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     

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.     

Provisioning of bioavailable carbon between the wet and dry phases in a semi-arid floodplain

Ecosystem functioning on arid and semi-arid floodplains may be described by two alternate traditional paradigms. The pulse-reserve model suggests that rainfall is the main driver of plant growth and subsequent carbon and energy reserve formation in the soil of arid and semi-arid regions. The flood pulse concept suggests that periodic flooding facilitates the two-way transfer of materials between a river and its adjacent floodplain, but focuses mainly on the period when the floodplain is inundated. We compared the effects of both rainfall and flooding on soil moisture and carbon in a semi-arid floodplain to determine the relative importance of each for soil moisture recharge and the generation of a bioavailable organic carbon reserve that can potentially be utilised during the dry phase. Flooding, not rainfall, made a substantial contribution to moisture in the soil profile. Furthermore, the growth of aquatic macrophytes during the wet phase produced at least an order of magnitude more organic material than rainfall-induced pulse-reserve responses during the dry phase, and remained as recognizable soil carbon for years following flood recession. These observations have led us to extend existing paradigms to encompass the reciprocal provisioning of carbon between the wet and dry phases on the floodplain, whereby, in addition to carbon fixed during the dry phase being important for driving biogeochemical transformations upon return of the next wet phase, aquatic macrophyte carbon fixed during the wet phase is recognized as an important source of energy for the dry phase. Reciprocal provisioning presents a conceptual framework on which to formulate questions about the resistance and ecosystem resilience of arid and semi-arid floodplains in the face of threats like climate change and alterations to flood regimes.     

Impacts of hydrological restoration on lowland river floodplain plant communities

Many lowland floodplain habitats have been disconnected from their rivers by flood defence banks. Removing or lowering these banks can reinstate regular flooding and thus restore these important wetland plant communities. In this study we analyse changes in wetland hydrology and plant community composition following the lowering of flood defence banks at a floodplain of the River Don in the United Kingdom (UK). The aim of the restoration project was to improve the quality of “floodplain grazing marsh” habitat, which is a group of wetland communities that are of conservation interest in the UK. We analyse changes in species richness and community composition over a period of 6 years, and compare the presence of indicator species from the target floodplain grazing marsh plant communities. The lowering of the flood banks increased the frequency of flood events, from an estimated average of 1.7 floods per year to 571 floods per year. The increased flooding significantly increased the proportion of time that the wetland was submerged, and the heterogeneity in hydrological conditions within the floodplain. There were significant differences in composition between the pre-restoration and restored plant communities. Plants with traits for moisture tolerance became more abundant, although the communities did not contain significantly more ‘target’ floodplain grazing marsh species at the end of the study period than prior to restoration. Colonisation by floodplain grazing marsh species may have been limited because environmental conditions were not yet suitable, or because of a shortage of colonising propagules. While the desired target plant community has not been achieved after 5 years, it is encouraging that the community has changed dynamically as a result of hydrological changes, and that moisture-tolerant species have increased in occurrence. Over the next few decades, the restored flood regime may cause further environmental change or colonisation events, thus helping increase the occurrence of desired floodplain grazing marsh indicator species.

     

Tree Phenology in Adjacent Amazonian Flooded and Unflooded Forests1

Most phenological studies to date have taken place in upland forest above the maximum flood level of nearby streams and rivers. In this paper, we examine the phenological patterns of tree assemblages in a large Amazonian forest landscape, including both upland (terra firme) and seasonally flooded (várzea and igapó) forest. The abundance of vegetative and reproductive phenophases was very seasonal in all forests types. Both types of flooded forest were more deciduous than terra firme, shedding most of their leaves during the inundation period. Pulses of new leaves occurred mainly during the dry season in terra firme, whereas those in the two floodplain forests were largely restricted to the end of the inundation period. Flowering was concentrated in the dry season in all forest types and was strongly correlated with the decrease in rainfall. The two floodplain forests concentrated their fruiting peaks during the inundation period, whereas trees in terra firme tended to bear fruits at the onset of the wet season. The results suggest that the phenological patterns of all forest types are largely predictable and that the regular and prolonged seasonal flood pulse is a major determinant of phenological patterns in várzea and igapó, whereas rainfall and solar irradiance appear to be important in terra firme. The three forest types provide a mosaic of food resources that has important implications for the conservation and maintenance of wide‐ranging frugivore populations in Amazonian forests.     

Is the field water use of Eucalyptus largiflorens F. Muell. affected by short-term flooding?

Abstract Transpiration of and water sources for Eucalyptus largiflorens F. Muell trees at three sites on a semi-arid floodplain of the lower River Murray in southern Australia were investigated during the course of a flood. Two of the sites were flooded for 32 days while the third was not flooded. At both flooded sites transpiration was not suppressed during the period when the sites were flooded, indicating that anoxia did not occur. Available literature suggests that E. largiflorens has very low O² consumption, and that lateral roots can absorb O² from soil water at very low partial pressures. The oxygen stored in the unsaturated region of the soil appeared sufficient, in this case, to meet the requirements of the trees. Transpiration at the flooded sites did not increase in comparison with that at the non-flooded site in the period following flooding. We suggest that this was possibly due to the above-average rainfall experienced in the months preceding the flood and/or physiological adaptations to the highly saline conditions that prevented the trees taking full advantage of the additional water. The sources of water used by the trees were investigated using the naturally occurring stable isotopes of water and measurements of soil water suction. These showed that prior to the flood shallow rain-derived soil water was being used at all three sites. Ten days after flooding ceased trees at both flooded sites were using shallow flood-derived soil water while at the non-flooded sites they were using groundwater. Results from the flooded sites 7 and 22 weeks after flooding ceased showed that as the flood- and rain-derived water became limited the trees supplemented their supplies with groundwater from the capillary fringe. The non-flooded trees used various combinations of rain-derived soil water and groundwater during this period.     

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.     

Aquatic ecosystem responses to fire and flood size in the Okavango Delta: observations from the seasonal floodplains

The frequency of fires in the Okavango Delta seasonal floodplains peaked at an intermediate frequency of flooding. Floodplains are commonly burnt every 3–5 years. This study showed fundamental changes in ecosystem properties due to burning. A burnt seasonal floodplain in the aquatic phase had oxygen levels well above saturation, 100–200%, while the levels in the un-burnt control site were below saturation and, at night, could decline to 10–40% saturation. The total phosphorous and total nitrogen concentrations were similar on both floodplains but considerably enriched relative to inflowing water, due to nutrient release from the flooded soil-sediment and animal droppings. Zooplankton biomass was very high in both systems although the abundance of fish fry was ten times higher on the un-burnt floodplain. In a low flood year the un-burnt floodplain water had high nutrient levels, primary production, methane emission, and subsequent uptake of methane in biota, as well as a high zooplankton biomass. The very high flood the following year showed the opposite with much lower production at all levels owing primarily to greater dilution of nutrients. The abundance of fish, however, was much higher during the high flood year. Macrophytes and litter provide direct shelter for fish fry but also promote low oxygen levels when decaying. Large flooded areas result in high fish production by removing obstacles related to congestion. This interplay between hydroperiod and fire may be crucial for the maintenance of high biological productivity both in the aquatic and terrestrial phases in a very nutrient poor wetland landscape. Understanding these interactions is crucial for optimal management.     

Early fish growth varies in response to components of the flow regime in a temperate floodplain river

1. The biological productivity of floodplain rivers is intimately related to their flow regimes and it has been proposed that fish production should be linked to components of the flow regime in productivity models. To assess applicability of existing models of productivity in floodplain rivers, we tested predictions about growth during the early life stages of a common, short‐lived fish (Australian smelt Retropinna semoni) in a non‐flow‐altered, temperate Australian floodplain river. 2. The morphometric condition of larval and juvenile fish measured over a five‐year period was positively related to annual discharge, but the highest average seasonal growth rates occurred in two years of contrasting hydrology, one with early spring flooding and the other with predominantly low flows and a late season (within channel) flow pulse. 3. Analysis of daily growth measures indicated that timing, river height, the duration of in‐channel flow events and antecedent flood events are all significant factors influencing the early growth of Australian smelt. The flexible manner in which fish growth responds to these factors appears to be an effective early life history strategy for a short‐lived species occupying a highly variable environment. 4. Growth rates conformed to some predictions of the Flood Pulse Concept (in particular the Extended Flood Pulse Concept), but specific growth responses suggest that the Riverine Productivity Model and tenets of the Low Flow Recruitment Hypothesis best describe the production of Australian smelt in this system. We suggest that none of the existing conceptual models adequately describes fish productivity in temperate Australian floodplain rivers but that aspects of each are likely to be relevant under different flow conditions.     

Pressurized gas transport in Amazonian floodplain trees

Trees of Central Amazonian whitewater floodplain forests are subjected to extended flood periods lasting several months. They must therefore be highly adapted to long-term flooding and to oxygen shortage in the soil and roots which is expected to be the main stress factor in this environment. The improvement of oxygen transport to the roots by pressurized gas transport, known for some tree species of the temperate zone, is one specific adaptation to flooding. To evaluate its significance, gas transport measurements were carried out under experimental conditions with saplings of five Amazonian tree species. Internal aeration of the roots was improved under conditions of pressurized gas transport as shown by polarographical measurements of oxygen exchange between root and rhizosphere. Tracer gas measurements using sulphur hexafluoride (SF₆) showed gas permeability of transport pathways between stem base and roots. Based on the data, we suppose that pressurized gas transport significantly contributes to internal aeration of roots in Amazonian floodplain saplings growing on the higher levels in the flooding gradient, with low water columns, and is an important adaptation for establishment in these temporarily inundated habitats also in the weeks of rising and lowering water levels.     

Effects of the Flooding Gradient on Tree Community Diversity in Várzea Forests of the Purus River,Central Amazon,Brazil

Amazonian floodplain forests have remarkable variation in tree diversity and structure related to flood duration. We assessed how floodplain forests respond to the continuum flooding gradient and found that Fisher's α values vary up to seven times across the landscape.     

The influence of flood cycle and fish predation on invertebrate production on a restored California floodplain

Although floodplains are known to be tightly controlled by the flood cycle, we know comparatively little about how flooding influences predators and their consumption of secondary production, particularly in highly seasonal floodplains typical of Mediterranean climates. In this study, we investigate how the seasonal dynamics of a central California floodplain influence the timing and magnitude of fish predation and the abundance and composition of invertebrates. For 3 years (2000–2002), we compared changes in abundances and size distributions of invertebrates through the flood season (January–June) with seasonal changes in the abundance of larval and juvenile fishes. Using diet analysis of fishes and manipulative feeding experiments with fishes in field enclosures, we link specific changes in invertebrate populations directly to feeding preferences of seasonally abundant fish. Early in the flood season prior to March, we found little influence of fish predation, consistent with the near absence of larval and juvenile fishes during this period. Coinciding with the midseason increase in the abundance of larval and juvenile fishes in April, we found significant declines in zooplankton abundance as well as declines in the size of zooplankton consistent with fish feeding preferences. Our results were consistent with results from feeding enclosure experiments that showed that fish rapidly depressed populations of larger cladocerans with much less effect on smaller cladocerans and calanoid copepods. At the end of the flood season, zooplankton abundances rapidly increased, consistent with a switch in the feeding of juvenile fish to aquatic insects and subsequent fish mortality. We also found that zooplankton biomass on the floodplain reached a maximum 2–3 weeks after disconnection with the river. We suggest that floodplain restoration in this region should consider management strategies that would ensure repeated flooding every 2–3 weeks during periods that would best match the peaks in abundance of native fishes. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Variability of fish diets between dry and flood periods in an arid zone floodplain river

The fish assemblages of an arid zone floodplain river, Cooper Creek, Queensland, Australia, were sampled during two dry periods in isolated waterholes and on the inundated floodplain during the early and late phase of a major flood event. Diets were described for nine native species and compared within and between dry and flood periods. In the dry season, when fishes were restricted to waterholes, diets were characteristically simple with narrow diet breadths. Movement onto the floodplain during flooding clearly increased feeding opportunities, with greater diet breadths evident in all species. Despite obvious potential for terrestrial inputs, diets tended to be dominated by aquatic resources in both the waterholes and on the floodplain. Stomach fullness, however, varied little between dry season waterhole and floodplain samples. Fishes appeared to feed on potentially lower value resources such as detritus and calanoid copepods during the dry season, when waterholes were isolated and food resources were limited. They were then able to capitalize on the 'boom' of aquatic production and more diverse food resources associated with episodic flood events.     

Relationships between vegetation patterns and hydroperiod on the Roanoke River floodplain,North Carolina

This study quantified relationships between forest composition and flooding gradients on the Roanoke River floodplain, North Carolina. Because flooding is highly variable in time and space, the research was designed to determine the specific hydrological parameters that control woody species abundance on the landscape scale. I specifically tested the importance of spring vs. yearly flood duration, as well as flood duration during hydrologically wet vs. dry years. Field vegetation samples of woody species composition were integrated with spatial data from a Landsat Thematic Mapper (TM) classification and a flood simulation model derived in part from synthetic aperture radar (SAR) imagery. Flood simulations were output and summarized for the periods 1912–1950 (before dams were constructed on the river) and 1965–1996 (after all of the dams were completed). Tenth percentile (dry), median, and 90th percentile (wet) hydroperiod (flood duration) regimes were generated for the spring and year, both pre- and post-dam. Detrended correspondence analysis (DCA) was used to ordinate the plot data, and correlation/regression between ordination axis scores and the flood variables were used to explore the relationships between flooding and species composition. Nineteenth percentile hydroperiod (i.e., wet conditions) correlated most strongly with DCA axis 1 (r>0.9), indicating that inundation during extremely wet years strongly controls species composition on the floodplain. The results were used to quantitatively determine the niche width for both species and mapped vegetation classes in terms of number of days flooded annually and during the spring growth period. The results suggest that spring hydroperiod is an important mechanism that may drive competitive sorting along the flooding gradient, especially during the early years of succession (i.e., pre-dam, which represents the period during which most of the forests sampled were established), and that annual hydroperiod affects the relative dominance of species as the forests mature.     

Fish recruitment in a large,temperate floodplain: the importance of annual flooding,temperature and habitat complexity

1. Large river floodplains are considered key nursery habitats for many species of riverine fish. The lower Volga River floodplains (Russian Federation) are still relatively undisturbed, serving as a suitable model for studying the influence of flooding and temperature on fish recruitment in floodplain rivers. 2. We examined the interannual variability in recruitment success of young‐of‐the‐year (YOY) fish in the lower Volga floodplain in relation to flood pulse characteristics and rising water temperatures in the spring. We sampled four areas with different flooding regimes, in three consecutive years (2006–2008). 3. Extensive areas with a long duration of flooding accommodated high densities of young fish. This suggests that extended inundation improves the recruitment success of river fish. In areas with extensive flooding, the biomass of YOY of most fish species was about three times higher in 2006 and 2007 than in 2008. We hypothesise that low spring temperatures in 2008 may have caused this reduced recruitment and that a flood synchronised with rising temperature enhances recruitment success. 4. Extensive flooding was particularly favourable for species characterised by large body size, delayed maturation, high fecundity and low parental investment, such as pike Esox lucius, roach Rutilus rutilus and ide Leuciscus idus. Gibel carp Carassius gibelio, a species tolerant of high temperature and hypoxia, did particularly well in small waterbodies in the driest parts of the floodplain. 5. Structural characteristics of floodplain waterbodies explained much of YOY fish density. These species–environment associations varied from year to year, but some species such as common bream Abramis brama, roach and gibel carp showed consistent relationships with structural habitat characteristics in all years, despite large interannual fluctuations in flood pulse and spring temperature.     

Drought responses of flood-tolerant trees in Amazonian floodplains

Background

Flood-tolerant tree species of the Amazonian floodplain forests are subjected to an annual dry period of variable severity imposed when low river-water levels coincide with minimal precipitation. Although the responses of these species to flooding have been examined extensively, their responses to drought, in terms of phenology, growth and physiology, have been neglected hitherto, although some information is found in publications that focus on flooding.

Scope

The present review examines the dry phase of the annual flooding cycle. It consolidates existing knowledge regarding responses to drought among adult trees and seedlings of many Amazonian floodplain species.

Main Findings

Flood-tolerant species display variable physiological responses to dry periods and drought that indicate desiccation avoidance, such as reduced photosynthetic activity and reduced root respiration. However, tolerance and avoidance strategies for drought vary markedly among species. Drought can substantially decrease growth, biomass and photosynthetic activity among seedlings in field and laboratory studies. When compared with the responses to flooding, drought can impose higher seedling mortality and slower growth rates, especially among evergreen species. Results indicate that tolerance and avoidance strategies for drought vary markedly between species. Both seedling recruitment and photosynthetic activity are affected by drought,

Conclusions

For many species, the effects of drought can be as important as flooding for survival and growth, particularly at the seedling phase of establishment, ultimately influencing species composition. In the context of climate change and predicted decreases in precipitation in the Amazon Basin, the effects of drought on plant physiology and species distribution in tropical floodplain forest ecosystems should not be overlooked.Key words: Drought responses, Amazonia, floodplain forests, tree ecology, várzea     

The impact of experimental sedimentation and flooding on the growth and germination of floodplain trees

Land-use changes in a forested floodplain’s watershed can lead to incremental changes in the hydrology and sedimentation rates of the floodplain. The impacts of these changes can be difficult to measure due to the slow response time of mature trees. Seedlings and saplings, on the other hand, may show an immediate response. Responses during these early life history stages can have major consequences for regeneration of floodplain forests and ultimately result in community alteration. This study tested the importance of changes in hydrology and sedimentation on the germination and growth rates of three common floodplain tree species: Acer rubrum, Fraxinus pennsylvanica and Quercus palustris. Two-year-old saplings were grown in a greenhouse under two hydrologic regimes, with or without the addition of sediment. Neither periodic flooding with or without sediment nor static flooding on its own affected the growth of the seedlings. With the addition of sediment, static flooding for two weeks lead to a significant decrease in sapling growth. There was a significant species x treatment interaction, suggesting that each species responded differently to the application of flooding and sediment. The timing of germination and the total percent germination for F. pennsylvanica and Q. palustris seeds were tested under the same conditions. Flooding and sediment acted in an additive manner to delay the germination of both F. pennsylvanica and Q. palustris and to reduce the total germination rate of Q. palustris. There was no difference in the total germination rate of F. pennsylvanica seeds under any treatment. During the growth trials, adventitious roots sprouted on saplings grown under sedimentation. Adventitious roots growing into sediment rather than floodwater should be able to utilize the sediment’s nutrients and may compensate for some of the stress of flooding. The results of this study suggest that sediment tolerances will vary among species, but will not necessarily correlate with flood tolerances, and that sedimentation may be as important as flooding in determining floodplain plant community composition.     

Base-line variations in stable isotope values in an Arctic marine ecosystem: effects of carbon and nitrogen uptake by phytoplankton

Floods are fundamental for the maintenance of floodplain biodiversity. As a result, well-functioning floodplains are characterized by a high spatio-temporal heterogeneity. Most floodplain-organisms need this shifting landscape mosaic to fulfil their environmental requirements and display a range of adaptations to survive floods. However, in temperate areas, where winter floods are common, extraordinary floods occurring in summer, a period of high physiological activity, may seriously impact the floodplain fauna. This is especially true for guilds characterized by low mobility, such as molluscs. Here we examined the immediate and longer-term response of Elbe grassland molluscs to the extreme 2002 Elbe summer flood in terms of abundance, diversity, and community composition by comparing pre- and post-flood data collected with identical methods. The flood favoured the colonization of aquatic species and led to a shift of the community towards a more hydrophilic composition. Both diversity and abundance increased significantly in the first year following the flood but decreased later gradually to the pre-flood levels. The high spatio-temporal habitat heterogeneity played an important part in the maintenance of mollusc diversity by increasing refuge opportunities and favouring the maintenance of various mollusc communities with different environmental requirements within the floodplain. Handling editor: S. I. Dodson     

The effects of nutrient pulsing on the threatened,floodplain species,Boltonia decurrens

Prior to anthropogenic disturbance, the floodplains of the Illinois River (USA) experienced moderate, cyclical flooding; a phenomenon termed a 'flood pulse'. Significant changes in the hydrology of the Illinois River caused by human alteration of the river channel resulted in a decline in populations of Boltonia decurrens, a native, herbaceous floodplain species of the Illinois River Valley and the demography of the species and underlying mechanisms causing the severe decline of B. decurrens are currently being studied. One component of the river flood pulse is the increased availability of nutrients in floodplain soils. We studied the effects of increased nutrient availability on growth and reproduction of B. decurrens and Aster pilosus, a species replacing B. decurrens in the floodplain. In this study, we examined the effects of a spring nutrient pulse and a late summer nutrient pulse, as compared to a control treatment, on rosettes of B. decurrens and A. pilosus grown in three soil types: sandy loam, silty loam and clay. Biomass, plant height, inflorescences per plant, seed production, and seed mass were measured and compared. Individuals of B. decurrens grown in sandy loam, the predominant soil type of the Illinois River Valley, had thegreatest increase in growth and fecundity in response to nutrient pulsing. Total biomass and inflorescences per plant were significantly greater than the control regardless of the timing of the nutrient pulse. Aster pilosus did not exhibit a clear pattern of increased growth and fecundity in a specific soil type or pulse treatment. The early pulse treatment appeared to have a greater effect across the range of soil types than the late pulse treatment; however, it did not seem to influence any particular variable(s) that we measured. A pulse of nutrients benefits B. decurrens more than A. pilosus, particularly in sandy loam. The increased vigor and fecundity of B. decurrens with a nutrient pulse could be a factor contributing to population survival during the period between disturbance events.     

Habitat specifity,endemism and the neotropical distribution of Amazonian white‐water floodplain trees

.The Amazon basin is covered by the most species‐rich forests in the world and is considered to house many endemic tree species. Yet, most Amazonian ecosystems lack reliable estimates of their degree of endemism, and causes of tree diversity and endemism are intense matters of debate. We reviewed the spatial distribution of 658 of the most important flood‐tolerant Amazonian white‐water (várzea) tree species across the entire Neotropics by using data from herbaria, floras, inventories and checklists. Our results show that 90% of the várzea tree species are partially or widely distributed across neotropical macro‐regions and biomes. Chi‐square analyses indicated that várzea species richness in non‐várzea macro‐regions was dependent on the flooding gradient and the longitudinal position. Cluster analysis combined with association tests indicated four significant patterns of várzea species distributions depending on species flood‐tolerance (low vs high) and spatial distribution (restricted vs widespread). We predict that the predominance of Andean substrates is the most important factor that determines the distribution of várzea tree species within and beyond the Amazon basin and explains the high floristic similarity to the Orinoco floodplains. Distribution patterns in other extra‐Amazonian macro‐regions are more likely linked to climatic factors, with rainforest climates housing more várzea species than savanna climates. 130 tree species were restricted to South‐American freshwater floodplains, and 68 (> 10%) were endemic to Amazonian várzea. We detected two centers of endemism, one in the western Amazon characterized by low and brief floods, and one in the central Amazon, characterized by high and prolonged floods. Differences in taxonomic composition of endemic centers in the western and central Amazon are the result of different abiotic factors (i.e. flood regimes), as well as the regional species pools from where the species are recruited from. We hypothesize that numerous morphological, physiological and biochemical adaptations permit survival of trees in flooded environments. Furthermore, these adaptations are independently derived across many taxa and result in a highly specialized flora. We attribute higher than expected levels of endemism to the great spatial extent and age of floodplain ecosystems in the Amazon basin, and highlight the role of Amazonian várzea as an potential driver in speciation and diversification processes.