Review
Winner–Loser Species Replacements in Human-Modified Landscapes

https://doi.org/10.1016/j.tree.2021.02.006Get rights and content

Highlights

  • Winner–loser species replacements (WLRs) result primarily from changes in available resources rather than from altered competition among species.

  • Species dispersal controls community assembly, and both WLRs and community reorganization involve species spread at multiple scales.

  • WLRs result in the functional reorganization of assemblages owing to the proliferation of 'supertramp' species, including non-forest species, that operate as human commensals.

  • Breaches in biome boundaries facilitated by human-assisted colonization represent a 'one-way street' for the spreading of species into abiotically suitable environments.

  • Transitions toward functionally novel assemblages result in new 'packages' of both ecosystem services and disservices provided by winners.

  • WLRs, novel assemblages, and the provision of services and disservices represent a valuable framework to understand the multiple transitions that tropical forests are experiencing.

Community assembly arguably drives the provision of ecosystem services because they critically depend on which and how species coexist. We examine conspicuous cases of 'winner and loser' replacements (WLRs) in tropical forests to provide a framework integrating drivers, impacts on ecological organization, and reconfiguration of ecosystem service provisioning. Most WLRs involve native species and result from changes in resource availability rather than from altered competition among species. In this context, species dispersal is a powerful force controlling community (re)assembly. Furthermore, replacements imply a nearly complete functional reorganization of assemblages and new 'packages' of ecosystem services and disservices provided by winners. WLRs can thus elucidate the multiple transitions experienced by tropical forests, and have theoretical/applied implications, including the role that human-modified landscapes may play in global-scale sustainability.

Section snippets

Biodiversity Responses to Human Disturbances

Community assembly at multiple spatial scales has been a central topic in community ecology [1,2], but has become increasingly relevant because the provision of ecosystem services (see Glossary) may crucially depend on which and how species coexist at local to landscape scales. This is particularly true for ecosystems that are intensely exposed to human disturbances, including land-use change and overharvesting of natural resources. For instance, the functional composition of plant assemblages

Land-Use Intensification and WLRs

Tropical forests worldwide are experiencing land-use intensification through similar trajectories involving conversion of old-growth forest into human-modified landscapes. We define here human-modified landscapes as mosaics typically composed of (i) a few remaining patches of old-growth forest, (ii) myriad small edge-dominated forest fragments of varying sizes, (iii) second-growth stands of varying age, and (iv) exotic tree plantations and agroforest patches. All of these are typically embedded

Mechanisms of Population Decline or Proliferation

WLRs involve population declines particularly as a result of (i) ecological filtering of forest-dependent species because physiological constraints prevent the use of open, highly illuminated, and desiccated edge-affected habitats [30,51,52], and (ii) reduced availability of key resources associated with old-growth forests [53], such as large trees [54] and their flowers and fruits [55]. Population depletion via direct overexploitation of forest products represents a complementary and often

WLRs and Community (Re)Assembly

WLRs reorganize local assemblages in terms of species richness and abundance [31,63], taxonomic and ecological composition [25,55], and phylogenetic structure [64,65]. However, because disturbance-adapted species can be diverse, WLRs do not necessarily result in impoverished or homogenous communities, at least at the landscape scale. Human-modified landscapes can support, at least temporarily, diverse and distinct assemblages, with alpha and beta diversity scores similar to those of old-growth

Functional Transitions, Ecosystem Services and Disservices

WLRs represent a widespread phenomenon across temperate and tropical landscapes that are (i) becoming increasingly hyper-disturbed, (ii) recovering many of their baseline conditions, or/and (iii) experiencing climate change, particularly higher temperatures [71,72]. In sum, the findings examined here highlight the emergence of novel tropical forest that are functionally distinct, but are not necessarily more impoverished for some taxa. This diffuse biotic reorganization largely relies on a

Concluding Remarks

Globally it is unlikely that hyper-fragmented landscapes across consolidated agricultural regions will soon return to higher levels of forest cover, habitat connectivity, and habitat heterogeneity; in other words, the conditions that are necessary to retain species assemblages closer to historical arrangements (reviewed in [38]). Furthermore, many tropical landscapes can gradually accumulate forest cover because of societal demographic transitions such as rural–urban migration [76]. Finally, we

Acknowledgments

This review was funded by the Coordenação de Aperfeiçoamento de Pessoal de Nivel Superior (CAPES, Finance code 001), the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, grants 403770/2012-2, 441386/2016-4), the Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco (FACEPE, grant APQ-0138-2.05/14), and a Newton Fund Institutional Partnership Award between the University of East Anglia and the Universidade Federal de Pernambuco. B.K.C.F. thanks the CNPq for a

Declaration of Interests

No interests are declared.

Glossary

Agroforestry
land use management system in which forests coexist with crops or pastureland.
Biotic homogenization
the systematic replacement of high-diversity biotas by low-diversity and more similar biotas.
CO2 fertilization
the increased rate of photosynthesis in plants that results from increased levels of CO2 in the atmosphere.
Ecosystem services
direct and indirect contributions of ecosystems to human well-being.
Edge-affected habitats
forest habitats containing high amounts of forest edges but

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