One sixth of Amazonian tree diversity is dependent on river floodplains.

Amazonia's floodplain system is the largest and most biodiverse on Earth. Although forests are crucial to the ecological integrity of floodplains, our understanding of their species composition and how this may differ from surrounding forest types is still far too limited, particularly as changing inundation regimes begin to reshape floodplain tree communities and the critical ecosystem functions they underpin. Here we address this gap by taking a spatially explicit look at Amazonia-wide patterns of tree-species turnover and ecological specialization of the region's floodplain forests. We show that the majority of Amazonian tree species can inhabit floodplains, and about a sixth of Amazonian tree diversity is ecologically specialized on floodplains. The degree of specialization in floodplain communities is driven by regional flood patterns, with the most compositionally differentiated floodplain forests located centrally within the fluvial network and contingent on the most extraordinary flood magnitudes regionally. Our results provide a spatially explicit view of ecological specialization of floodplain forest communities and expose the need for whole-basin hydrological integrity to protect the Amazon's tree diversity and its function.

The Brazilian freshwater wetscape: Changes in tree community diversity and composition on climatic and geographic gradients.

Wetlands harbor an important compliment of regional plant diversity, but in many regions data on wetland diversity and composition is still lacking, thus hindering our understanding of the processes that control it. While patterns of broad-scale terrestrial diversity and composition typically correlate with contemporary climate it is not clear to what extent patterns in wetlands are complimentary, or conflicting. To elucidate this, we consolidate data from wetland forest inventories in Brazil and examine patterns of diversity and composition along temperature and rainfall gradients spanning five biomes. We collated 196 floristic inventories covering an area >220 ha and including >260,000 woody individuals. We detected a total of 2,453 tree species, with the Amazon alone accounting for nearly half. Compositional patterns indicated differences in freshwater wetland floras among Brazilian biomes, although biomes with drier, more seasonal climates tended to have a larger proportion of more widely distributed species. Maximal alpha diversity increased with annual temperature, rainfall, and decreasing seasonality, patterns broadly consistent with upland vegetation communities. However, alpha diversity-climate relationships were only revealed at higher diversity values associated with the uppermost quantiles, and in most sites diversity varied irrespective of climate. Likewise, mean biome-level differences in alpha-diversity were unexpectedly modest, even in comparisons of savanna-area wetlands to those of nearby forested regions. We describe attenuated wetland climate-diversity relationships as a shifting balance of local and regional effects on species recruitment. Locally, excessive waterlogging strongly filters species able to colonize from regional pools. On the other hand, increased water availability can accommodate a rich community of drought-sensitive immigrant species that are able to track buffered wetland microclimates. We argue that environmental conditions in many wetlands are not homogeneous with respect to regional climate, and that responses of wetland tree communities to future climate change may lag behind that of non-wetland, terrestrial habitat.

Phenology, fruit production and seed dispersal of Astrocaryum jauari (Arecaceae) in Amazonian black water floodplains

Astrocaryum jauari Mart. (Arecaceae) is one of the commonest palm species occurring in nutritionally poor Amazonian black water floodplains. It is an emergent or subcanopy tree that grows on river banks and islands, with a wide distribution along the entire flooding gradient, tolerating flood durations between 30 and 340 days. The species is important for fish nutrition in the floodplains, and is also used for hearts of palm. In the present study, the auto-ecology of A. jauari was analysed over a period of two years in the Anavilhanas Archipelago, Rio Negro, Brazil, with a focus on phenology, fruit production, and seed dispersal. Fruit fall is annual and synchronized with high water levels, with a production of 1.6 ton of fruit ha(-1). The fruits are eaten by at least 16 species of fish which either gnaw the pulp, fragment the seed, or ingest the entire fruit, thus acting as dispersal agents. Besides ichthyocory, barochory (with subsequent vegetative propagation) is an important dispersal mode, enhancing the occurrence of large masses of individuals in the Anavilhanas islands and in the region of maximum palm heart extraction near Barcelos.

Astrocaryum jauari Mart. (Arecaceae) es una de las especies más comunes de palma en las llanuras de inundación por las llamadas “aguas negras”, aguas ricas en taninos que tienen pocos nutrientes para la fauna. Habita el subdosel que se desarrolla en riberas e islas, con una distribución amplia en toda la gradiente de inundación (resiste entre 30 y 340 días bajo el agua). La especie es importante para la nutrición de los peces y en la producción de palmito. La autoecología de A. jauari fue analizada por dos años en el Archipiélago Anavilhanas, río Negro, Brazil, con énfasis en fenología, producción de frutas, y dispersores de semillas. La caída de los frutos es anual y sincronizada con el aumento de los niveles de agua, con una producción de 1.6 ton de fruta ha-1. Las frutas son comidas por al menos de 16 especies de peces que roen la pulpa o fragmentos de semilla, o ingieren la fruta entera y actúan como agentes dispersores. Además de la ictiocoria, la barocoria y la subsecuente progragación asexual son básicos para la alta densidad de la especie.

Phenology, fruit production and seed dispersal of Astrocaryum jauari (Arecaceae) in Amazonian black water floodplains.

Astrocaryum jauari Mart. (Arecaceae) is one of the commonest palm species occurring in nutritionally poor Amazonian black water floodplains. It is an emergent or subcanopy tree that grows on river banks and islands, with a wide distribution along the entire flooding gradient, tolerating flood durations between 30 and 340 days. The species is important for fish nutrition in the floodplains, and is also used for hearts of palm. In the present study, the auto-ecology of A. jauari was analysed over a period of two years in the Anavilhanas Archipelago, Rio Negro, Brazil, with a focus on phenology, fruit production, and seed dispersal. Fruit fall is annual and synchronized with high water levels, with a production of 1.6 ton of fruit ha(-1). The fruits are eaten by at least 16 species of fish which either gnaw the pulp, fragment the seed, or ingest the entire fruit, thus acting as dispersal agents. Besides ichthyocory, barochory (with subsequent vegetative propagation) is an important dispersal mode, enhancing the occurrence of large masses of individuals in the Anavilhanas islands and in the region of maximum palm heart extraction near Barcelos.

Wood growth patterns of Macrolobium acaciifolium (Benth.) Benth. (Fabaceae) in Amazonian black-water and white-water floodplain forests.

Macrolobium acaciifolium (Benth.) Benth. (Fabaceae) is a dominant legume tree species occurring at low elevations of nutrient-poor black-water (igapó) and nutrient-rich white-water floodplain forests (várzea) of Amazonia. As a consequence of the annual long-term flooding this species forms distinct annual tree rings allowing dendrochronological analyses. From both floodplain types in Central Amazonia we sampled cores from 20 large canopy trees growing at identical elevations with a flood-height up to 7 m. We determined tree age, wood density (WD) and mean radial increment (MRI) and synchronized ring-width patterns of single trees to construct tree-ring chronologies for every study site. Maximum tree age found in the igapó was more than 500 years, contrary to the várzea with ages not older than 200 years. MRI and WD were significantly lower in the igapó (MRI=1.52+/-0.38 mm year(-1), WD=0.39+/-0.05 g cm(-3)) than in the várzea (MRI=2.66+/-0.67 mm year(-1), WD=0.45+/-0.03 g cm(-3)). In both floodplain forests we developed tree-ring chronologies comprising the period 1857-2003 (n=7 trees) in the várzea and 1606-2003 (n=13 trees) in the igapó. The ring-width in both floodplain forests was significantly correlated with the length of the terrestrial phase (vegetation period) derived from the daily recorded water level in the port of Manaus since 1903. In both chronologies we found increased wood growth during El Niño events causing negative precipitation anomalies and a lower water discharge in Amazonian rivers, which leads to an extension of the terrestrial phase. The climate signal of La Niña was not evident in the dendroclimatic proxies.

Internal oxygen transport in cuttings from flood-adapted várzea tree species.

Tree species from the Central Amazon inundation areas are subjected to extreme flooding, with trees being partially submerged for up to 10 months. The rapidly advancing floodwater table at the onset of the aquatic phase interrupts the inward diffusion of oxygen from the atmosphere to submerged plant parts. Salix martiana (Leyb.) (Salicaceae) and Tabernaemontana juruana ((Markgr.) Schumann ex J. F. Macbride) (Apocynaceae), tree species typical of Amazon floodplains, respond to low oxygen concentrations by forming adventitious roots capable of longitudinal oxygen transport. Cuttings of these tree species were subjected to simulated flooding and the oxygen concentration of the root cortex was temporally monitored by oxygen microelectrodes that penetrated the roots. Changes in the floodwater table made it possible to localize precisely the entry points of atmospheric oxygen. Under experimental conditions, mathematical description of the transport kinetics revealed that longitudinal transport of O2 in both species was mainly attributable to diffusion. Based on the finding that diffusion was inhibited by a small increase in the floodwater table, we conclude that internal oxygen transport during a rising water table is only attainable when adventitious roots are continuously and rapidly developed, as is the case in S. martiana. In T. juruana, slow growth of adventitious roots and low root porosity suggest that other adaptations are required to overcome long flooding periods.

Apoplasmic barriers and oxygen transport properties of hypodermal cell walls in roots from four amazonian tree species.

The formation of suberized and lignified barriers in the exodermis is suggested to be part of a suite of adaptations to flooded or waterlogged conditions, adjusting transport of solutes and gases in and out of roots. In this study, the composition of apoplasmic barriers in hypodermal cell walls and oxygen profiles in roots and the surrounding medium of four Amazon tree species that are subjected to long-term flooding at their habitat was analyzed. In hypodermal cell walls of the deciduous tree Crateva benthami, suberization is very weak and dominated by monoacids, 2-hydroxy acids, and omega-hydroxycarboxylic acids. This species does not show any morphological adaptations to flooding and overcomes the aquatic period in a dormant state. Hypodermal cells of Tabernaemontana juruana, a tree which is able to maintain its leaf system during the aquatic phase, are characterized by extensively suberized walls, incrusted mainly by the unsaturated C(18) omega-hydroxycarboxylic acid and the alpha,omega-dicarboxylic acid analogon, known as typical suberin markers. Two other evergreen species, Laetia corymbulosa and Salix martiana, contained 3- to 4-fold less aliphatic suberin in the exodermis, but more than 85% of the aromatic moiety of suberin are composed of para-hydroxybenzoic acid, suggesting a function of suberin in pathogen defense. No major differences in the lignin content among the species were observed. Determination of oxygen distribution in the roots and rhizosphere of the four species revealed that radial loss of oxygen can be effectively restricted by the formation of suberized barriers but not by lignification of exodermal cell walls.

Impact of root morphology on metabolism and oxygen distribution in roots and rhizosphere from two Central Amazon floodplain tree species.

Adaptation to prolonged flooding was investigated using cuttings of two tree species from the Central Amazon white-water floodplain (Várzea). Morphological features and oxygen distribution patterns were correlated with metabolic changes under hypoxia, such as alterations in alcohol dehydrogenase (ADH) activity and adenylate energy charge (AEC) of root cells. Salix martiana (Leyb.) was able to react to hypoxic growth conditions with formation of adventitious roots rich in lysigenous aerenchyma, which facilitates root aeration by longitudinal oxygen transport and rhizosphere oxidation by radial oxygen loss (ROL). The oxygen concentration on the surface of adventitious roots of S. martiana reached 2-3 mg O2 L-1. The low resistance to gas exchange in Salix roots was reflected by low ADH activities, which ranged between 0.03-0.1 µmol NADH mg -1 min-1, and AEC values of 0.8-1 under hypoxic conditions. Adventitious roots were also formed by Tabernaemontana juruana ([Markgr.] Schumann ex. J.F. Macbride) during growth under low-oxygen conditions, although at a later stage. The gas-space continuum in roots of T. juruana was less pronounced, resulting in a 10-fold lower oxygen concentration in the root cortex under oxygen stress compared with adventitious roots of Salix. The lower oxygen content was reflected in 6-fold higher ADH activities and decreased AEC values. ROL occurred only at the non-suberized root tip, suggesting that the suberized hypodermis functions as a barrier against gas exchange between the root and the rhizosphere. These findings indicate that different strategies of adaptation to low oxygen levels are realized in the two species under investigation that occur naturally in the same ecosystem but inhabit different elevation sites.