Our results suggest that açaí stems and clumps density (i.e., a measure of açaí management intensification) reorganize woody plant assemblages in the estuarine forest landscapes devoted to açaí fruit production. Açaí intensification reduces the density and alfa taxonomic diversity of both adult woody plant assemblages inhabiting forest canopy and the emergent layer, but also of those assemblages inhabiting forest understory. We refer to assemblages by small-statured tree species, palms and large shrubs. In fact, such understory assemblage appears to be much more diverse than canopy and emergent assemblages, as they combine individuals from species occurring across all forest vertical habitats. On the other hand, açaí intensification appear to increment taxonomic beta diversity, particularly due to species turnover in response to reduction on alpha diversity and increasing levels of rarity as açaí density increases. The estuarine forest also supports an ecologically diverse team of sensitive species (from large trees to shrubs), which decline before the regulatory threshold of 400 açaí clumps ha, while a reduced number of species respond positively. Accordingly, açaí intensification reorganize plant assemblages taxonomically but also ecologically towards impoverished species sets, probably including the assemblages responsible for forest regeneration and structural integrity (i.e., saplings from canopy and emergent tree species).
Our findings reinforce the notion that açaí intensification promotes de establishment of low-density and impoverished adult tree assemblages, which become taxonomically distinct as açaí density increases as already documented (Freitas et al., 2015, 2021). However, we offer evidence suggesting that species loss among adult tree assemblages occurs across forest stands still supporting relatively low-density açaí stands; species loss occurs as soon as intensification starts to alter açaí abundance. On the other hand, few species respond positively to açaí intensification; i.e., more losers than winners sensu Tabarelli et al. (2012). Furthermore, our findings indicate the occurrence of a diverse woody plant assemblage inhabiting forest understory, which combines saplings from canopy tree species, but also a variety of large shrubs, small-statured tree and palm species; i.e., a typical understorey flora across tropical forests (Whitmore, 1984; Gentry and Emmons, 1987). This neglected forest component in açaí-related studies, but not necessarily less diverse than the canopy woody flora, appears to be negatively affected by açaí intensification and thus call attention to other forest components being impacted by açaí intensification. In fact, a reasonable portion of the Amazonian woody plant flora consists of shrubs, small-statured trees and palms inhabiting forest understory by being shade-tolerant species; i.e., the whole life-cycle in the shaded forest understory (Bloor and Grubb, 2003). The few estimates available report a minimum 16% of tree species flora belonging to this ecological group in the Amazon region (Assis and Wittmann 2011). However, by considering herbs, shrubs and small palms, the understory flowering species flora is expected to achieve over 20% of all species (see Costa and Magnusson 2002). Unfortunately, there is little information about the Amazon estuarine forest, even for the tree flora, although this unique forest represents the core area of açaí production and intensification (Brondízio, 2008) and an ecologically unique component of an area of species endemism considering an immense number of taxa, from insects to primates (Moraes et al., 2020; Silva et al., 2022).
Although we did not explicitly address the baseline mechanisms driving the changes in plant assemblages, it has been argued the elimination of undesirable trees in order to provide better microclimatic conditions and the space demand to increment clump density is the main driver for the impoverishment tree assemblages experience as açaí clump density increases (Freitas et al., 2015, 2021). Such a forest thinning or coppicing is one of the prescribed management procedures in order to increment açaí fruit productivity (Anderson, 1988) and in a large extent responds for a complete degradation of forest spatial structure and the impoverishment of tree assemblages, although it has never been experimentally tested. In this context, decreasing species richness and increasing rarity as açaí density increases is probably the mechanism behind higher beta diversity as intensification proceeds, a similar phenomenon in response to habitat loss and fragmentation resulting in low gamma diversity as already documented; i.e., species richness at landscape/regional level (Sfair et al., 2016).
In fact, it is reasonable to propose that changes promoted by forest thinning in forest physical structure and woody plant assemblages will alter a myriad of processes limiting further plant recruitment upon which thinning operations will occur. Three examples, elimination of large trees probably results into much more illuminated habitats filtering shade-tolerant species while favoring light demanding such as sun-loving grasses and açaí recruitment (Rutishauser et al., 2016; Pinho et al., 2020). Similarly, the elimination of large buttressed trees, including their large dead trunks over the ground, is likely to increase seed runoff during water descending (Junk et al., 2010). Increased runoff causing lower soil fertility is another potential effect by “açaization”, the local expression to describe intensification leading to the emergence of monospecific stands, which are not permitted according to current regulation but continue to proliferate (Hiraoka, 1993; Freitas et al., 2021). As we documented here soil fertility is positively associated with several community- and ecosystem-attributes across tropical forests, including species richness, forest productivity and aboveground biomass with cascading effects relative to support for herbivores/frugivores assemblages (Gentry and Emmons, 1987; Peres, 2008; Cleveland at al., 2011). It is interesting that forest stand accessibility was positively with adult species richness. Such a priori unexpected result may in fact result from a higher number of useful species being spared close to households such as Theobroma cacao, Hevea brasiliensis, Spondias mobim, Mauritia flexuosa (Nogueira et al., 2005; Tregidgo et al., 2020). Additionally, açaí dropped fronds may cover seedlings on the ground and reduce recruitment of woody plant species as already documented for other palm species (Peters et al., 2004). Finally, there is evidence suggesting that açaí intensification disturbs plant pollination (Campbell et al., 2018) and seed dispersal by frugivorous vertebrates (Moegenburg and Levey, 2002). We refer to both abiotic and biotic changes affecting plant recruitment and then contributing to forest impoverishment as “managed forest/landscapes” experience the intensification trajectory.
In this context, the current regulation requiring a maximum of 400 açaí clumps per ha plus the persistence of 50 palm trees of other species and 200 trees hardwoods, 40 thick (> 45 cm DBH), 40 medium (20 to 45 cm DBH) and 120 thin (5 to 20 cm DBH) is highly welcome (see Queiroz and Mochiutti, 2001). However, to acknowledge the açaí management as sustainable and then stimulate legislation compliance by riverine traditional producers, it is highly recommendable to examine in which extent these thresholds are able to guarantee forest integrity considering and all ecosystem services this forest provide, from biodiversity persistence to the apport of organic material (e.g., leaves, flowers, fruits, seeds) to a highly productive estuarine system (Sodré et al., 2011). Accordingly, it is required a reference ecosystem sensu (Suganuma and Durigan, 2015) as a baseline (probably landscapes still covered by old-growth forests) to investigate forest changes and responses to the whole package involving the açaí management as it involves more forest thinning and increments on açaí density.
It is true the estuarine forest has long been submitted to the extraction forest products, including management to favor cocoa, rubber tree and açaí; i.e., a typical land-sharing approach (Green et al., 2005). Like in the past, such a “disturbed forest” currently experiencing açaí intensification continues to be exposed to a management exclusively restricted to açaí forestry/agricultural demands rather than a management able to guarantee both forest products but also forest integrity. Such a broad perspective of forest management is likely to demand land-use zoning and the establishment of strictu sensu protected areas in addition to more comprehensive and robust set of guidelines to manage açaí stands considering multiple spatial scales (e.g., land-sparing vs. land-sharing). This more ambitious approach remains neglected due a lack of (1) a substantial research effort focused on the multiple impacts by açaí intensification (from local to regional scales) such as the collapse of the understory flora, and (2) a detailed reference on the estuarine forest relative to patterns of species diversity, community organization, physical structure and ecosystem functioning, including primary productivity and biomass.
In synthesis, we shall recognize that rather than the extension of the flooding period (Parolin and Junk, 2002), açaí intensification currently represents the major driver of plant species assembly and community organization operating across several floodplain forests. Applied, açaí intensification moves forward into a “blind context” and thus it can be considered a major threat to estuarine/floodplain forests, offering a disturbing message about the role played by NTFPs as an opportunity for sustainability. In fact, the “açaí case” inform us that NTFPs can only be sustainably extracted in the case appropriate public policies/regulations are incorporated to guarantee traditional low impact forest management approaches rather than market-oriented management stimulating intensification.