Sex-driven neighborhood effects on herbivory in the dioecious Mediterranean palm Chamaerops humilis L.

Although it is well recognized that the strength of plant-herbivore interactions can vary with the plant sex, the distance, and the density of conspecific neighbors, no study has yet assessed their combined influence. Here, we filled this knowledge gap by focusing on the dioecious palm Chamaerops humilis L., and its two main herbivores, the invasive moth Paysandisia archon Burmeister and the feral goat Capra hircus L. We evaluated levels and spatial patterns of herbivory, as well as those of plant size and number of inflorescences in two palm populations in Mallorca (Balearic Islands, Spain). Our spatial point pattern analyses revealed that palms not affected by moth herbivory or goat florivory were spatially aggregated, goats fed more strongly upon inflorescences in palms with more neighbors, but they consumed more leaves in isolated palms. Interestingly, we could reveal for the first time that plant sex is a key plant trait modulating neighborhood effects. For instance, whereas aggregated female palms experienced lower intensity of goat florivory than isolated ones, male palms showed the opposite pattern. Palm size and number of inflorescences also showed sex-related differences, suggesting that sexual dimorphism is a key driver of the observed neighborhood effects on herbivory. Our study highlights the importance of considering relevant plant traits such as sex when investigating plant neighborhood effects, calling for further research to fully understand the dynamics governing plant-herbivore interactions in dioecious systems.

Accelerated forest fragmentation leads to critical increase in tropical forest edge area.

Large areas of tropical forests have been lost through deforestation, resulting in fragmented forest landscapes. However, the dynamics of forest fragmentation are still unknown, especially the critical forest edge areas, which are sources of carbon emissions due to increased tree mortality. We analyzed the changes in forest fragmentation for the entire tropics using high-resolution forest cover maps. We found that forest edge area increased from 27 to 31% of the total forest area in just 10 years, with the largest increase in Africa. The number of forest fragments increased by 20 million with consequences for connectivity of tropical landscapes. Simulations suggest that ongoing deforestation will further accelerate forest fragmentation. By 2100, 50% of tropical forest area will be at the forest edge, causing additional carbon emissions of up to 500 million MT carbon per year. Thus, efforts to limit fragmentation in the world's tropical forests are important for climate change mitigation.

Consequences of spatial patterns for coexistence in species-rich plant communities.

Ecology cannot yet fully explain why so many tree species coexist in natural communities such as tropical forests. A major difficulty is linking individual-level processes to community dynamics. We propose a combination of tree spatial data, spatial statistics and dynamical theory to reveal the relationship between spatial patterns and population-level interaction coefficients and their consequences for multispecies dynamics and coexistence. Here we show that the emerging population-level interaction coefficients have, for a broad range of circumstances, a simpler structure than their individual-level counterparts, which allows for an analytical treatment of equilibrium and stability conditions. Mechanisms such as animal seed dispersal, which result in clustering of recruits that is decoupled from parent locations, lead to a rare-species advantage and coexistence of otherwise neutral competitors. Linking spatial statistics with theories of community dynamics offers new avenues for explaining species coexistence and calls for rethinking community ecology through a spatial lens.

General statistical scaling laws for stability in ecological systems.

Ecological stability refers to a family of concepts used to describe how systems of interacting species vary through time and respond to disturbances. Because observed ecological stability depends on sampling scales and environmental context, it is notoriously difficult to compare measurements across sites and systems. Here, we apply stochastic dynamical systems theory to derive general statistical scaling relationships across time, space, and ecological level of organisation for three fundamental stability aspects: resilience, resistance, and invariance. These relationships can be calibrated using random or representative samples measured at individual scales, and projected to predict average stability at other scales across a wide range of contexts. Moreover deviations between observed vs. extrapolated scaling relationships can reveal information about unobserved heterogeneity across time, space, or species. We anticipate that these methods will be useful for cross-study synthesis of stability data, extrapolating measurements to unobserved scales, and identifying underlying causes and consequences of heterogeneity.

Habitat filtering drives the local distribution of congeneric species in a Brazilian white-sand flooded tropical forest.

The investigation of ecological processes that maintain species coexistence is revealing in naturally disturbed environments such as the white-sand tropical forest, which is subject to periodic flooding that might pose strong habitat filtering to tree species. Congeneric species are a good model to investigate the relative importance of ecological processes that maintain high species diversity because they tend to exploit the same limiting resources and/or have similar tolerance limits to the same environmental conditions due to their close phylogenetic relationship. We aim to find evidence for the action and relative importance of different processes hypothesized to maintain species coexistence in a white-sand flooded forest in Brazil, taking advantage of data on the detailed spatial structure of populations of congeneric species. Individuals of three Myrcia species were tagged, mapped, and measured for diameter at soil height in a 1-ha plot. We also sampled seven environmental variables in the plot. We employed several spatial point process models to investigate the possible action of habitat filtering, interspecific competition, and dispersal limitation. Habitat filtering was the most important process driving the local distribution of the three Myrcia species, as they showed associations, albeit of different strength, to environmental variables related to flooding. We did not detect spatial patterns, such as spatial segregation and smaller size of nearby neighbors, that would be consistent with interspecific competition among the three congeneric species and other co-occurring species. Even though congeners were spatially independent, they responded to differences in the environment. Last, dispersal limitation only led to spatial associations of different size classes for one of the species. Given that white-sand flooded forests are highly threatened in Brazil, the preservation of their different habitats is of utmost importance to the maintenance of high species richness, as flooding drives the distribution of species in the community.

Network science applied to forest megaplots: tropical tree species coexist in small-world networks.

Network analysis is an important tool to analyze the structure of complex systems such as tropical forests. Here, we infer spatial proximity networks in tropical forests by using network science. First, we focus on tree neighborhoods to derive spatial tree networks from forest inventory data. In a second step, we construct species networks to describe the potential for interactions between species. We find remarkably similar tree and species networks among tropical forests in Panama, Sri Lanka and Taiwan. Across these sites only 32 to 51% of all possible connections between species pairs were realized in the species networks. The species networks show the common small-world property and constant node degree distributions not yet described and explained by network science. Our application of network analysis to forest ecology provides a new approach in biodiversity research to quantify spatial neighborhood structures for better understanding interactions between tree species. Our analyses show that details of tree positions and sizes have no important influence on the detected network structures. This suggests existence of simple principles underlying the complex interactions in tropical forests.

Sperm storage reduces the strength of the mate-finding Allee effect.

Mate searching is a key component of sexual reproduction that can have important implications for population viability, especially for the mate-finding Allee effect. Interannual sperm storage by females may be an adaptation that potentially attenuates mate limitation, but the demographic consequences of this functional trait have not been studied. Our goal is to assess the effect of female sperm storage durability on the strength of the mate-finding Allee effect and the viability of populations subject to low population density and habitat alteration. We used an individual-based simulation model that incorporates realistic representations of the demographic and spatial processes of our model species, the spur-thighed tortoise (Testudo graeca). This allowed for a detailed assessment of reproductive rates, population growth rates, and extinction probabilities. We also studied the relationship between the number of reproductive males and the reproductive rates for scenarios combining different levels of sperm storage durability, initial population density, and landscape alteration. Our results showed that simulated populations parameterized with the field-observed demographic rates collapsed for short sperm storage durability, but were viable for a durability of one year or longer. In contrast, the simulated populations with a low initial density were only viable in human-altered landscapes for sperm storage durability of 4 years. We find that sperm storage is an effective mechanism that can reduce the strength of the mate-finding Allee effect and contribute to the persistence of low-density populations. Our study highlights the key role of sperm storage in the dynamics of species with limited movement ability to facilitate reproduction in patchy landscapes or during population expansion. This study represents the first quantification of the effect of sperm storage durability on population dynamics in different landscapes and population scenarios.

Competition for light and persistence of rare light-demanding species within tree-fall gaps in a moist tropical forest.

Current evidence suggests that tree-fall gaps can influence forest structure and dynamics by enabling certain species guilds to persist over the long term. Here we assessed the development of local size hierarchies and asymmetric competition for light in tree-fall gaps, and the role played by these two processes for the persistence of rare light-demanding species in the Barro Colorado Island Forest Dynamics Plot (Panama). We performed spatial point pattern analysis, considering both the spatial locations (x,y) and the diameter at breast height (DBH) of all the woody plant recruits from the 1985 and 2000 censuses located in tree-fall gaps, and followed their fate up to the 1990-2010 and 2005-2010 censuses, respectively. For these two recruit cohorts, we found that, from the initial census until 5-10 yr later, close neighbors presented a larger DBH than the mean DBH of all individuals within gaps, which points to a positive growth response of recruits to the increased light levels in the gap centers. However, close neighbors of the 1985 cohort also showed larger than expected DBH differences that disappeared in subsequent censuses, indicating an enhancement of size differences between neighbors and the mortality of the smaller individuals. Finally, for both recruit cohorts, we found that 10-15 yr after gap formation, surviving individuals of rare light-demanding species had a negative impact on survival of neighboring individuals of other species. Our results indicate that gaps favor the persistence of rare light-demanding species through the development of local size hierarchies and asymmetric competition for light. The strength of this process, however, apparently depends upon gap size and the role played by the woody plants already existing at the time of gap formation in early colonization. Moreover, our findings suggest that in this forest, gaps may enhance colonization of plant species typical of nearby dry tropical areas, and that, over the coming decades, similar processes could strongly modify the structure and dynamics of moist tropical forests in the region.

Integrating the underlying structure of stochasticity into community ecology.

Stochasticity is a core component of ecology, as it underlies key processes that structure and create variability in nature. Despite its fundamental importance in ecological systems, the concept is often treated as synonymous with unpredictability in community ecology, and studies tend to focus on single forms of stochasticity rather than taking a more holistic view. This has led to multiple narratives for how stochasticity mediates community dynamics. Here, we present a framework that describes how different forms of stochasticity (notably demographic and environmental stochasticity) combine to provide underlying and predictable structure in diverse communities. This framework builds on the deep ecological understanding of stochastic processes acting at individual and population levels and in modules of a few interacting species. We support our framework with a mathematical model that we use to synthesize key literature, demonstrating that stochasticity is more than simple uncertainty. Rather, stochasticity has profound and predictable effects on community dynamics that are critical for understanding how diversity is maintained. We propose next steps that ecologists might use to explore the role of stochasticity for structuring communities in theoretical and empirical systems, and thereby enhance our understanding of community dynamics.

Dataset on the diversity of plant-parasitic nematodes in cultivated olive trees in southern Spain.

Datasets presented here were employed in the main work "Spatial structure and soil properties shape local community structure of plant-parasitic nematodes in cultivated olive trees in southern Spain" Archidona-Yuste et al., 2020. In this research, we aimed to unravel the diversity of plant-parasitic nematodes (PPN) associated with cultivated olive (Olea europaea subsp. europaea var. europaea) in southern Spain, Andalusia. The olive growing area of Andalusia is of high agriculture and socio-economic importance with an extensive distribution of this crop. To this end, we conducted a systematic survey comprising 376 commercial olive orchards covering the diversity of cropping systems applied. Data showed 128 species of PPN belonging to 38 genera and to 13 families. In addition, an extensive data set regarding to potential factors in structuring the community patterns of PPN found in the 376 commercial olive orchards sampled is provided. Three variables data set were compiled including above-ground environment, soil and agronomic management. Overall, 48 explanatory variables were selected as determinist processes on shaping the diversity of PPN. Finally, data also showed the values regarding to the partition of beta diversity into contributions of single sites to overall beta diversity (LCBD) and intro contributions of individual species to overall beta diversity (SCBD). Data may serve as benchmarks for other groups working in the field of PPN diversity associated with crops and of belowground communities and ecosystems.

Projected impacts of climate change on functional diversity of frugivorous birds along a tropical elevational gradient.

Climate change forces many species to move their ranges to higher latitudes or elevations. Resulting immigration or emigration of species might lead to functional changes, e.g., in the trait distribution and composition of ecological assemblages. Here, we combined approaches from biogeography (species distribution models; SDMs) and community ecology (functional diversity) to investigate potential effects of climate-driven range changes on frugivorous bird assemblages along a 3000 m elevational gradient in the tropical Andes. We used SDMs to model current and projected future occurrence probabilities of frugivorous bird species from the lowlands to the tree line. SDM-derived probabilities of occurrence were combined with traits relevant for seed dispersal of fleshy-fruited plants to calculate functional dispersion (FDis; a measure of functional diversity) for current and future bird assemblages. Comparisons of FDis between current and projected future assemblages showed consistent results across four dispersal scenarios, five climate models and two representative concentration pathways. Projections indicated a decrease of FDis in the lowlands, an increase of FDis at lower mid-elevations and little changes at high elevations. This suggests that functional dispersion responds differently to global warming at different elevational levels, likely modifying avian seed dispersal functions and plant regeneration in forest ecosystems along tropical mountains.

Disentangling the functional trait correlates of spatial aggregation in tropical forest trees.

Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.

The Latitudinal Diversity Gradient: Novel Understanding through Mechanistic Eco-evolutionary Models.

The latitudinal diversity gradient (LDG) is one of the most widely studied patterns in ecology, yet no consensus has been reached about its underlying causes. We argue that the reasons for this are the verbal nature of existing hypotheses, the failure to mechanistically link interacting ecological and evolutionary processes to the LDG, and the fact that empirical patterns are often consistent with multiple explanations. To address this issue, we synthesize current LDG hypotheses, uncovering their eco-evolutionary mechanisms, hidden assumptions, and commonalities. Furthermore, we propose mechanistic eco-evolutionary modeling and an inferential approach that makes use of geographic, phylogenetic, and trait-based patterns to assess the relative importance of different processes for generating the LDG.

Global patterns of tropical forest fragmentation.

Remote sensing enables the quantification of tropical deforestation with high spatial resolution. This in-depth mapping has led to substantial advances in the analysis of continent-wide fragmentation of tropical forests. Here we identified approximately 130 million forest fragments in three continents that show surprisingly similar power-law size and perimeter distributions as well as fractal dimensions. Power-law distributions have been observed in many natural phenomena such as wildfires, landslides and earthquakes. The principles of percolation theory provide one explanation for the observed patterns, and suggest that forest fragmentation is close to the critical point of percolation; simulation modelling also supports this hypothesis. The observed patterns emerge not only from random deforestation, which can be described by percolation theory, but also from a wide range of deforestation and forest-recovery regimes. Our models predict that additional forest loss will result in a large increase in the total number of forest fragments-at maximum by a factor of 33 over 50 years-as well as a decrease in their size, and that these consequences could be partly mitigated by reforestation and forest protection.

Spatio-temporal arrangement of Chamaerops humilis inflorescences and occupancy patterns by its nursery pollinator, Derelomus chamaeropsis.

Background and Aims: Nursery pollination is a highly specialized interaction in which pollinators breed inside plant reproductive structures. Pollinator occupancy of host plants often depends on plant location, flowering synchrony and sex. The nursery pollination system between the dioecious dwarf palm Chamaerops humilis (Arecaceae) and the host-specific palm flower weevil Derelomus chamaeropsis was investigated. For the first time, sex, flowering synchrony and spatial distribution of plants was related to the occupancy probability and the abundance of D. chamaeropsis larvae, important traits influencing both pollinator and plant fitness. Methods: During the flowering season, all inflorescences in anthesis were counted every 12 d and a flowering synchrony index was calculated taking into account all possible correlations with generalized linear mixed models. To analyse the spatial structure of plants, larva occupancy and abundance, different techniques of spatial point pattern analysis were used. Key results: In total, 5986 larvae in 1063 C. humilis inflorescences were recorded over three consecutive seasons. Male inflorescences showed a higher presence and abundance of weevil larvae than females, but interestingly approx. 30 % of the females held larvae. Also, larvae occurred mainly in highly synchronous plants with a low number of inflorescences, perhaps because those plants did not lead to a resource dilution effect. There was no evidence of spatial patterns in larva occupancy or abundance at any spatial scale, suggesting high dispersal ability of adult weevil. Conclusions: The results in a nursery-pollinated dioecious palm demonstrate that plant sex, flowering display and flowering synchrony act as additive forces influencing the presence and abundance of the specialized pollinator larvae. Contradicting previous results, clear evidence that female dwarf palms also provide rewarding oviposition sites was found, and thus the plant 'pays' for the pollination services. The findings highlight that plant local aggregation is not always the main determinant of pollinator attraction, whereas flower traits and phenology could be critical in specialized plant-pollinator interactions.

Distance-dependent seedling mortality and long-term spacing dynamics in a neotropical forest community.

Negative distance dependence (NDisD), or reduced recruitment near adult conspecifics, is thought to explain the astounding diversity of tropical forests. While many studies show greater mortality at near vs. far distances from adults, these studies do not seek to track changes in the peak seedling curve over time, thus limiting our ability to link NDisD to coexistence. Using census data collected over 12 years from central Panama in conjunction with spatial mark-connection functions, we show evidence for NDisD for many species, and find that the peak seedling curve shifts away from conspecific adults over time. We find wide variation in the strength of NDisD, which was correlated with seed size and canopy position, but other life-history traits showed no relationship with variation in NDisD mortality. Our results document shifts in peak seedling densities over time, thus providing evidence for the hypothesized spacing mechanism necessary for diversity maintenance in tropical forests.

What drives the spatial distribution and dynamics of local species richness in tropical forest?

Understanding the structure and dynamics of highly diverse tropical forests is challenging. Here we investigate the factors that drive the spatio-temporal variation of local tree numbers and species richness in a tropical forest (including 1250 plots of 20 × 20 m2). To this end, we use a series of dynamic models that are built around the local spatial variation of mortality and recruitment rates, and ask which combination of processes can explain the observed spatial and temporal variation in tree and species numbers. We find that processes not included in classical neutral theory are needed to explain these fundamental patterns of the observed local forest dynamics. We identified a large spatio-temporal variability in the local number of recruits as the main missing mechanism, whereas variability of mortality rates contributed to a lesser extent. We also found that local tree numbers stabilize at typical values which can be explained by a simple analytical model. Our study emphasized the importance of spatio-temporal variability in recruitment beyond demographic stochasticity for explaining the local heterogeneity of tropical forests.

Functionally specialised birds respond flexibly to seasonal changes in fruit availability.

Interactions between resource and consumer species result in complex ecological networks. The overall structure of these networks is often stable in space and time, but little is known about the temporal stability of the functional roles of consumer species in these networks. We used a trait-based approach to investigate whether consumers (frugivorous birds) show similar degrees of functional specialisation on resources (plants) in ecological networks across seasons. We additionally tested whether closely related bird species have similar degrees of functional specialisation and whether birds that are functionally specialised on specific resource types within a season are flexible in switching to other resource types in other seasons. We analysed four seasonal replicates of two species-rich plant-frugivore networks from the tropical Andes. To quantify fruit preferences of frugivorous birds, we projected their interactions with plants into a multidimensional plant trait space. To measure functional specialisation of birds, we calculated a species' functional niche breadth (the extent of seasonal plant trait space utilised by a particular bird) and functional originality (the extent to which a bird species' fruit preference functionally differs from those of other species in a seasonal network). We additionally calculated functional flexibility, i.e. the ability of bird species to change their fruit preference across seasons in response to variation in plant resources. Functional specialisation of bird species varied more among species than across seasons, and phylogenetically similar bird species showed similar degrees of functional niche breadth (phylogenetic signal λ = 0·81) and functional originality (λ = 0·89). Additionally, we found that birds with high functional flexibility across seasons had narrow functional niche breadth and high functional originality per season, suggesting that birds that are seasonally specialised on particular resources are most flexible in switching to other fruit resources across seasons. The high flexibility of functionally specialised bird species to switch seasonally to other resources challenges the view that consumer species rely on functionally similar resources throughout the year. This flexibility of consumer species may be an important, but widely neglected mechanism that could potentially stabilise consumer-resource networks in response to human disturbance and environmental change.

Spatial patterns of an endemic Mediterranean palm recolonizing old fields.

Throughout Europe, increased levels of land abandonment lead to (re)colonization of old lands by forests and shrublands. Very little is known about the spatial pattern of plants recolonizing such old fields. We mapped in two 21-22-ha plots, located in the Doñana National Park (Spain), all adult individuals of the endozoochorous dwarf palm Chamaerops humilisL. and determined their sex and sizes. We used techniques of spatial point pattern analysis (SPPA) to precisely quantify the spatial structure of these C. humilis populations. The objective was to identify potential processes generating the patterns and their likely consequences on palm reproductive success. We used (1) Thomas point process models to describe the clustering of the populations, (2) random labeling to test the sexual spatial segregation, and (3) mark correlation functions to assess spatial structure in plant sizes. Plants in both plots showed two critical scales of clustering, with small clusters of a radius of 2.8-4 m nested within large clusters with 38-44 m radius. Additional to the clustered individuals, 11% and 27% of all C. humilis individuals belonged to a random pattern that was independently superimposed to the clustered pattern. The complex spatial pattern of C. humilis could be explained by the effect of different seed-dispersers and predators' behavior and their relative abundances. Plant sexes had no spatial segregation. Plant sizes showed a spatial aggregation inside the clusters, with a decreasing correlation with distance. Clustering of C. humilis is strongly reliant on its seed dispersers and stressful environmental conditions. However, it seems that the spatial patterns and dispersal strategies of the dwarf palm make it a successful plant for new habitat colonization. Our results provide new information on the colonization ability of C. humilis and can help to develop management strategies to recover plant populations.

Correction: Persistence of Neighborhood Demographic Influences over Long Phylogenetic Distances May Help Drive Post-Speciation Adaptation in Tropical Forests.

[This corrects the article DOI: 10.1371/journal.pone.0156913.].