1 Introduction

Whether plants can continue and spread in different environments often depends on the effect of seed propagation. It not only affects the survival of individual species, but also influences community structure and the diversity and resilience of ecosystems (McConkey et al., 2024; Pereira et al., 2025). After seeds leave their mother plants, interspecific competition will be alleviated, and density-dependent mortality rates will also decline, making them more adaptable to the constantly changing environment. It can thus be seen that the dissemination process itself has become an important force driving ecological functions and evolution (Rogers et al., 2021; Pereira et al., 2025).

As for the mode of dissemination, there is no single path. Wind and water can carry away seeds, while the role of animals is particularly prominent. They often complete the long-distance transportation of plants in casual feeding or activities (Nathan et al., 2008; Harrer and Levi, 2018; Aguado et al., 2022). Many terrestrial plant populations are in fact highly dependent on this animal-mediated transmission (Reiserer et al., 2018; McConkey et al., 2024). These interactions not only sustain vegetation renewal and forest regeneration, but are also closely related to biodiversity conservation and the long-term interests of human society (Banos-Villalba et al., 2017; Godo et al., 2023). However, such a network is not stable and secure. With the fragmentation of habitats, the change of animal behavioral patterns and the continuous fluctuations of climate, the transmission process has gradually been disturbed, and the stability of the ecosystem has thus endured greater uncertainty (McConkey et al., 2012; Spengler, 2020; Rogers et al., 2021).

This study aims to analyze the existing knowledge on seed dispersal mechanisms and patterns in terrestrial ecosystems, with a focus on exploring the diversity of dispersal media, the ecological and evolutionary consequences of dispersal. By integrating the results of cross group, cross landscape, and different research methods, we strive to clarify the roles of well studied and overlooked transmission factors, highlighting the interaction between transmission processes and ecosystem dynamics. This study aims to provide a comprehensive framework for understanding how seed dispersal shapes terrestrial ecosystems and to offer reference strategies for maintaining plant diversity and ecosystem functionality in a rapidly changing world.

2 Theoretical Background of Seed Propagation

2.1 The evolutionary significance of communication

Seed propagation is a key evolutionary strategy that enhances plant fitness by reducing competition among closely related individuals, avoiding density-dependent death near the mother plant, and promoting colonization in new or disturbed habitats. Propagation enables plants to track changes in environmental conditions, maintain gene flow, and buffer populations from local extinction, thus playing a core role in the persistence and adaptability of plant species (Beckman et al., 2019; Chen et al., 2020). The trade-off between transmission and dormancy is also regarded as an important risk diversification strategy. Different species often choose spatial transmission or temporal transmission (dormancy) based on environmental variability and life history characteristics (Chen et al., 2020).

2.2 Classical communication model

In the study of seed propagation, theoretical models have long been important tools for understanding this process. Researchers rely on both empirical analysis and deduction, and the "propagation core" is the most common approach. It describes the distance of seed diffusion from the mother plant through probability distribution (Bullock et al., 2017; Kim et al., 2022). This distribution has no uniform form. It may be exponential, power-law or logarithmic hyperbolic. Different forms often reflect the differences in propagation mechanisms and environmental backgrounds (Bullock et al., 2017; Kim et al., 2022). Unlike this, the mechanism model pays more attention to the propagation process itself. For example, in the study of wind propagation, parameters such as seed morphology, release height, wind speed and turbulence often need to be considered to predict the distance and pattern of propagation (Nathan et al., 2011; Kim et al., 2022).

2.3 Main ecological consequences

People tend to view seed dispersal as a matter of individual continuity, but in reality, it has already left traces on a larger scale - genetic diversity, population expansion, and even the form of community structure are all closely related to it. The flow of genes between different populations essentially relies on this transmission process. As a result, the risk of inbreeding is reduced, variations are preserved, and species have more opportunities to adapt to long-term environmental changes (Beckman et al., 2019). However, transmission is not only linked to genetic communication, it also determines the speed and mode of population movement in space. Sometimes it is opening up new territories, sometimes it is forced migration, and success or failure often depends on the efficiency of dissemination (Nathan et al., 2008; Beckman et al., 2019).

Looking at it at the scale of a community, things are even more complex. If the transmission mechanism is properly utilized, species can coexist better and the renewal process can be driven, thereby affecting the composition and diversity of the community (Schupp et al., 2010). But this influence is not always the same, it will be constrained by various specific conditions. The number and effectiveness of communication events, the ecological habits of the communicators themselves, and even the differences in the surrounding landscape patterns can all lead to significant differences in the results (Schupp et al., 2010; Nevo et al., 2023).

3 The Mechanism of Seed Propagation

3.1 Non-biological transmission

Abiotic seed dispersal refers to the transportation of seeds by non-living environmental forces, thereby shaping the distribution and community structure of plants in terrestrial ecosystems. The main abiotic transmission methods include wind transmission (anemochory), water transmission (hydrochory), and gravity transmission (barochory), each accompanied by unique seed adaptability and ecological background.

Wind propagation is a widespread mechanism, especially common in open areas and plants with adaptive seeds. Seeds propagated by wind usually have special morphological characteristics, such as wings, crown hairs or down. These structures can increase the surface area and reduce the terminal velocity, allowing the seeds to stay in the air for a longer time and spread farther (Nathan et al., 2002; Soons et al., 2017; Snell et al., 2019). For instance, seeds with samaras and feather-like crown hairs are common adaptive characteristics that can be lifted and horizontally moved by turbulent updraft (Nathan et al., 2002; Soons et al., 2017; Snell et al., 2019). Mechanism models indicate that successful long-distance wind propagation depends on seed release time, canopy structure and meteorological conditions. Among them, updraft and turbulence play a key role in lifting seeds above the canopy for long-distance transportation (Nathan et al., 2002; Snell et al., 2019; Kim et al., 2022). These adaptive characteristics enable plants to colonalize in new or disturbed habitats and play a core role in species expansion and ecosystem resilience (Nathan et al., 2002; Nathan et al., 2008; Kim et al., 2022).

Water dispersal is very common in riverbank, wetland and coastal ecosystems, and its seeds are transported by streams, rivers or ocean currents. Seeds adapted to water propagation usually have floating structures, waterproof shells or inflatable structures, which enable the seeds to float and remain active during prolonged soaking (Nathan et al., 2008; Soons et al., 2017). This mechanism enables plants to take advantage of the dynamic aquatic environment, promoting gene flow and connectivity among populations, and even achieving diffusion in fragmented landscapes. Ocean currents can also drive long-distance transmission and contribute to the colonization of remote islands and coastal habitats (Nathan et al., 2008). In areas prone to flooding or seasonal flooding, water dispersal is particularly important because water body movement is the dominant ecological force (Nathan et al., 2008; Soons et al., 2017).

Gravity propagation, also known as gravity-driven propagation, refers to the process where seeds fall from their mother plants by their own weight and roll to nearby areas. This mechanism is common in plants with large and heavy seeds or fruits. Terrain features such as slopes or uneven ground can further promote the movement of seeds (Soons et al., 2017). Although gravity propagation usually causes a limited propagation distance compared with wind propagation or water propagation, it still has important ecological significance in dense forests or steep terrains because seeds may aggregate in suitable microhabitats, which is conducive to germination and establishment (Soons et al., 2017). The spatial patterns produced by gravitational propagation often affect the local genetic structure and seedling renewal dynamics (Horn et al., 2001).

3.2 Biological transmission

Biology helps plants spread seeds, which is not accidental, but the result of long-term interaction. It not only affects how populations expand in space, but is also closely related to whether genetic diversity can be maintained and whether ecosystems can be restored. The ways of transmission are not singular, they may involve animals or humans, and each method will leave its own distinct imprint at the ecological level.

Take Epizoochory as an example, which is quite intuitive. The seeds do not enter the animal's digestive system, but choose another path - hanging tightly on the hair, feathers, or skin. Some plants have even evolved special structures for this, such as hooks or sticky shells, which make it easier to "hitchhike" and even achieve long-distance diffusion (Zhang and Wang, 2023). This approach is particularly prominent in fragmented or highly differentiated environments, as animal movements often cross isolation zones, connecting dispersed plants and promoting gene flow and population exchange (Cruzan and Hendrickson, 2020).

Endozoochory refers to the internal transmission achieved by animals through feeding, especially in fruit-eating animals. The seeds are ingested along with the fleshy fruits and excreted with feces at a distance from the mother plant. After passing through the digestive tract, the germination rate of seeds is often increased due to the erosion of the seed coat or the removal of the inhibitory pulp. For example, in the spread among primates, the germination speed and success rate of seeds of many tropical plants have significantly increased (Fuzessy et al., 2016; Nevo et al., 2023). Large vertebrates, including mammals and birds, are important vectors of long-distance transmission and play a crucial role in forest regeneration, population connectivity and the maintenance of plant diversity (Nathan et al., 2008; Fuzessy et al., 2016; Tol et al., 2017). However, the loss of these spreaders due to deanimalization or habitat change may disrupt plant regeneration and alter community structure (Beckman and Rogers, 2013; Donoso et al., 2022).

Among the various ways of seed dispersal, Myrmecocorry is often mentioned. The reason is not difficult to understand: many plant seeds have elaiosomes on their surface, which are small structures rich in nutrients that can attract ants. Ants carry seeds back to their nest, gnaw off oily bodies, and usually throw the seeds around the nest. The result is that the seeds both evade predators and fall into a fertile, relatively safe microenvironment, greatly increasing the chances of seedling survival (Zhang and Wang, 2023). This "mutually beneficial arrangement" can be seen in different ecosystems, often quietly changing the distribution pattern of plants and long-term affecting the structure and diversity of communities (Zhang and Wang, 2023).

In contrast, the story of Anthropochory is not so 'natural'. It is not a product of mutualistic coexistence, but more like a trace left by human activities. Farming, gardening, and even transcontinental trade may inadvertently bring seeds to faraway places (McConkey et al., 2012; Soons et al., 2017). Once brought into a new area, some plants will rapidly spread and even become invasive species. Due to its fast speed and wide range, human mediated transmission often causes significant changes in plant communities in a short period of time, which impacts local ecosystems and presents more challenging management and protection (McConkey et al., 2012; Soons et al., 2017).

Biological transmission mechanisms, with their diversity and efficiency, play a crucial role in plant supplementation, genetic communication and ecosystem resilience. Especially in the current situation where habitat fragmentation and environmental change are constantly intensifying, the importance of these mechanisms is even more prominent (McConkey et al., 2012; Beckman and Rogers, 2013; Fuzessy et al., 2016; Zhang and Wang, 2023).

4 Patterns of Seed Propagation in Ecosystems

4.1 Spatial pattern

Seed propagation can form various spatial patterns ranging from highly concentrated to relatively uniform distribution, which depends on the propagation mechanism and ecological environment background. The distance of propagation varies greatly. Most seeds deposit near the mother plant, but there are also some seeds that can spread over long distances. Hybrid models (such as 2Dt kernel functions) can better describe both local propagation and long-distance propagation events simultaneously, and are more in line with reality compared with traditional models. The mode of transmission has an important influence on spatial aggregation. For example, seeds transmitted by animals and gravity tend to be distributed in clusters, while seeds transmitted by wind are more likely to spread widely and evenly (Seidler and Plotkin, 2006). These spatial patterns are crucial for community structure. They affect species coexistence and the spatial arrangement of plant populations (Seidler and Plotkin, 2006).

4.2 Time mode

The temporal pattern of seed propagation is jointly influenced by plant phenology and the activity cycle of the propagation medium. Many plant species exhibit seasonal transmission events, which often coincide with peak animal activity or optimal environmental conditions for seedling establishment (Cruz et al., 2013; Valdesolo et al., 2022). For example, in coastal dune communities, different plant groups spread their propagation time in spring, summer and late summer, thereby reducing direct competition and possibly promoting community formation (Valdesolo et al., 2022). In the animal-mediated propagation system, the time of fruit ripening and the feeding behavior of animals form the impulse effect of seed propagation, and fruit-eating animals and other propagators drive the temporal changes of seed rain (Cruz et al., 2013; Morales et al., 2013).

4.3 The influence of landscape scale

The landscape structure has a profound influence on the seed propagation pattern. Habitat fragmentation typically reduces the number of plant and animal species, the diversity of transmission interactions, and the number of interspecies connections, thereby leading to functional homogeneity and the loss of specific transmission relationships (Emer et al., 2019). In fragmented landscapes, the propagation network often transforms into a more universal form, that is, universal species are maintained while specific species gradually decline (Emer et al., 2019). Ecological corridors and residual patches can promote the movement of disseminators and seeds and alleviate the negative effects brought by fragmentation to a certain extent (Garcia et al., 2010; Morales et al., 2013). Edge effect and landscape heterogeneity can also affect the location of seed deposition, as the activities and behaviors of animals respond to vegetation structure, patch quality and resource availability (Garcia et al., 2010; Morales et al., 2013; Russo et al., 2024). In urban and human-transformed landscapes, land use history and habitat connectivity further shape the functional composition and spatial distribution of plant communities by influencing seed propagation (Johnson et al., 2018).

5 The Ecological and Evolutionary Significance of Seed Propagation

5.1 Role in species coexistence and biodiversity maintenance

Seed dispersal plays an important role in maintaining species coexistence and diversity in terrestrial ecosystems. Without it, many seeds would die near the mother plant due to density dependence; With it, they can enter new or disturbed environments, reduce competitive pressure, and maintain diversity in their colonies (Beckman and Sullivan, 2023). The form of communication networks is not accidental, but is shaped by the ecological background and evolutionary history. The interaction patterns of different species, the modularity and specificity of the network, all affect the resilience and diversity maintenance of the community (Schleuning et al., 2014). But once the spread is interrupted, such as the disappearance of key animal spreaders or the intervention of invasive species, plant diversity often decreases and community structure changes accordingly (McConkey et al., 2012; Donoso et al., 2022).

5.2 Impact on forest renewal and succession

Seed propagation is the core driving force for forest regeneration and ecological succession. Effective propagation can promote the recolonization of disturbed areas, the diffusion of pioneer species and the establishment of late succession species, thereby shaping succession paths and forest structures (Andresen et al., 2018). Animal-mediated transmission, especially the role of primates and birds, has been proven to promote the restoration of degraded habitats and maintain tree species diversity in tropical forests (Andresen et al., 2018). The plasticity of propagation strategies, such as the increase in the propagation rate in the early succession stage, can accelerate species substitution and enhance the resilience of the ecosystem after disturbance.

5.3 The impact of transmission restrictions on cenetic structure and adaptation

Restricted transmission, meaning that seeds cannot enter suitable habitats or distant populations, can have a profound impact on genetic structure and adaptive potential. Insufficient dissemination often leads to higher inter population differentiation, weakened gene flow, hindered local adaptation, and even increased extinction risk, especially in fragmented landscapes (Snell et al., 2019; Grasty et al., 2020; Beckman and Sullivan, 2023). At the same time, intraspecific transmission traits are not consistent, and different differences can affect the fitness, population dynamics, and evolutionary direction of plants, as well as their response to environmental changes (Snell et al., 2019). Previous studies have shown that habitat fragmentation can drive rapid evolution of communication strategies, which precisely reflects the dynamic game between ecological conditions and evolutionary processes (Cheptou et al., 2008).

6 Case Study: Seed Propagation in Tropical Rainforests

6.1 The role of neotropical forests and large mammals

In the neotropical rainforest, tapirs, primates and rodents such as the brown-haired rat are key seed disseminators, especially for large-seeded tree species. These animals can promote both primary and secondary transmission: primates and birds often transport seeds over long distances after eating fruits, while terrestrial mammals such as the brown-haired rat bury seeds through storage behavior for future use, thus enabling successful germination and establishment of seeds far from the mother plant (Seidler and Plotkin, 2006). The diversity and abundance of disseminators support the high plant diversity and complex community structure unique to tropical rainforests (Figure 1) (Seidler and Plotkin, 2006; Kakishima et al., 2015).

Figure 1 Schematic diagram of the population and evolution models (Adopted from Kakishima et al., 2015) Image caption: (a) Schematic diagram with two tree species (P1 and P2 on plant layer L0) and the corresponding seed dispersers (birds A1 on layer L1 and squirrels A2 on L2). In each animal layer, the grey cells (U1 and U2) indicate unvisited sites (where animals never visit). (b) An animal (bird) eats a fruit (apple) at the tree (P1) and drops a seed at an unoccupied site (O1) that results in a new apple tree. (c) The fragmentation process repeats at every 6000 MCS, such that (stage I: a single habitat) →3000 MCS→ (stage II: habitat separation and stage III: species differentiation) →3000 MCS→ (stage IV: habitat reunification and resulting species coexistence=stage I) (Adopted from Kakishima et al., 2015)

6.2 Loss of communicators and "hollow forest syndrome"

Hunting and deanimalization have led to a sharp decline in the number of large vertebrates, resulting in what is known as the "hollow forest syndrome". In Guam and the overhunted Amazon forest, the loss of key disseminators has led to a sharp decline in seedling species richness and increased the spatial aggregation of seedlings, eventually forming plant communities with reduced diversity and greater homogeneity (Wandrag et al., 2017; Culot et al., 2017). Although some small fruit-eating animals can to some extent make up for the loss of large propagators, their effect is limited, especially for large seed species that rely on large animals for propagation (Babweteera and Brown, 2009; Culot et al., 2017). Due to the irreplaceability of the transmission network, the extinction of large mammals cannot be completely replaced by small species (Culot et al., 2017).

6.3 Impacts on tree renewal, carbon storage and forest resilience

The disruption of the seed propagation process has a chain effect on forest regeneration, carbon storage and long-term resilience. The reduction in dissemination will lead to a decrease in the recruitment rate of large-seed hardwood tree species, which are often key contributors to forest carbon storage (Culot et al., 2017). Empirical studies have shown that deanimalization may reduce plant renewal of certain species by up to 95%, and changes in tree species community composition may endanger the maintenance of carbon storage in tropical forests (Culot et al., 2017). Furthermore, the loss of disseminators weakens the ability of forests to recover from disturbances, thereby threatening biodiversity and ecosystem services (Wandrag et al., 2017; Culot et al., 2017).

7 The Influence of Human Activities on Seed Propagation

7.1 Habitat destruction and fragmentation

When habitats are reduced and divided, the number and diversity of seed spreaders are often the first to be affected. Large fruit-eating animals and some highly dependent spreaders are most likely to disappear, which directly leads to shortened transmission distance, increased genetic similarity among communities, and decreased effective population size (Fonturbel et al., 2015; Perez-Mendez et al., 2016; Borah and Beckman, 2024). Meanwhile, the original activity patterns of animals will also be disrupted, seeds will be difficult to deposit in suitable habitats, and the coverage of seed rain will subsequently shrink (Borah and Beckman, 2024; Fonturbel et al., 2015). Due to the lack of sufficient redundancy in the dissemination function, once key species disappear, it is often difficult to be replaced by other species. As a result, there are chain changes in both plant renewal and community structure (Leal et al., 2013; Perez-Mendez et al., 2016; Da Silva et al., 2024).

7.2 Climate change

The impact of climate change is more reflected in temporal and spatial patterns. With the changes in temperature and precipitation patterns, the original synchronicity between the fruiting period of plants and animal activities was disrupted, and thus the success rate of seed propagation decreased (Teixido et al., 2022; Hernandez et al., 2023). Frequent droughts and extreme weather further reduce the number and activity range of disseminators. For example, ants significantly decrease under such conditions, and the speed and distance of seed dissemination also decrease accordingly, especially in areas where the ecosystem is already fragile (Oliveira et al., 2019; Hernandez et al., 2023). These changes not only threaten the ability of plants to migrate along with climate niches, but also weaken genetic connectivity (Hernandez et al., 2023).

7.3 Invasive species

Once invasive species enter, the original seed propagation network is often disrupted. Invasive plants not only compete with native plants but may also directly vie for resources of disseminators. Invasive animals, on the other hand, can alter the efficiency and composition of spreaders (Teixido et al., 2022; Da Silva et al., 2024). This influence is even more pronounced in an environment where human intervention is intense. Whether in urban areas or large-scale plantations, communication networks tend to be single, randomness increases, and the diversity and number of native communicators decline significantly (Da Silva et al., 2024). Furthermore, humans themselves often unintentionally promote the spread. Road and vehicle transportation became the medium of diffusion, enabling invasive plants and even genetically modified crops to expand rapidly in a short period of time (Garnier et al., 2008; Beckman et al., 2019; Johnson et al., 2020).

8 Methodological Approaches to Seed Propagation Research

8.1 Field observation and seed tracking methods

Direct field observations, including seed collectors, population censuses, and tracking of labeled seeds, remain fundamental methods for quantifying propagation distance and sedimentary patterns. These methods are crucial for capturing the propagation process at the local scale and verifying the model, but they are often limited by logistical challenges and operational difficulties when tracking long-distance propagation or propagation in complex environments (Schupp et al., 2010; Beckman et al., 2019; Kim et al., 2022). Experimental studies, such as feeding experiments in animal-mediated transmission (e.g., fish transmission), can quantify different stages of transmission and reveal the influence of plant and vector traits (Pollux, 2011). Combining phytocentric and zoocentric sampling methods contributes to a more comprehensive understanding of propagation networks and their interactions (Lussier et al., 2024).

8.2 Genetic markers and parent-child analysis

Genetic techniques, such as molecular markers and parent-child analysis, have revolutionized researchers' ability to correlate scattered seeds or seedlings with their mother plants, making accurate measurements of long-distance transmission possible, which were often difficult to achieve in traditional methods (Cain et al., 2000; Wang and Smith, 2002; Jones and Muller-Landau, 2008). These methods can be combined with classical methods to improve the propagation distribution estimation at the population level, but sampling bias and interference from external germline sources need to be fully considered (Cain et al., 2000; Jones and Muller-Landau, 2008). The advancement of genetic attribution and likelihood analysis methods has continuously expanded the scope and resolution of seed propagation research (Cain et al., 2000; Wang and Smith, 2002).

8.3 Propagation core modeling and landscape connectivity analysis

Mathematical and simulation models, including mechanism models and statistical propagation kernels, have been widely used to describe and predict seed propagation patterns across landscapes (Russo et al., 2006; Cousens et al., 2010; Bullock et al., 2017; Kim et al., 2022). Propagation nuclei, as the probability distribution of seed movement, are usually fitted based on empirical data for generalization among different plant types and propagation patterns (Bullock et al., 2017). The mechanism model gradually incorporated animal behavior, landscape heterogeneity and environmental variables, thereby improving the prediction of "seed shadow" and connectivity (Russo et al., 2006; Cousens et al., 2010). However, the accuracy of the model depends on the quality of empirical data and the integration of related biological processes (Beckman et al., 2019; Kim et al., 2022).

9 The Future Prospects and Research Directions of Seed Propagation

9.1 The integration of seed propagation ecology and conservation biology

The value of seed propagation research will be greatly discounted if it is disconnected from conservation practices. The problems of biodiversity loss, habitat fragmentation and degradation of ecosystem functions all force communication research to respond more directly to conservation needs (McConkey et al., 2012). When formulating strategies, merely maintaining the minimum viable population is often insufficient. What is more crucial is to preserve those communicators who have unique functional roles and ensure that they continue to fulfill their ecological roles. Such a goal requires research to be conducted on a larger spatial scale and in a complex landscape context, considering how multiple driving factors have a superimposed impact on the communication network at the community level. At the same time, taking plant functional groups as the analytical unit not only helps to generalize conclusions across species and guide practice, but also enhances reliability under the verification of species-specific research.

9.2 Prediction of ecosystem resilience under global change

In the context of global change, the importance of seed dissemination is becoming increasingly crucial. It is related to whether the plant population can continue, how the community can be rebuilt, and whether the ecosystem can recover after disturbance. However, the dissemination process is not simple and is often limited by specific ecological contexts, which also makes the prediction results uncertain (Beckman et al., 2019; Snell et al., 2019; Beckman et al., 2020). To enhance predictive ability, future research requires more mechanistic models that closely integrate propagation with population dynamics and environmental variability, in order to more accurately assess diffusion rates, distribution range shifts, and ecosystem responses to climate and human disturbances. A often overlooked detail is the significant differences in intraspecific transmission traits, which have profound impacts on plant fitness and community dynamics. If it can be included in the analysis, the prediction accuracy will be significantly improved under global change scenarios (Snell et al., 2019).

9.3 Emerging tools for large-scale propagation research

In recent years, large-scale research has gradually opened up new prospects for seed dispersal ecology. With the help of remote sensing, environmental sensor networks, and long-term global collaborative projects such as the Long Term Ecological Research Station and ForestGEO, cross community transmission patterns have been captured at unprecedented scales (Beckman et al., 2019). However, data alone cannot solve all problems. Only when combined with statistics, computational methods, mathematical modeling, and interdisciplinary collaboration can they truly realize their value (Beckman et al., 2019; Beckman et al., 2020). At the same time, the model is constantly evolving. For example, the application of mechanistic propagation kernels and improved models that take into account secondary propagation processes are gradually improving their predictive power for natural regeneration and ecosystem dynamics (Kim et al., 2022).

Acknowledgments

The author would like to thank Anita W.W. for her insightful suggestions throughout the development of this study.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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