 a^-1。近16年来东北地区年均植被NPP空间上呈现南高北低、东高西低的分布格局,整体变化趋势以增加为主,其中轻微增加面积占该地区总面积的45.9%。不同土地覆盖类型的年均NPP差异明显,其中灌木最高为400.34 g C m^-2 a^-1,草地最低为300.49 g C m^-2 a^-1。东北地区植被NPP与气温的相关性不明显,而与降水量主要表现为正效应。水热波动对该地区不同土地覆盖类型NPP总量变化的贡献大于土地覆盖变化的贡献,其中对森林和农田的贡献最大,均达到70%以上。     

Central Amazon Floodplain Forests: Root Adaptations to Prolonged Flooding

The floodplains of Central Amazonia represent a complex system of inundated river valleys and shallow lakes along the Solimões–Amazonas river, which is subjected to an annual flood-pulse lasting up to 10 months. Such flooding reaching an amplitude of about ten meters causes dramatic changes in the bioavailability of nutrients and oxygen levels and poses extreme constraints for plant survival and reproductivity. Tree species of inundation forests in Central Amazonia had to evolve adaptive mechanisms to both desiccation of soils and partial or full submergence. To adapt to flooded conditions, some trees overcome the flood period by dormancy accompanied by defoliation and formation of annual rings in the wood. Other species maintain metabolism and retain the foliage during the flooding, representing another adaptive mechanism to low oxygen availability. This investigation focused on the root physiology and morphology of six species typical of white-water inundation areas (várzea) led to a preliminary classification of adaptive strategies of trees inhabiting forest communities in floodplains of the Amazon basin.     

Terrestrial invertebrates inhabiting lowland river floodplains of Central Amazonia and Central Europe: a review

1. Amazonian terrestrial invertebrates produce high population densities during favourable periods and may suffer a drastic decrease during occasional floods and droughts. However, the monomodal, predictable flood pulse of the larger Amazonian rivers favours the development of morphological (respiratory organs, wing‐dimorphism), phenological (synchronization of life cycles, univoltine mode of life), physiological (flooding ability, gonad dormancy, alternating number of developmental stages), and behavioural adaptations (migration, temporal diving) with numerous interactions. 2. In lowlands of Central Europe, the flood pulse of large rivers is less predictable than in Central Amazonia and is superimposed by the seasonal light/temperature pulse (summer/winter regime). Some terrestrial invertebrates show physiological resistance against inundation or drought, phenologies fitting the normal annual rhythm of water level fluctuation (quiescence or diapause of eggs or adult invertebrates), high dispersal ability and migration. However, most species survive simply using a `risk strategy', combining high reproduction rates, dispersal and reimmigration following catastrophic events. 3. The diversity of species in terrestrial invertebrates is lower in lowland riverine ecosystems of Central Amazonia and Central Europe compared with the respective uplands because of flood stress in these systems. However, floodplains in Central Amazonia possess a greater number of endemic species in comparison with Central European floodplains because of long periods of fairly stable climatic conditions in comparison with large palaeoclimatic changes in Central Europe.     

Submerged in darkness: adaptations to prolonged submergence by woody species of the Amazonian floodplains

Background

In Amazonian floodplain forests, >1000 tree species grow in an environment subject to extended annual submergence which can last up to 9 months each year. Water depth can reach 10 m, fully submerging young and also adult trees, most of which reproduce during the flood season. Complete submergence occurs regularly at the seedling or sapling stage for many species that colonize low-lying positions in the flooding gradient. Here hypoxic conditions prevail close to the water surface in moving water, while anaerobic conditions are common in stagnant pools. Light intensities in the floodwater are very low.

Questions and Aims

Despite a lack of both oxygen and light imposed by submergence for several months, most leafed seedlings survive. Furthermore, underwater growth has also been observed in several species in the field and under experimental conditions. The present article assesses how these remarkable plants react to submergence and discusses physiological mechanisms and anatomical adaptations that may explain their success.Key words: Adaptation, Amazonian floodplains, darkness, environmental stress, flooding, hypoxia, submergence tolerance, trees, underwater photosynthesis, woody species     

黄土高原草地净初级生产力时空趋势及其驱动因素

草地净初级生产力是生态系统碳循环的关键环节和重要组成部分.本研究使用分段线性回归分析和Pearson相关分析,分析了黄土高原2000-2015年间土地利用类型未改变的草地净初级生产力(NPP)的变化趋势及气候核心因子(年降水量、年强降水量、年有效降水日数、年平均温度、年最高温度、年最低温度)对NPP变化的影响,并借助增...     

Effects of hydrological variation on the aquatic plant community in a floodplain palustrine wetland of southern Brazil

This study analyzed macrophyte richness, biomass, and composition under flooding of brief duration (less than 3 days) and drawdown events over an annual cycle in a floodplain palustrine wetland in the south of Brazil. The study was carried out to test the hypothesis that floods of brief and very brief duration are not long enough to compromise the richness and the biomass of aquatic macrophytes and that the alternation between wet and drawdown phases may cause variations in the macrophyte richness and composition. A total of 26 aquatic macrophyte species were observed from April 2003 to May 2004: 13 species were observed during the wet phase, and 24 during the drawdown phase. The mean richness was higher during the drawdown phase than during the wet phase, however, the mean biomass was similar in both phases. Although macrophyte richness was not modified after the three flooding events, mean biomass was modified after two events. The number of macrophyte species of which the biomass was modified after the first flooding event increased with subsequent floods. These results illustrate the importance of the dynamics between brief floods and drawdown events to the aquatic plant community in floodplain wetlands in southern Brazil.     

Do flood pulses structure amphibian communities in floodplain environments?

Beta diversity can provide insights into the processes that regulate communities subjected to frequent disturbances, such as flood pulses, which control biodiversity in floodplains. However, little is known about which processes structure beta diversity of amphibians in floodplains. Here, we tested the influence of flood pulses on the richness, composition, and beta diversity of amphibians in Amazonian floodplain environments. We also evaluated indicator species for each environment. We established linear transects in three environments: low várzea, high várzea, and macrophyte rafts. Species richness decreased and beta diversity increased according to the susceptibility of habitats to flood pulses. Indicator species differed among environments according to forest succession promoted by the flood pulse. The decrease in species richness between high and low várzea is due to non‐random extinctions. The higher rates of species turnover between várzeas and macrophyte rafts are driven by the colonization of species adapted to open areas. Our results highlight that the maintenance of complex environments is needed to protect biodiversity in floodplains.     

Spatial and temporal variations in densities of small fishes across different temporary floodplain types of the lower Okavango Delta,Botswana

Small-sized and juvenile fishes as well as physicochemical water parameters in various primary, secondary and rarely flooded temporary floodplains of the Okavango Delta were assessed during the different hydrological phases of the 2009–2010 flooding season. Small fishes were sampled in the marginal zone of the floodplains using a throw-trap net and a suite of physicochemical properties were measured. Both physicochemical water quality parameters and densities of small fishes showed spatiotemporal variations across the temporary floodplain types. Turbidity, dissolved oxygen, conductivity, woody debris and chlorophyll a were all highest in rarely flooded floodplains compared to both primary and secondary floodplains (ANOVA, p < 0.05). Fish from 38 fish taxa belonging to 11 families were identified and classified during the study. The fish families Cichlidae (dominated by tilapias), Poeciliidae (Aplocheilichthys spp.) and Cyprinidae (Barbus spp.), were the most abundant across the study floodplains. The poeciliids were most abundant in frequently flooded primary and secondary floodplains, whereas juvenile cichlids dominated in rarely flooded floodplains. During high floods the rarely flooded portions of the delta function as important nursery habitats for juvenile cichlids, implying that a significant reduction in flooding may have negative effects on the delta's tilapia stocks due to reduced recruitment success.     

Flow-induced alterations to fish assemblages, habitat and fish–habitat associations in a regulated lowland river

Understanding the cause–effect response of aquatic biota to hydrological variability is fundamental to the restoration of regulated rivers. Spatio-temporal variation in fish assemblage structure, microhabitat cover and fish–habitat associations were investigated in the main channel of the regulated lower River Murray, Australia, during a prolonged period of low within-channel flows and following a high flow event and flood. Several small-bodied species (e.g. carp gudgeon, Hypseleotris spp.), were abundant and significantly associated with submerged macrophytes during low flows, but were absent or significantly less abundant following flooding, and the loss of these microhabitats. Large-bodied riverine species that spawn in response to increases in flow (e.g. golden perch, Macquaria ambigua ambigua) or spawn and recruit in inundated floodplain habitats (e.g. common carp, Cyprinus carpio), exhibited flexible microhabitat use and were significantly more abundant following flooding. In the lower River Murray, high flow events appear integral in structuring fish assemblages, indirectly influencing the abundance of small-bodied fish by re-structuring macrophyte cover and directly influencing the abundance of large-bodied species by facilitating critical life history processes (e.g. recruitment). These results highlight species-specific differences in cause–effect responses to flow variability and have implications for managing flow in regulated rivers.     

Controls on water balance of shallow thermokarst lakes and their relations with catchment characteristics: a multi‐year,landscape‐scale assessment based on water isotope tracers and remote sensing in Old Crow Flats,Yukon (Canada)

Many northern lake‐rich regions are undergoing pronounced hydrological change, yet inadequate knowledge of the drivers of these landscape‐scale responses hampers our ability to predict future conditions. We address this challenge in the thermokarst landscape of Old Crow Flats (OCF) using a combination of remote sensing imagery and monitoring of stable isotope compositions of lake waters over three thaw seasons (2007–2009). Quantitative analysis confirmed that the hydrological behavior of lakes is strongly influenced by catchment vegetation and physiography. Catchments of snowmelt‐dominated lakes, typically located in southern peripheral areas of OCF, encompass high proportions of woodland/forest and tall shrub vegetation (mean percent land cover = ca. 60%). These land cover types effectively capture snow and generate abundant snowmelt runoff that offsets lake water evaporation. Rainfall‐dominated lakes that are not strongly influenced by evaporation are typically located in eastern and northern OCF where their catchments have higher proportions of dwarf shrub/herbaceous and sparse vegetation (ca. 45%), as well as surface water (ca. 20%). Evaporation‐dominated lakes, are located in the OCF interior where their catchments are distinguished by substantially higher lake area to catchment area ratios (LA/CA = ca. 29%) compared to low evaporation‐influenced rainfall‐dominated (ca. 10%) and snowmelt‐dominated (ca. 4%) lakes. Lakes whose catchments contain >75% combined dwarf shrub/herbaceous vegetation and surface water are most susceptible to evaporative lake‐level drawdown, especially following periods of low precipitation. Findings indicate that multiple hydrological trajectories are probable in response to climate‐driven changes in precipitation amount and seasonality, vegetation composition, and thermokarst processes. These will likely include a shift to greater snowmelt influence in catchments experiencing expansion of tall shrubs, greater influence from evaporation in catchments having higher proportions of surface water, and an increase in the rate of thermokarst lake expansion and probability of drainage. Local observations suggest that some of these changes are already underway.     

What controls variation in carbon use efficiency among Amazonian tropical forests?

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPP_wood), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (R_a; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times <40 yrs had significantly lower maintenance respiration for both wood and fine roots than forests with residence times >40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long‐term mean annual temperatures do not impact CUE.