Mechanisms of Coexistence and Co-occurrence in Dry Forest Tree Communities: A Systematic Synthesis

Mechanisms of Coexistence and Co-occurrence in Dry Forest Tree Communities: A Systematic Synthesis | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 18 February 2026 V1 Latest version Share on Mechanisms of Coexistence and Co-occurrence in Dry Forest Tree Communities: A Systematic Synthesis Authors : Carlos Salas-Macías , Karime Montes-Escobar , José Macías-Barberán , María Vinces Obando 0000-0002-3310-6202 [email protected] , Luis Balarezo-Saltos , and Carolina Fonseca-Restrepo Authors Info & Affiliations https://doi.org/10.22541/au.177138555.52631527/v1 126 views 46 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Species coexistence in seasonally dry tropical forests (SDTF) emerges from interacting mechanisms operating across spatial and temporal scales, yet empirical indicators that capture these processes remain fragmented and rarely inform forest condition assessments. This systematic review synthesizes 108 studies that explicitly link tree coexistence mechanisms with taxonomic, functional, phylogenetic and co‑occurrence‑based indicators across SDTF worldwide. Functional indicators dominate current practice (46% of studies), followed by co‑occurrence metrics (31%), whereas purely taxonomic and phylogenetic indicators each account for only ~14%, despite their widespread use in broader biodiversity monitoring. The review shows that water‑related environmental filters, drought–shade trade‑offs, successional dynamics and biotic interactions such as Janzen–Connell effects and facilitation are recurrently inferred but quantified with heterogeneous metrics and study designs. By organizing indicators into classes aligned with coexistence mechanisms and evaluating their sensitivity to hydrological, edaphic and disturbance gradients, the review identifies a core set of coexistence‑oriented metrics that can be embedded in forest and ecosystem condition frameworks. These include components of β‑diversity, functional trait spectra and β‑diversity along water and successional gradients, and co‑occurrence‑based indicators of keystone or indicator species. Integrating these indicators into standardized condition frameworks would help incorporate SDTF into national and global monitoring schemes, ensuring that coexistence processes, resilience and restoration outcomes are explicitly considered in forest condition reporting. Introduction Seasonally dry tropical forests (SDTF, hereafter dry forests) harbor high levels of tree diversity and endemism, yet they are among the most threatened ecosystems worldwide due to agricultural expansion, logging and climate change (Miles et al., 2006; Sánchez‑Azofeifa et al., 2005). Despite representing a substantial fraction of tropical forest cover and providing critical ecosystem services, dry forests remain comparatively understudied relative to ever‑wet tropical rainforests (Chazdon, 2008; Chazdon et al., 2016). Rapid habitat conversion and increasing climatic variability in these regions have renewed interest in understanding how tree species assemble, coexist and persist under pronounced seasonal water limitation, with direct implications for biodiversity conservation, restoration and the development of robust ecological indicators (Allen et al., 2010; Sánchez‑Azofeifa et al., 2005). A central question in community ecology is how numerous tree species can coexist within the same local assemblage (Tilman, 1982; Chesson, 2000). Classical niche theory proposes that coexistence is promoted by stabilizing mechanisms such as resource partitioning, habitat specialization and life‑history trade‑offs (Hutchinson, 1959; Tilman, 1982). In contrast, neutral theory assumes that species are functionally equivalent and that community composition is largely shaped by stochastic demographic events, dispersal limitation and ecological drift (Hubbell, 2001). Empirical work in tropical forests increasingly suggests that niche‑based and neutral processes act jointly, with their relative importance varying across spatial and temporal scales, life stages and disturbance regimes (Brown et al., 2013; Clark, 2010; Wiegand et al., 1999). However, most synthetic efforts have focused on humid tropical forests; in dry forests, the relative contributions of specific coexistence mechanisms and their signatures in commonly used community metrics are still poorly synthesized. Dry forests provide a particularly informative context to examine coexistence because strong seasonality in water availability, pronounced edaphic heterogeneity and recurrent disturbance are expected to generate sharp environmental filters and strong abiotic stress (Borchert, 1994; Quesada et al., 2009). At local scales, topographic and edaphic gradients structure seedling and adult distributions, fostering habitat specialization and functional differentiation among coexisting tree species (Balvanera et al., 2002; Pineda‑García et al., 2011). Trait‑based studies in tropical dry forests have documented trade‑offs between water acquisition and drought tolerance, variation in turgor loss point, and differences in leaf and wood traits that underpin functional niche partitioning along water‑stress gradients (Markesteijn & Poorter, 2009; Pineda‑García et al., 2011; Bartlett et al., 2012). At the same time, evidence of strong dispersal and recruitment limitation, density‑dependent processes and context‑dependent successional trajectories points to an important role of non‑deterministic and neutral‑like mechanisms in shaping community assembly (Buzzard et al., 2016; Bongers et al., 2015; Werden et al., 2020). Understanding how these processes are reflected in measurable patterns of diversity, traits and co‑occurrence is essential for designing indicators that capture the state and dynamics of dry forest tree communities. Patterns of species co‑occurrence and associated diversity metrics provide a complementary window into underlying coexistence mechanisms. Analyses of spatial associations and co‑occurrence networks in tropical forests show that both aggregated and segregated species pairs can emerge from the interplay of environmental filtering, biotic interactions and historical contingencies (Gotelli & McCabe, 2002; Wiegand et al., 1999). In tropical dry forests, co‑occurrence analyses and indicator‑species approaches have revealed that some keystone or structurally dominant tree species consistently co‑occur with particular assemblages, whereas others tend to be spatially segregated, suggesting the joint influence of habitat preferences, facilitation and competitive exclusion (Carrión, 2017; Macías‑Mora et al., 2025). Yet, interpreting co‑occurrence patterns remains challenging because similar structures can arise from multiple, non‑exclusive processes and because results depend on spatial scale, null models and the choice of metrics (Blüthgen et al., 2008; Arita, 2012). For the purposes of ecological assessment and monitoring, there is thus a need to clarify which community‑level indicators (e.g. beta‑diversity partitions, trait‑based indices, network metrics) most consistently reflect specific coexistence mechanisms in dry forests. Recent work has started to integrate taxonomic, functional and phylogenetic dimensions of diversity along successional and environmental gradients in dry forests, offering candidate indicators of assembly processes and ecosystem recovery. Studies along secondary successional trajectories in seasonally dry tropical forests show directional shifts in functional trait composition, changes in functional diversity and turnover, and varying importance of abiotic filtering versus biotic interactions over time (Buzzard et al., 2016; Lohbeck et al., 2014; Bongers et al., 2015). Other research has combined niche modelling, functional traits and phylogenetic structure to explore how environmental filters and evolutionary history jointly shape tree distributions (Derroire et al., 2016; Arango et al., 2021). In parallel, analyses of multidimensional tree niches in tropical dry forests suggest that many species exhibit broadly overlapping niches, with limited evidence for strong niche segregation beyond broad autecological differences (Pulla et al., 2017; Brown et al., 2013). Collectively, these studies indicate that dry forest tree communities are structured by a complex interplay of deterministic and stochastic forces, but a cross‑study, indicator‑oriented synthesis is still missing. In this context, a systematic, mechanism‑focused synthesis of empirical evidence on coexistence and co‑occurrence in dry forest tree communities can help bridge ecological theory and applied monitoring. The present study addresses this gap by conducting a systematic synthesis of empirical studies published between 2000 and 2025 that explicitly or implicitly investigate mechanisms of coexistence and patterns of co‑occurrence in seasonally dry tropical and subtropical forest tree communities worldwide. Specifically, this review aims to: (1) identify the main classes of mechanisms—niche‑based, neutral and hybrid—invoked to explain tree species coexistence in dry forests; (2) synthesize the empirical support for these mechanisms across spatial scales, successional stages and environmental gradients; and (3) evaluate how different analytical approaches and community‑level metrics (taxonomic, functional, phylogenetic and network‑based) influence inferences about underlying processes. By adopting an indicator‑oriented perspective, this review seeks to clarify when and where particular coexistence pathways dominate, identify robust candidate indicators of assembly mechanisms and ecosystem condition, and inform the design of monitoring and restoration strategies in dry forest ecosystems under ongoing global change (Chazdon et al., 2016; Bongers et al., 2015). Review framework This systematic synthesis followed the general principles of the Preferred Reporting Items for Systematic Reviews and Meta‑Analyses (Figure 1) extension for ecology and evolutionary biology (PRISMA‑EcoEvo; O’Dea et al., 2021), adapting the checklist to a question‑driven review of coexistence mechanisms and community‑level indicators in dry forest tree communities. The protocol defined a priori the research questions, eligibility criteria, search strategy, screening procedures, and data‑extraction scheme, with a particular emphasis on variables and metrics that can serve as indicators of underlying coexistence processes and ecosystem condition, in line with recent work on forest and ecosystem indicators (Abas et al., 2023; Grima et al., 2023). Information sources The primary bibliographic source was Scopus (Elsevier), selected for its broad coverage of ecological and forestry journals. To reduce database bias, complementary exploratory searches were conducted in Web of Science Core Collection and ScienceDirect for key authors and concepts that appeared under‑represented in the Scopus export (e.g. “tropical dry forest coexistence”, “co‑occurrence patterns dry forest trees”, “multidimensional niches dry forest”). Reference lists of highly relevant articles and recent systematic reviews on forest indicators and biomonitoring were also scanned to identify additional studies not captured in the initial search (Abas et al., 2023; Gatica‑Saavedra et al., 2017; Grima et al., 2023). No grey literature or theses were systematically considered. Search strategy The search strategy was designed to capture studies that: (i) focused on seasonally dry tropical or subtropical forests sensu Miles et al. (2006) and Sánchez‑Azofeifa et al. (2005), and (ii) reported empirical analyses related to coexistence mechanisms, co‑occurrence patterns, or community‑level structure of tree assemblages. Search strings combined four concept blocks applied to title, abstract and keywords in Scopus: 1. Ecosystem block: “tropical dry forest” OR “seasonally dry forest” OR “dry tropical forest” OR “seasonally dry tropical forest” OR “bosque seco” 2. Focal group block: tree* OR “woody species” OR “arboreal” 3. Process/structure block: coexist* OR “co‑occurrence” OR “species co‑occurrence” OR “species association*” OR “community assembly” OR “species assembly” OR “beta diversity” OR “functional diversity” OR “phylogenetic structure” 4. Mechanism/indicator block: “niche differentiation” OR “neutral theory” OR “density dependence” OR “Janzen‑Connell” OR “environmental filtering” OR “functional traits” The final Scopus query, restricted to 2000–2025, was of the general form: (TITLE‑ABS‑KEY(“tropical dry forest” OR “seasonally dry forest” OR “dry tropical forest” OR “bosque seco”) AND TITLE‑ABS‑KEY(tree* OR “woody species” OR arboreal) AND TITLE‑ABS‑KEY(coexist* OR “co‑occurrence” OR “species co‑occurrence” OR “species association*” OR “community assembly” OR “beta diversity” OR “functional diversity” OR “phylogenetic structure”) AND TITLE‑ABS‑KEY(“niche differentiation” OR “neutral theory” OR “density dependence” OR “Janzen‑Connell” OR “environmental filtering” OR “functional traits”)) AND PUBYEAR > 1999 AND PUBYEAR < 2026 No language filter was applied at the search stage, but subsequent screening prioritized articles in English and Spanish. Eligibility criteria Eligibility criteria were defined to retain studies directly informative about mechanisms of coexistence or patterns of co‑occurrence in dry forest tree communities and about community‑level indicators potentially relevant for assessment and monitoring: Inclusion criteria: • Peer‑reviewed empirical studies (observational or experimental) published between 2000 and 2025. • Study system primarily classified as seasonally dry tropical or subtropical forest, including Neotropical, African and Asian dry forests sensu Miles et al. (2006) and Quesada et al. (2009). • Focal community composed mainly of woody plants where trees form the dominant stratum; mixed tree–liana systems were included when tree assemblages were explicitly analysed (e.g. Pulla et al., 2017). • Explicit analysis or discussion of: • coexistence mechanisms (e.g. niche partitioning, environmental filtering, density dependence, storage effects, neutral processes), and/or • co‑occurrence patterns, spatial or temporal associations, beta diversity, or community structure of tree assemblages. • Reporting of at least one community‑level metric or indicator related to taxonomic, functional, phylogenetic or network structure (e.g. species richness and β‑diversity components, functional diversity indices, phylogenetic dispersion, co‑occurrence or network metrics) that could be interpreted in relation to coexistence processes. Exclusion criteria: • Purely conceptual, theoretical or modelling studies without empirical data from dry forests. • Studies on fauna, herbaceous plants or other life forms without a clearly delimited tree component. • Silvicultural trials or plantation studies focusing on single species or mixtures without community‑level analyses. • Broad biogeographic or macroecological studies that included dry forests but did not separate them from other forest types in analyses or reporting. Study selection and screening All records retrieved from Scopus were exported in CSV format, including bibliographic information, abstracts, author keywords and index keywords. Duplicate records across databases were identified using DOI and title and removed. Screening proceeded in two stages following PRISMA‑EcoEvo recommendations (O’Dea et al., 2021). Title and abstract screening: Records were first screened based on title and abstract to exclude clearly irrelevant studies (e.g. non‑forest biomes, non‑woody communities, purely physiological or agronomic trials without community context). At this stage, broad inclusion was favoured for any study that mentioned dry forests, tree communities and at least one of the target concepts (coexistence, co‑occurrence, community assembly, diversity metrics, functional or phylogenetic structure). Full‑text screening: For the remaining records, full texts were assessed against the eligibility criteria. Studies that only mentioned coexistence or co‑occurrence in passing, without any community‑level analysis or indicator that could be related to mechanisms, were excluded. Reasons for exclusion (e.g. biome mismatch, no tree community focus, insufficient data on community metrics) were recorded qualitatively. The numbers of records identified, screened, deemed eligible and included will be reported in a PRISMA flow diagram following O’Dea et al. (2021) and Abas et al. (2023). Data extraction For each included study, data was extracted using a standardized template focused on both ecological mechanisms and indicators: • Bibliographic information: authors, year, journal, region and country, coordinates or broad location, and biome classification. • Study design: forest type (e.g. mature vs secondary, protected vs managed), plot size and number, sampling design (e.g. permanent plots, chronosequences, transects), and temporal extent. • Focal level and life stage: seedlings, saplings, adults, multiple ontogenetic stages. • Mechanisms addressed: environmental filtering, niche partitioning (resource, habitat, temporal), density dependence (e.g. Janzen‑Connell), neutral or dispersal limitation, facilitation, priority effects, or combinations thereof, as explicitly interpreted by the authors. • Community‑level metrics and indicators: • Taxonomic: species richness, evenness, α‑ and β‑diversity (including partitioning into turnover and nestedness), spatial or temporal turnover indices. • Functional: trait distributions (community‑weighted means), multidimensional trait space, functional richness/evenness/divergence, functional β‑diversity. • Phylogenetic: indices of phylogenetic diversity, clustering or overdispersion (e.g. NRI, NTI), phylogenetic β‑diversity and related metrics. • Co‑occurrence and network: pairwise co‑occurrence indices, null‑model outputs, modularity, centrality or indicator‑species values when available (Gotelli & McCabe, 2002; Carrión, 2017; Macías‑Mora et al., 2025). • Main conclusions about coexistence mechanisms and their relationship to the reported metrics or indicators. When quantitative effect sizes or summary statistics were not directly comparable across studies, metrics were retained qualitatively (e.g. “functional divergence increased along succession”, “phylogenetic clustering at local scale with overdispersion at regional scale”) and coded into broad response categories to enable synthesis. Quality assessment Although no formal risk‑of‑bias tool exists for ecological coexistence studies, a simple quality‑assessment scheme was applied to aid interpretation of the evidence base (O’Dea et al., 2021). Each study was scored on three dimensions (0–2 scale each): 1. Design and reporting clarity: Adequacy of site description, forest type classification and sampling design (e.g. clear plot sizes, replication, temporal coverage). 2. Suitability of methods for inferring mechanisms: Appropriateness and transparency of methods used to infer coexistence processes or co‑occurrence structure (e.g. null models specified, assumptions discussed, link between metrics and mechanisms explicitly justified; Gotelli & McCabe, 2002; Wiegand et al., 1999). 3. Relevance and interpretability of indicators: Extent to which reported diversity, trait, phylogenetic or co‑occurrence metrics were interpretable as indicators of underlying mechanisms and/or ecosystem condition (Abas et al., 2023; Grima et al., 2023). These scores were not used to exclude studies but to weigh their contribution in the qualitative synthesis and in highlighting robust versus tentative inferences regarding mechanisms and indicators. Results Study selection and general characteristics The initial Scopus search returned 120 records for the period 2000–2025 after duplicate removal. A keyword‑based screening focused on coexistence, co‑occurrence, community assembly and diversity metrics indicated that 108 records (90%) contained at least one of these mechanism‑ or indicator‑related terms in the title, abstract or keywords, and were therefore retained for full‑text assessment. Following application of the predefined eligibility criteria, the final dataset consisted of studies spanning the entire period, with a marked increase in publications after 2010 and peaks in 2016–2017 and 2021–2025 (Figure 2), reflecting growing interest in dry forest community assembly and associated indicators Temporal coverage of the selected studies was uneven, with only a few contributions published before 2005 and a sustained rise from 2010 onwards, paralleling broader trends in trait‑based ecology and systematic indicator development for forests (Miles et al., 2006; Abas et al., 2023). Many recent papers explicitly framed their analyses in terms of functional or phylogenetic diversity, spatial co‑occurrence or resilience indicators, often linking these metrics to management or restoration questions in seasonally dry forests. Representative examples of the selected literature include work on functionally deterministic but taxonomically stochastic secondary succession in a Mexican dry forest, studies of regenerative niche divergence in co‑invasive woody species, analyses of keystone species and co‑occurrence patterns in Ecuadorian dry forests, and assessments of trait‑based responses to environmental gradients such as topographic water availability or salinity stress in Neotropical dry forests. Geographical coverage was strongly biased towards Neotropical seasonally dry forests, with 72 of the 108 mechanism‑related studies conducted in the Neotropics, 12 in Asian dry forests, 2 in African dry forests, and one explicitly spanning both Africa and the Neotropics, while 33 records did not provide a clear regional tag in the title, abstract or keywords (Table 1). Study characteristics and indicator types The set of mechanism‑related studies encompassed a wide variety of community‑level metrics and indicators, with a clear predominance of functional and co‑occurrence‑based approaches. A simple keyword‑based classification of abstracts and keywords indicated that roughly one third of the records focused primarily on functional indicators (e.g. trait distributions, functional diversity or functional space occupation), another third combined functional metrics with explicit co‑occurrence or association analyses, and smaller subsets emphasized purely co‑occurrence, taxonomic or phylogenetic indicators. Only a very limited number of studies simultaneously integrated taxonomic, functional and phylogenetic dimensions in a unified indicator framework, despite recurrent calls for multidimensional biodiversity metrics in forest assessment (Abas et al., 2023; Grima et al., 2023). Functional indicators were frequently derived from leaf, wood and root traits related to water use, drought tolerance and resource acquisition, and were used to infer environmental filtering, niche differentiation and response diversity along gradients of water availability, disturbance or succession (e.g. Buzzard et al., 2016; Pineda‑García et al., 2011; Bartlett et al., 2012). Co‑occurrence‑based indicators included pairwise co‑occurrence indices, null‑model outputs and, in a few cases, network‑ or indicator‑species analyses to identify keystone or characteristic assemblages in Neotropical dry forests (Gotelli & McCabe, 2002; Carrión, 2017; Macías‑Mora et al., 2025). Taxonomic and phylogenetic metrics (e.g. α‑ and β‑diversity partitions, phylogenetic clustering or overdispersion) were less frequently the focus but often complemented trait‑based or co‑occurrence analyses, providing additional insight into the scale and evolutionary depth of assembly processes in dry forest tree communities (Brown et al., 2013; Pulla et al., 2017). To further characterize how studies combine different dimensions of community structure, we summarized the frequency of single‑ and multi‑dimensional indicator sets (taxonomic, functional, phylogenetic and co‑occurrence) across the 108 mechanism‑related studies. Figure 3 illustrates that most studies rely on functional indicators alone or in combination with co‑occurrence metrics, whereas fully multidimensional approaches that integrate three or four indicator types within a single analytical framework remain comparatively rare. In addition, the overall frequency of each indicator type (Figure 4) shows that approximately 50 studies reported functional indicators (traits and functional diversity), 33 included co‑occurrence or association metrics, and about 15 each used taxonomic or phylogenetic diversity indicators as primary or complementary metrics, highlighting the predominance of functional and co‑occurrence‑based approaches in the dry forest coexistence literature. Mechanisms of coexistence and associated indicators Across the selected literature, four broad classes of mechanisms were most frequently invoked to explain tree species coexistence in seasonally dry forests: (i) environmental filtering and habitat specialization, (ii) functional niche differentiation and trade‑offs, (iii) density‑dependent and other biotic interaction processes, and (iv) dispersal limitation and neutral‑like dynamics. These mechanisms were rarely considered in isolation; instead, most studies interpreted observed community patterns as the joint outcome of multiple processes acting at different spatial and temporal scales, with community‑level indicators reflecting this interplay rather than a single dominant driver (Brown et al., 2013; Pulla et al., 2017). Environmental filtering by water availability, soils and microtopography emerged as a pervasive mechanism, particularly in studies that related tree distributions or community composition to fine‑scale gradients in water status and edaphic conditions (Balvanera et al., 2002; Quesada et al., 2009). Indicators of this process included strong associations between topographic position and functional trait values (e.g. turgor loss point, leaf and root traits), reduced functional space in the driest microsites, and consistent shifts in community‑weighted means along moisture gradients (Pineda‑García et al., 2011; Bartlett et al., 2012; Cely et al., 2025). Such patterns were typically interpreted as evidence that only species or individuals with trait combinations conferring sufficient drought tolerance can persist in the most stressful habitats, generating non‑random functional and, in some cases, phylogenetic clustering at local scales (Markesteijn & Poorter, 2009; Arango et al., 2021). Functional niche differentiation and life‑history trade‑offs were commonly inferred from trait‑based indicators and successional trajectories. Studies of secondary succession in dry forests reported that while taxonomic composition can be highly stochastic, functional trait composition tends to follow more deterministic trajectories, with convergence on particulars combinations of traits associated with water use, wood density and regeneration strategies (Buzzard et al., 2016; Werden et al., 2020). Indicators of these mechanisms included increases or directional changes in functional richness and divergence along successional gradients, shifts in the occupation of multidimensional trait space, and evidence of regenerative niche divergence among co‑occurring or coinvasive woody species (Simian et al., 2025). Together, these results suggest that multiple, partially overlapping trait syndromes can facilitate coexistence by reducing niche overlaps in key resource or regeneration axes, even when species share similar life forms and habitat ranges. Density‑dependent processes and other biotic interactions were less frequently quantified directly but often invoked to explain residual variation in recruitment, survival and spatial structure after accounting for environmental filtering. Studies examining recruitment limitation and conspecific neighbourhood effects in dry forests reported strong dispersal and establishment limitations that varied among species and ontogenetic stages, with some evidence that local conspecific density influences seedling performance and community functional composition (Gee et al., 2025; Pulla et al., 2017). Co‑occurrence indicators, such as negative associations between particular species pairs or reduced conspecific aggregation relative to null expectations, were interpreted as indirect signals of negative density dependence, competition or Janzen–Connell–type effects, whereas positive associations and co‑occurrence of structurally or functionally complementary species were taken as possible evidence of facilitation, especially under high abiotic stress (Carrión, 2017; Macías‑Mora et al., 2025). Finally, dispersal limitation and neutral‑like dynamics were highlighted in several studies that documented high stochasticity in species composition, idiosyncratic species responses to climate variability and weak relationships between some diversity metrics and measured environmental gradients (Clark, 2010; Brown et al., 2013). Indicators of these processes included: (i) large unexplained variation in species composition among nearby sites with similar abiotic conditions, (ii) strong spatial autocorrelation in species occurrences not fully accounted for by environment, and (iii) successional trajectories where species turnover was only weakly linked to functional shifts (Derroire et al., 2016; Werden et al., 2020). Several authors argued that neutral or nearly neutral processes may be particularly important in early successional stages or in highly fragmented landscapes, where dispersal constraints and historical contingencies can override fine‑scale environmental filtering, at least transiently. At a synthetic level, the relationships between mechanisms, indicators and applications in seasonally dry forests can be summarized in a simple conceptual framework. In this framework, environmental filtering and habitat specialization are primarily reflected in shifts in community‑weighted trait means, functional diversity and, in some cases, phylogenetic clustering, whereas functional niche differentiation is captured by patterns of functional divergence and β‑diversity along environmental and successional gradients. Density‑dependent processes and other biotic interactions are most consistently expressed through non‑random co‑occurrence and network metrics, while neutral‑like dynamics are associated with high taxonomic and functional turnover that is only weakly explained by measured environmental gradients. These linkages between coexistence mechanisms, community‑level indicators and management applications are summarized in Figure 5, which provides a conceptual guide for selecting indicator sets to diagnose underlying processes and to assess ecosystem condition, resilience and restoration trajectories in seasonally dry forests. Discussion Indicators of coexistence mechanisms in dry forests The synthesis of empirical studies on dry forest tree communities reveals that different classes of community‑level indicators vary markedly in their diagnostic power for underlying coexistence mechanisms. Functional trait‑based indicators emerge as the most consistently informative for environmental filtering and functional niche differentiation, particularly when traits are mechanistically linked to water use, drought tolerance and regeneration strategies (Markesteijn & Poorter, 2009; Pineda‑García et al., 2011; Buzzard et al., 2016). Community‑weighted means and functional diversity indices (e.g. functional richness, evenness, divergence) responded predictably to gradients of water availability, successional stage and disturbance, supporting their use as indicators of filtering strength and the degree of trait convergence or divergence in dry forest tree communities (Bartlett et al., 2012; Lohbeck et al., 2014). However, the interpretation of these metrics still depends on careful trait selection, scale, and the consideration of intraspecific variability, which several recent studies show can be substantial along fine‑scale topographic gradients (Cely et al., 2025; Pulla et al., 2017). Co‑occurrence‑based indicators provide complementary, but more equivocal, information about biotic interactions and density‑dependent processes. Null‑model analyses of species associations have detected both aggregated and segregated patterns in dry forest tree assemblages, yet similar co‑occurrence structures can arise from environmental filtering, dispersal limitation or species interactions, making them inherently multi‑causal (Gotelli & McCabe, 2002; Wiegand et al., 1999). In the context of dry forests, where strong abiotic gradients and habitat heterogeneity are pervasive, co‑occurrence metrics appear most interpretable when combined with explicit environmental and trait data, as in studies that link negative conspecific aggregation to density‑dependent mortality or that identify keystone species whose presence structures local assemblages (Carrión, 2017; Macías‑Mora et al., 2025). From an indicator perspective, this suggests that co‑occurrence metrics alone are insufficient as mechanistic indicators but can serve as useful red flags of non‑random spatial structure when cross‑validated against functional and environmental information. Taxonomic and phylogenetic indicators, widely used in broader forest health and biodiversity assessments, play a more supportive role for mechanistic inference in dry forests. Species richness and β‑diversity partitions capture important information on turnover and nestedness along environmental and successional gradients, but they are largely agnostic about process and can be reproduced under multiple assembly scenarios (Miles et al., 2006; Brown et al., 2013). Phylogenetic structure indices (e.g. clustering or overdispersion) have been used to infer the balance between environmental filtering and competitive exclusion, yet their interpretation is hampered by incomplete phylogenies, trait conservatism assumptions and scale dependence, particularly in species‑rich tropical systems (Arango et al., 2021; Pulla et al., 2017). The strongest inferences arise in studies that triangulate taxonomic, functional and phylogenetic indicators, for instance when local phylogenetic clustering coincides with functional convergence under strong drought stress, or when high functional turnover is decoupled from relatively modest taxonomic turnover along successional or hydrological gradients (Buzzard et al., 2016; Derroire et al., 2016). Taken together, these findings point to the value of multidimensional indicator sets that combine taxonomic, functional, phylogenetic and co‑occurrence metrics to disentangle coexistence pathways in dry forest tree communities. This aligns with broader reviews in Ecological Indicators emphasizing the need for composite or integrative indicators to capture forest condition and resilience (Abas et al., 2023; Grima et al., 2023). For monitoring and management, functional trait‑based indicators linked to water relations and regeneration, coupled with carefully interpreted co‑occurrence metrics and β‑diversity components, appear particularly promising for diagnosing the relative importance of environmental filtering, niche differentiation and neutral‑like dynamics in seasonally dry forests. At the same time, the synthesis highlights substantial gaps, including limited coverage of African and Asian dry forests, under‑representation of long‑term and multi‑life‑stage studies, and uneven reporting of the methodological details needed to evaluate indicator robustness—issues that should be addressed in future research and in the development of standardized indicator frameworks for dry forest ecosystems. Limitations This synthesis has several limitations that should be considered when interpreting its findings. First, the evidence base is derived primarily from studies indexed in Scopus, which may under‑represent literature published in regional journals or in languages other than English and Spanish, particularly for African and Asian dry forests. Second, the classification of mechanisms and indicator types relied on information reported in titles, abstracts, keywords and methods, combined with keyword‑based text screening, which can misclassify studies that use indicators without naming them explicitly or that discuss mechanisms only in the discussion. Third, we did not perform a formal meta‑analysis of effect sizes, and many of the reported metrics are not directly comparable across studies due to differences in design, scale, trait selection and analytical choices, which constrains the strength of quantitative inferences about the relative importance of specific coexistence mechanisms and indicators. Finally, because most of the included studies are observational and cross‑sectional, causal interpretations remain tentative, and experimental and long‑term data are still needed to validate the mechanistic links between indicators, coexistence processes and ecosystem condition in seasonally dry forests. Implications and future directions The synthesis underscores that indicators of coexistence mechanisms in seasonally dry forests must explicitly acknowledge scale, context and multidimensionality. Functional trait‑based metrics derived from hydraulics, leaf and root traits are particularly powerful at local to landscape scales for diagnosing environmental filtering and niche differentiation, but their interpretation depends on integrating intraspecific variation, life‑stage differences and disturbance history (Pineda‑García et al., 2011; Pulla et al., 2017). Co‑occurrence and network‑based indicators, while attractive for their apparent simplicity, should be applied in conjunction with environmental and trait information and with transparent null‑model specifications if they are to inform management decisions about keystone species, facilitative interactions or density‑dependent processes (Gotelli & McCabe, 2002; Macías‑Mora et al., 2025). Several of the indicator classes highlighted in this review can be directly mapped onto emerging forest and ecosystem condition frameworks. Functional trait spectrum and functional β‑diversity along hydrological and successional gradients, for example, correspond closely to indicators of ecosystem structure and function that have been proposed for forest health assessment and ecosystem condition reporting (Abas et al., 2023; Grima et al., 2023). Likewise, taxonomic and phylogenetic β‑diversity, together with co‑occurrence‑based identification of keystone or indicator species, align with biodiversity and compositional indicators used in international initiatives on ecosystem condition and nature’s contributions to people. Embedding these coexistence‑oriented metrics within standardized indicator frameworks would facilitate the inclusion of seasonally dry forests in national and global monitoring schemes, and help ensure that coexistence processes and resilience are explicitly considered in assessments of forest condition and restoration success. From a monitoring and policy perspective, the results support ongoing efforts in Ecological Indicators to move towards standardized, composite indicators of forest condition and resilience that combine taxonomic, functional and structural information (Abas et al., 2023; Grima et al., 2023). For seasonally dry forests, promising candidate indicator sets include: (i) β‑diversity components capturing spatial and temporal turnover, (ii) functional trait spectra and functional β‑diversity along hydrological and successional gradients, and (iii) co‑occurrence‑derived metrics identifying keystone or indicator species that structure assemblages and may signal shifts in coexistence regimes. Future research should prioritize under‑represented regions (especially African and Asian dry forests), long‑term and multi‑life‑stage studies, and explicit testing of hypothesized links between specific indicators and coexistence mechanisms under ongoing climate change and land‑use intensification. Conclusions This systematic synthesis shows that tree species coexistence and co‑occurrence in seasonally dry tropical and subtropical forests are governed by a complex interplay of environmental filtering, functional niche differentiation, density‑dependent interactions and neutral‑like dynamics, with the relative importance of each mechanism varying across spatial scales, ontogenetic stages and disturbance contexts. Among the diverse community‑level metrics used in the literature, functional trait‑based indicators and multidimensional diversity measures (taxonomic, functional, phylogenetic) provide the most consistent and interpretable signals of underlying coexistence processes, especially when combined with carefully designed co‑occurrence analyses and robust null models. Advancing towards standardized, indicator‑oriented frameworks for dry forests will require integrating these metrics into practical monitoring schemes, improving geographic and temporal coverage, and explicitly evaluating how well proposed indicators track changes in coexistence pathways and ecosystem condition in the face of accelerating global change. The authors express their gratitude to the Universidad Técnica de Manabí for the institutional support provided. We also thank the Research Group FAGROCLIM and the Latin American Seasonally Dry Tropical Forest Floristic Network (DRYFLOR) for their valuable contributions and support to this research. Author contributions CASM: Conceptualization, Supervision, Writing – review & editing. KME: Formal analysis, Writing – original draft (Results). JRMB: Writing – original draft, Investigation. MBVO: Formal analysis, Writing – original draft. LDBS: Writing – original draft, Investigation, Writing – review & editing. 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Keywords 10: forest 27: trees 36: biodiversity 5: species interaction community assembly forest functional traits and indicators multidimensional biodiversity metrics species co‑occurrence patterns Authors Affiliations Carlos Salas-Macías Universidad Técnica de Manabí Facultad de Ingeniería Agronómica View all articles by this author Karime Montes-Escobar Universidad Técnica de Manabí View all articles by this author José Macías-Barberán Universidad Técnica de Manabí View all articles by this author María Vinces Obando 0000-0002-3310-6202 [email protected] Universidad Técnica de Manabí Facultad de Ingeniería Agronómica View all articles by this author Luis Balarezo-Saltos Universidad Técnica de Manabí View all articles by this author Carolina Fonseca-Restrepo Universidad Técnica de Manabí View all articles by this author Metrics & Citations Metrics Article Usage 126 views 46 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Carlos Salas-Macías, Karime Montes-Escobar, José Macías-Barberán, et al. 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