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The spatiotemporal genetic variation at early plant life stages may substantially affect the natural recolonization of human-altered areas, which is crucial to understand plant and habitat conservation. In animal-dispersed plants, dispersers’ behavior may critically drive the distribution of genetic variation. Here, we examine how genetic rarity is spatially and temporally structured in seedlings of a keystone pioneer palm (Chamaerops humilis) and how the variation of genetic rarity could ultimately affect plant recruitment. We intensively monitored the seed rain mediated by two medium-sized carnivores during two consecutive seasons in a Mediterranean human-altered area. We genotyped 143 out of 309 detected seedlings using 12 microsatellite markers. We found that seedlings emerging from carnivore-dispersed seeds showed moderate to high levels of genetic diversity and no evidence of inbreeding. We found inflated kinship among seedlings that emerged from seeds within a single carnivore fecal sample, but a dilution of such FSGS at larger spatial scales (e.g. latrine). Seedlings showed a significant genetic sub-structure and the sibling relationships varied depending on the spatial scale. Rare genotypes arrived slightly later throughout the dispersal season and tended to be spatially isolated. However, genetic rarity was not a significant predictor by itself which indicates that, at least, its influence on seedling survival was smaller than other spatiotemporal factors. Our results suggest strong C. humilis resilience to genetic bottlenecks due to human disturbances. We highlight the study of plant-animal interactions from a genetic perspective since it provides crucial information for plant conservation and the recovery of genetic plant resilience.

Seed dispersal is an essential ecological process for plant (re)colonization, especially in intensively human-altered habitats such as old-fields (i.e. abandoned farmlands) where seed arrival is often limited. Assessment of spatial patterns of mammal-generated seed rain and their matching with the spatial distribution of adult plants provides essential information on the patterns and pace of the (re)colonization processes. For instance, increased seedling survival far from adult plants could indicate density-dependent mortality (Janzen-Connell) effects whereas increased survival close to adult plants could suggest facilitative effects. Here, we characterized during two dispersal seasons the spatial distribution of feces from four frugivorous mammals and quantified its spatial association with the distribution of adult plants of the pioneer Mediterranean dwarf palm (Chamaerops humilis) in two old-fields in southwestern Spain. We also estimated the dispersal kernels of both emerged and surviving seedlings and assessed potential evidence for Janzen-Connell and/or facilitative effects. Using a spatially explicit approach, we revealed strong differences between study sites in the strength and scale of spatial associations between mammal feces with C. humilis seeds and adult dwarf palms, being strongly positive at small scales in one site and slightly positive at larger scales in the other one. Further, we found some evidence of both Janzen-Connell and facilitative effects depending on the study site. Altogether, our results emphasize the central role as seed disperser of the Eurasian badger (Meles meles) in the natural (re)colonization of Mediterranean old-fields and the spatial variations of the underlying mechanisms and demographic consequences for plant populations.

[Question]: Nurse–beneficiary plant interactions are often used to restore degraded habitats. However, whether and how shifts in plant–plant interactions along the facilitation–competition continuum alter revegetation success has been seldom considered. To test whether and how shifts in plant–plant interactions (due to woody species identity, study site, early life stage, and individual nurse) might alter plant recruitment and thus the success of revegetation projects, we chose a system comprising the Mediterranean dwarf palm (Chamaerops humilis) and seven common woody plant species. [Location]: Two human-degraded sites within Doñana National Park (southwestern Spain). [Methods]: We carried out several well-replicated field experiments to compare plant performance (seed survival, seedling emergence, seedling survival, seedling recruitment) in the presence and absence of Chamaerops humilis. [Results]: Chamaerops humilis had marked effects on the performance of woody species that, however, changed among life stages. Depending on woody species identity, seed survival was up to 193 times greater in adjacent open spaces than beneath Chamaerops humilis. Conversely, seedling survival and recruitment were up to 19 times greater beneath Chamaerops humilis than in open spaces. Importantly, none of the studied woody species showed greater accumulated recruitment in open spaces than beneath Chamaerops humilis. Interestingly, we found strong inter-individual palm variation in the sign and strength of their effect on woody plant performance. [Conclusions]: We found strong seed–seedling conflicts the strength of which was species-specific. The strong inter-individual palm variation depicts a facilitation–competition continuum with important implications for restoration. We propose several management recommendations across different hierarchical levels (i.e., from individuals to communities) that may increase plant recruitment and therefore the success of revegetation projects. Our

Investigating spatial variation in the relative importance of sexual reproduction and clonal propagation is critical to obtain more accurate estimates of future effective population sizes and genetic diversity, as well as to identify ecological correlates of clonality. We combined a stratified sampling scheme with microsatellite genetic analyses to estimate variation in the proportion of sexual versus clonal recruits among saplings in five populations of the tree Pyrus bourgaeana. Using a likelihood framework, we identified clones among the genotypes analysed and examined variation among populations regarding the proportion of saplings coming from clonal propagation. We also examined the relationship between the relative abundance of clonal shoots across the studied populations and their herbivory levels. Our results revealed that one third of the saplings examined (N = 225 saplings) had a probability above 0.9 of being clones of nearby (<10 m) trees, with the ratio between clonal propagation and sexual recruitment varying up to eight-fold among populations. A small portion of these putative clonal shoots reached sexual maturity. Relative abundance of clonal shoots did not significantly relate to the herbivory by ungulates. Our results call into question optimistic expectations of previous studies reporting sufficient levels of recruitment under parental trees without animal seed dispersal services. Nevertheless, given that some of these clonal shoots reach sexual maturity, clonal propagation can ultimately facilitate the long-term persistence of populations during adverse periods (e.g. environmental stress, impoverished pollinator communities, seed dispersal limitation).

Restricted seed dispersal frequently leads to fine-scale spatial genetic structure (i.e., FSGS) within plant populations. Depending on its spatial extent and the mobility of pollinators, this inflated kinship at the immediate neighbourhood can critically impoverish pollen quality. Despite the common occurrence of positive FSGS within plant populations, our knowledge regarding the role of long-distance pollination preventing reproductive failure is still limited. Using microsatellite markers, we examined the existence of positive FSGS in two low-density populations of the tree Pyrus bourgaeana. We also designed controlled crosses among trees differing in their kinship to investigate the effects of increased local kinship on plant reproduction. We used six pollination treatments and fully monitored fruit production, fruit and seed weight, proportion of mature seeds per fruit, and seed germination. Our results revealed positive FSGS in both study populations and lower fruit initiation in flowers pollinated with pollen from highly-genetically related individuals within the neighbourhood, with this trend intensifying as the fruit development progressed. Besides, open-pollinated flowers exhibited lower performance compared to those pollinated by distant pollen donors, suggesting intense qualitative pollen limitation in natural populations. We found positive fine-scale spatial genetic structure is translated into impoverished pollen quality from nearby pollen donors which negatively impacts the reproductive success of trees in low-density populations. Under this scenario of intrapopulation genetic rescue by distant pollen donors, the relevance of highly-mobile pollinators for connecting spatially and genetically distant patches of trees may be crucial to safeguarding population recruitment.

One advantageous strategy for the restoration of human-disturbed landscapes is the use of ecologically “important” species such as nurse plants. We propose a field-based approach to measure the functional importance of nurse species (i.e. their relative facilitative effects on other plant species) and to identify which species will yield more efficient revegetation programs regarding their abundance. We identified 30 nurse-beneficiary spatial associations, with the functional importance varying largely among four nurse species and three human-disturbed areas. A Mediterranean endemic palm was the most important nurse species, thus showing its potential key role in revegetation programs by promoting spatial associations with late-successional plant species. We encourage restorers to use nurse species with a disproportionate (regarding their relative abundance) impact on ecosystems to save additional resources.

The study of plant-frugivore interactions is essential to understand the ecology and evolution of many plant communities. However, very little is known about how interactions among frugivores indirectly affect plant reproductive success. In this study, we examined direct interactions among vertebrate frugivores sharing the same fruit resources. Then, we inferred how the revealed direct interspecific interactions could lead to indirect (positive or negative) effects on reproductive success of fleshy fruited plants. To do so, we developed a new analytical approach that combines camera trap data (spatial location, visitor species, date and time, activity) and tailored null models that allowed us to infer spatial-temporal interactions (attraction, avoidance or indifference) between pairs of frugivore species. To illustrate our approach, we chose to study the system composed by the Mediterranean dwarf palm, Chamaerops humilis, the Iberian pear tree, Pyrus bourgaeana, and their shared functionally diverse assemblages of vertebrate frugivores in a Mediterranean area of SW Spain. We first assessed the extent to which different pairs of frugivore species tend to visit the same or different fruiting individual plants. Then, for pairs of species that used the same individual plants, we evaluated their spatial-temporal relationship. Our first step showed, for instance, that some prey frugivore species (e.g. lagomorphs) tend to avoid those C. humilis individuals that were most visited by their predators (e.g. red foxes). Also, the second step revealed temporal attraction between large wild and domestic frugivore ungulates (e.g. red deer, cows) and medium-sized frugivores (e.g. red foxes) suggesting that large mammals could facilitate the C. humilis and P. bourgaeana exploitation to other smaller frugivores by making fruits more easily accessible. Finally, our results allowed us to identify direct interaction pathways, that revealed how the mutualistic and antagonistic relations

[Background]: Disentangling the drivers of genetic differentiation is one of the cornerstones in evolution. This is because genetic diversity, and the way in which it is partitioned within and among populations across space, is an important asset for the ability of populations to adapt and persist in changing environments. We tested three major hypotheses accounting for genetic differentiation—isolation-by-distance (IBD), isolation-by-environment (IBE) and isolation-by-resistance (IBR)—in the annual plant Arabidopsis thaliana across the Iberian Peninsula, the region with the largest genomic diversity. To that end, we sampled, genotyped with genome-wide SNPs, and analyzed 1772 individuals from 278 populations distributed across the Iberian Peninsula., [Results]: IBD, and to a lesser extent IBE, were the most important drivers of genetic differentiation in A. thaliana. In other words, dispersal limitation, genetic drift, and to a lesser extent local adaptation to environmental gradients, accounted for the within- and among-population distribution of genetic diversity. Analyses applied to the four Iberian genetic clusters, which represent the joint outcome of the long demographic and adaptive history of the species in the region, showed similar results except for one cluster, in which IBR (a function of landscape heterogeneity) was the most important driver of genetic differentiation. Using spatial hierarchical Bayesian models, we found that precipitation seasonality and topsoil pH chiefly accounted for the geographic distribution of genetic diversity in Iberian A. thaliana., [Conclusions]: Overall, the interplay between the influence of precipitation seasonality on genetic diversity and the effect of restricted dispersal and genetic drift on genetic differentiation emerges as the major forces underlying the evolutionary trajectory of Iberian A. thaliana.

Understanding the mechanisms underlying the recolonization of old-fields is critical to promote the recovery of the ecosystem functioning, particularly in regions where agricultural abandonment has increased in the last 60 years. Given that seed arrival and seedling survival often limit the recolonization process by woody species in many Mediterranean habitats, the ‘perching’ and ‘nursing’ effects exerted by some pioneer species could be crucial for the restoration of such abandoned lands. We examined the role of an endemic Mediterranean palm (Chamaerops humilis) on the recolonization of old-fields by woody species in southern Iberian Peninsula. We chose three independent old-fields differing in their shrub encroachment levels. To identify potential facilitation by C. humilis, we used a spatially-explicit approach and analyzed its spatial associations with ten common woody species (e.g. Asparagus spp., Daphne gnidium, Olea europaea var. sylvestris, Pyrus bourgaeana). We detected positive spatial associations between C. humilis and woody species at small-scale (1–5 m) in the three plots. Most of such small-scale associations were linked to the bird-dispersal of woody species. Nonetheless, there were marked differences among plots in spatial associations between C. humilis and woody plants, being Asparagus spp. the only species positively associated with C. humilis within the three studied old-fields. These species-specific differences were likely related to variations among old-fields in encroachment level and the legacy of human management. Such small-scales spatial associations between C. humilis and woody species across Iberian old-fields were linked to the perching and nursing effects exerted by the palm. We emphasize the strong potential of this pioneer Mediterranean palm for the restoration of native ecosystems and the recovery of ecosystems services.

Plant provenance, plant facilitation and the characteristics of restoration sites are critical determinants of revegetation success. Surprisingly, however, very little is known about how these key factors interact, both with each other and to complex patterns of recruitment in target plant species. We investigated, for the first time, the potential of combining seed provenance, plant facilitation and restoration site characteristics to guide revegetation efforts. As a model system, we used the Iberian pear (Pyrus bourgaeana) and three potential nurse shrub species. To this end, we completed a comprehensive reciprocal transplant experiment in five P. bourgaeana populations within a fragmented landscape at the Doñana Biosphere Reserve (SW Spain). There were marked differences among provenances for seedling emergence and growth, as well as for recruitment, but no evidence of local adaptation was found. Shrubs generally had a positive or neutral effect on P. bourgaeana seedling performance with, for example survival underneath shrubs being, on average, twice as high as in open microhabitats. Interestingly, however, the magnitude and even the sign of such plant–plant interactions changed with both sowing locality and seed provenance. For instance, shrubs enhanced seedling survival and recruitment at the end of the study in four of the five sowing localities, whereas it decreased it in the remaining locality. Also, although nurse shrubs lessened seedling size of two provenances, it did not have a significant effect on the other three. Synthesis and applications. We show how revegetation success can be maximized, not only by selecting the most appropriate plant material, taking advantage of plant facilitative interactions and selecting the right restoration site, but also by using these factors in an integrated fashion. We propose local managers efficiently undertake future revegetation campaigns by mixing seeds from several genetically adequate source populations in numbe

Seed predation is a key demographic process for plant population dynamics and community structure. The strength of seed predation for a given plant species often varies among populations depending on both plant traits (e.g. seed size) and extrinsic factors (e.g. seed predator abundance). Because both types of factors tend to covary spatially, identifying their relative contribution to changes in seed predation through observational approaches is challenging. During two consecutive years, we reciprocally translocated seeds of Iberian pear (Pyrus bourgaeana) from five localities in SW Spain to estimate changes in interaction strength between the tree and its seed-eaters (mostly rodents). We also measured relevant seed traits, the abundance of granivore rodents, and the density of fruiting-shrubs in the same five populations. Both seed traits and extrinsic factors varied among the five studied populations. As predicted, we found significant spatial covariance between some extrinsic factors (i.e. rodent abundance and fruiting-shrub density) as well as between extrinsic factors and plant traits (seed size). Seed removal likelihood varied highly among offering localities, with differences up to 11.6-fold. However, and despite differences in seed traits among populations, no evidence of seed predator preference for seeds from any locality was found in any of the five local experiments. Our study indicated that extrinsic factors (e.g. seed predator abundance) rather than measured seed traits mediated among-population variation in seed predation. Selection pressures exerted by seed predators on seed traits could thus be less pervasive than previously thought. Our experimental approach can help to better understand the potential of different functional animal groups (pollinators, herbivores, granivores, frugivores) to exert selection pressures on plant traits.

Margins of distribution of plant species constitute natural areas where the impact of the antagonistic interactions is expected to be higher and where changes in the dynamics of plant-herbivore coevolution could promote intraspecific differentiation in (co)evolving plant traits. In the present study, we investigated how differences in the average herbivory level affect maternal fitness in core continuous and marginal disjunct populations of Daphne laureola in an effort to assess the role of herbivores limiting plant distribution. Furthermore, we investigated intraspecific differentiation in vegetative traits and their potential connection to divergent selection by herbivores in both groups of populations. Our results did not support increased herbivory at the species margin but did support a difference in the effect of herbivory on maternal fitness between core continuous and marginal disjunct populations of D. laureola. In addition, herbivores did not exert phenotypic selection consistent with the geographic variation in studied plant traits. Therefore, the geographic variation of vegetative traits of D. laureola seems to be consequence of environmental heterogeneity more than result of geographically divergent selection by herbivores.

Several ecological conditions and processes occurring naturally in plant populations may lead to spatial aggregation of sexes within populations of sexually polymorphic species. In addition, ecological disturbances such as forest management or fire could also affect the spatial distribution of sexes within populations. Spatial aggregation of sexes can have important consequences for the fitness of the individuals in sex-biased patches through increased pollen limitation and/or variation in the male fitness of hermaphrodites. Therefore, spatial aggregation of sexes could be relevant for the maintenance of the sexual polymorphism in plant species. Here, we used point pattern analysis to study the spatial distribution of female and hermaphrodite individuals within a single population of the gynodioecious understory shrub Daphne laureola, inhabiting a young pine reforestation and the adjacent natural undisturbed area. In the undisturbed area, we found that hermaphrodites were distributed randomly whereas females were spatially aggregated at short distances. Such aggregation could result from narrower ecological amplitude, cyto-nuclear determination of sex and limited dispersal and/or increased cloning in females. In the reforested area, females were slightly more abundant and the two sexes were randomly distributed among all plants. Recolonization processes after certain forest disturbances could thus favour the establishment of females and alter the spatial distribution of sexes in this understory species. © 2012.

Según la teoría del mosaico geográfico de coevolución, las variaciones espaciales en la magnitud del flujo génico y en el signo e intensidad de la selección ejercida por animales sobre los rasgos de las plantas, son los dos elementos esenciales que dan lugar a la aparición de mosaicos de selección y determinan el grado de adaptación entre las especies de plantas y animales. Hay ciertas situaciones naturales donde cabe esperar que se produzcan, de manera predecible y a lo largo de extensiones geográficas relativamente pequeñas, gradientes en la magnitud del flujo génico, así como en la forma e intensidad de selección por probables variaciones en la composición y/o abundancia de los polinizadores y herbívoros. Uno de estos lugares dotados de una singularidad ecológica propia son los márgenes de distribución de las especies vegetales. Además de sus bien conocidas peculiaridades ecológicas, los márgenes de distribución también presentan peculiaridades genéticas que los convierten en lugares ideales para verificar las predicciones de la teoría del mosaico geográfico de coevolución. El trabajo de esta tesis doctoral se diseñó para verificar la siguiente predicción general, basada en los postulados de la teoría del mosaico geográfico de coevolución: gradientes en la magnitud del flujo génico que sean consistentes con gradientes de selección por animales (polinizadores y herbívoros) sobre algunos caracteres vegetales, darán lugar a diferenciación intraespecífica en los caracteres sujetos a selección. Esta predicción se abordó mediante el estudio de patrones de selección por animales (polinizadores y herbívoros), variación fenotípica y flujo génico replicado en grupos de poblaciones centrales y marginales del arbusto ginodioico Daphne laureola. A continuación se presenta un resumen de cada uno de los capítulos incluidos en esta tesis doctoral, señalando los resultados principales de cada uno de ellos.

Biogeographic models predict that marginal populations should be more geographically isolated and smaller than central populations, linked to more stressful conditions and likely also to a reduction in density of individuals, individual growth, survival and reproductive output. This variation in population features could have important consequences for different aspects of plant ecology such as individual reproductive success, population genetic structure or plant-animal interactions. In this study, we analyze if individuals of the evergreen shrub Daphne laureola at disjunt populations in a local border of its distribution area in southern Iberian Peninsula differ in individual size, shoot growth, reproductive output and the pollination environment from central continuous populations within the area. Plants of central continuous populations were larger and produced more flowers and fruits than plants of marginal disjunct populations suggesting more optimal conditions, although they had lower annual shoot growth. In contrast, fruit set was higher in plants at the local border, suggesting a more efficient pollinator service in these populations where the main pollinator in central continuous populations, the pollen beetle Meligethes elongatus, was not present. Our results do not support strong differences in the ecological stress between marginal disjunct and central continuous populations of D. laureola in the south of the Iberian Peninsula but indicate some changes in plant-pollinator interactions that could be relevant for the sexual polymorphism in this gynodioecious species.