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., This work was supported by the Portuguese Foundation for Science and Technology, P.J.G. (SFRH/BD/130527/2017), A.R.C.(SFRH/BPD/115781/2016), and by a grant of the Spanish Min-istry of Education and Science (PGC2018-094808-B-I00) toJ.M.F.

[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 results are particularly relevant for restoring arid, semi-arid and alpine landscapes worldwide where the nurse–beneficiary plant interactions are critical to ameliorating stressful conditions., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017) and ARC (SFRH/BPD/115781/2016); and by a grant of the Spanish Ministry of Education and Science to JMF (PGC2018-094808-B- I00).

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., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017) and ARC (SFRH/BPD/115781/2016); and by a grant of the Spanish Ministry of Science and Innovation to JMF (PGC2018-094808-B-I00). Logistic and technical support was provided by ICTS-RBD., Peer reviewed

Although farmlands are the most extensive terrestrial biomes, the abandonment of traditional agriculture in many parts of the world has brought opportunities and challenges for the restoration of such human-disturbed habitats. Seed arrival is a crucial necessary ecological process during plant recolonization that can be enhanced by the use of the so-called “perch plants”. Little is known, however, about whether the seed arrival via frugivorous birds is affected by the spatial distribution of the perch plants in disturbed habitats. To evaluate several spatial aspects of “perching” effect, we used a spatially explicit approach in two disturbed plots within the Doñana National Park (SW Spain). Specifically, we chose as study system the pioneer Mediterranean dwarf palm Chamaerops humilis L., which is often used as a perch by a variety of frugivorous bird species. A total of 289 C. humilis individuals were sampled in search of bird feces (N = 2998) and dispersed seeds (N = 529). Recorded seeds belonged to six different woody species from five different families. Nine bird species from six different families were recorded using C. humilis as perches. GLMs analyses indicated that taller C. humilis males with higher numbers of spatially associated woody species received more dispersed seeds. We detected a random spatial structure of bird feces and dispersed seeds in one study plot, while a nonrandom spatial structure was found in the other one, where isolated C. humilis received a higher number of bird feces and dispersed seeds than expected under spatial null models. The difference in spatial patterns between both study plots could relate, among other factors, to their different state of development in the ecological succession. Most of dispersed seeds were concentrated in a small number of C. humilis individuals, usually male and large ones, that acted as “hotspots” of seed arrival. The fact that frugivorous birds in one study site visited most often isolated C. humilis questions the aggregated spatial structure of revegetation designs typically used in restoration projects. This study reveals novel spatial aspects of the “perching” effect which could be helpful in the restoration of human-disturbed habitats worldwide., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017) and by a grant of the Spanish Ministry of Education and Science to JMF (PGC2018-094808-B-I00). Logistic and technical support was provided by ICTS-RBD. The authors acknowledge support of the publication fee by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI)

Pollination service is crucial to achieve successful plant sexual reproduction and long-term population persistence. This pollination service can be affected by plant conspecific density and also by intrinsic features of individuals related to their flowering phenology and floral display. However, studies examining intrinsic and extrinsic traits on pollinator visitation and reproductive success of Mediterranean trees with limited reproduction are still scarce. We analyzed the effects of plant isolation, flowering phenology, flower weight and tree size on flower visitation probability, flowering patch visitation probability, fruit set and crop size. To this end, we intensively monitored pollinator visitation and fruit production of 67 (in 2019) and 73 (in 2020) Pyrus bourgaeana Decne trees within a threatened Mediterranean population. Our results revealed that isolated individuals received more pollinators than those on conspecific aggregations, suggesting intraspecific competition for pollinators in dense flowering neighborhoods. However, fruit set was higher in trees close to flowering conspecifics despite having fewer visits from pollinators, suggesting pollen limitation but not pollinator limitation in spatially isolated trees. Interestingly, we found increased crop sizes in spatially isolated trees which could be related to reduced intraspecific competition for resources in low-density neighborhoods (water, nutrients) and/or to higher reproductive investment (i.e. higher flower production). Overall, our results indicated pollen but not pollinator limitation in spatially isolated trees. Under this scenario of sexual reproduction mediated by pollinators, our findings stress the relevance of individuals' spatial distribution for self-incompatible trees exhibiting low individuals’ densities., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017) and by a grant of the Spanish Ministry of Education and Science to JMF (PGC2018-094808-B-I00).

Plant–animal interactions fall within a mutualism–antagonism continuum, exerting a wide range of effects on plant reproductive success. These effects become even more complex and diverse when several disparate animal species interact with the same plant species. Despite the increasing number of studies about the influence of herbivory on plant performance, the outcomes mediated by pollination and the combined impact of multiple herbivores on pollination-specialized plants are underexplored. In this study, we chose the Mediterranean dwarf palm Chamaerops humilis (Arecaceae) to illustrate the isolated and joint effect of two contrasting introduced herbivores, the palm borer Paysandisia archon (Lepidoptera, Castniidae) and feral goats, on pollinator abundance and plant reproductive success. To this aim, we monitored moth herbivory and goat herbivory in four palm populations in Mallorca (Balearic Islands) during 2019 and 2020. The effect of herbivory varied widely depending on both the herbivore and the pollinator species. Moth herbivory had a positive effect on pollinator abundance and fruit initiation, whereas goat herbivory had a negative effect on inflorescence production, pollinator abundance and fruit initiation. In addition, both herbivores exerted unexpected nonadditive effects on palm reproduction. Palms attacked by both herbivore species produced many more inflorescences (up to 18-fold) but had a lower fruit initiation success (close to zero) than unattacked palms or those attacked by a single herbivore species. Interestingly, only one of the two main pollinator species (the nitidulid beetle Meligethinus pallidulus) was impacted by herbivory. Our study highlights the need to investigate the possible nonadditive effects of all coexisting herbivores on plant performance, especially when establishing conservation plans and pest control strategies., This research was supported by project MISFAUNATION (PGC2018-094808-B-I00), financed by the Spanish Ministry of Science, Education and Universities. Raquel Muñoz-Gallego was funded by an FPU fellowship from the Spanish Ministry of Universities. Pau E. Serra was funded by the Youth Guarantee Program from the European Commission.

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., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017) and ARC (SFRH/BPD/115781/2016). Funding received from Spanish Ministry of Science, Education and Universities (PGC2018-094808-B-I00) to JMF. FXP was funded by grant (PID2019-104135GB-I00) from the Agencia Estatal de Investigación (AEI/https://doi.org/10.13039/501100011033) of Spain and the European Regional Development Fund (FEDER, UE).

Interspecific ecological interactions are inherently context-dependent. They may vary in both magnitude and sign depending on the biotic and abiotic conditions, depicting a mutualism–antagonism continuum. However, how population abundances and the activity of interacting species modulate these interactions remains underexplored. Here, we chose the interaction between the Mediterranean palm Chamaerops humilis and the feral goat Capra hircus in Mallorca (Balearic Islands, Spain). We selected three study plots with low, intermediate and high intensities of goat activity where we characterized palm distribution, seed rain, seed predation and early palm recruitment during two consecutive years. Since goats can cause both costs (e.g. florivory) and benefits (e.g. seed dispersal) to C. humilis performance, we investigated the following three questions: 1) does the spatial distribution of adult palms vary depending on the intensity of goat activity? 2) Does the intensity of goat activity influence seed rain and its potential spatial association with adult palms? 3) To what extent does the intensity of goat activity determine post-dispersal events such as seed predation and seedling emergence? We found that adult palms showed a more clumped and complex distribution (double-cluster process) in plots with low and intermediate goat activity compared to that with high goat activity (simple-cluster process). In the low goat activity plot, dispersed seeds were spatially aggregated around adult palms, showing twice as much insect-seed predation and nearly three times lower seed germination success than those in the intermediate goat activity plot. Palm seed dispersal and recruitment were almost nil in the high goat activity plot due to heavy consumption of palm inflorescences and developing fruits by goats. Our findings demonstrate how the net outcome of plant–animal interactions can change from mutualism to antagonism, from reproductive service to reproductive collapse, depending on the abundance and the activity of the interacting species., This study is framed within project MISFAUNATION (PGC2018-094808-B-I00) financed by the Spanish Ministry of Science, Education and Universities. RM-G was funded by a FPU fellowship from the Spanish Ministry of Universities., Peer reviewed

Landscape fragmentation and defaunation have major impacts on plant dispersal and dynamics. However, whether the impact of such perturbations on seed dispersal and recruitment change in sign and strength across habitats and spatial scales, and whether they amplify or buffer each other, remains largely unknown. To evaluate, for the first time, the joint impact of fragmentation and defaunation on seed dispersal effectiveness (SDE) across spatial scales (e.g., short- and long-distance seed dispersal), we utilized the long-term field data of a mammal-dispersed tree (Pyrus bourgaeana) in a spatially explicit individual-based model. By means of simulation experiments, we evaluated the effects of different levels of landscape fragmentation and defaunation on SDE and tree recruitment. Our simulation results revealed that the direction and magnitude of the fragmentation effect on SDE depended on its strength (mild, severe) and the spatial scale considered. Severe fragmentation decreased SDE for short- and intermediate-distance seed dispersal. Interestingly, mild and severe fragmentation increased SDE of long-distance seed dispersal, suggesting a positive effect of such perturbations (i.e., an increase in the proportion of successful long-distance dispersal events). Though defaunation had a consistently negative effect on overall SDE, its magnitude was highly species- and spatial–scale-dependent. The impact of defaunation on seed dispersal distance was also species-specific: the proportion of long-distance dispersal increased under total badger (Meles meles) defaunation but decreased under total fox (Vulpes vulpes) defaunation. A pervasive integration of seed dispersal distance into the SDE framework is essential to most comprehensively understand the scale-dependent nature of human activity impacts on plant dynamics. In the long term, strong perturbations (landscape fragmentation and defaunation) could select either for or against long-distance seed dispersal, altering in different ways the ability of plants to cope with climate change., Science, Innovation, and University Ministry of Spain, Grant/Award Number:PGC2018-094808-B-I00, Peer reviewed

This new version of DisPear allows to evaluate changes in seed dispersal effectiveness of Pyrus bourgaeana under scenarios of fragmentation and defaunation., Spanish Ministry of Science, Education and Universities (PGC2018-094808-B-I00), Peer reviewed

Plant–ungulate interactions are critical in shaping the structure of Mediterranean plant communities. Nevertheless, there is a dearth of knowledge on how plant intrinsic and extrinsic factors mediate the sign and strength of plant–ungulate interactions. This is most relevant when addressing natural or assisted restoration of plant communities in human-disturbed areas.
We conducted field-clipping experiments simulating how different intensities of ungulate herbivory may affect the natural regeneration and establishment of the Mediterranean dwarf palm (Chamaerops humilis), a keystone species in Mediterranean ecosystems. We quantified seedling survival and size in two human-disturbed sites (SW Spain) where wild and domestic ungulates exert high herbivory pressure on vegetation.
Severe clipping and seedling aging reduced rates of seedling survival. In contrast, moderate clipping did not affect seedling survival, suggesting a certain degree of C. humilis tolerance to herbivory. Severe clipping reduced seedling height strongly but not seedling diameter, and these effects seem to have decreased seedling survival. Nurse shrubs increased seedling size, which likely improved seedling survival. We also found seedling compensatory growth which varied between study sites.
Field-clipping experiments can help disentangle effects of plant extrinsic and intrinsic factors on the sign and strength of plant–ungulate interactions and their ecological consequences on the dynamics of human-disturbed ecosystems. We call attention to the importance of appropriately managing scenarios of severe herbivory and summer droughts, particularly frequent in Mediterranean ecosystems, as synergic effects of such key drivers can negatively affect the structure and dynamics of plant communities and endanger their conservation., This work was supported by the Portuguese Foundation for Science and Technology to PJG (SFRH/BD/130527/2017); and a grant from the Spanish Ministry of Science and Innovation to JMF (PGC2018-094808-B-I00)., Peer reviewed

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., Spanish Ministry of Science, Innovation and Universities, PGC2018-094808-B-I00, Jose M. Fedriani. Spanish Ministry of Universities, FPU17/05463, Raquel Muñoz Gallego., Peer reviewed

Although it is well recognized that the strength of plant-herbivore interactions can vary with plant sex and the distance and 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 the 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 the 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., PGC2018-094808-B-I00,Spanish Ministry of Science, Innovation and Universities
FPU17/05463, Spanish Ministry of Universities, Peer reviewed

The spatial distribution pattern of plant species is frequently driven by a combination of biotic and abiotic factors that jointly influence the arrival, establishment, and reproduction of plants. Comparing the spatial distribution of a target plant species in different populations represents a robust approach to identify the underlying mechanisms. We mapped all reproductive individuals of the Iberian pear (Pyrus bourgaeana) in five plots (1.39-8.57 km2) differing in the activity of seed dispersers and vertebrate herbivores in southern Iberian Peninsula. We used Thomas point process models to quantify the consistency in the spatial pattern and the level of spatial aggregation of this mammal-dispersed tree among the five populations. We tested two hypotheses: (i) because the clumped defecation behavior of some dispersers can lead to local tree aggregation, and because denser groups of fruiting trees can limit seed dispersal by attracting frugivores to specific sites, we expected a consistent small-scale aggregation pattern across all populations; and (ii) because ungulates reduce recruitment by preying on seeds and seedlings, we hypothesize that ungulate activity will show negative relationships with tree density and level of aggregation. Our spatial analysis revealed consistent and highly aggregated small-scale patterns of all Iberian pear populations, with one critical scale aggregation, a low density of clusters and high variability in the number of trees per cluster. Ungulate activity and the number of trees per cluster showed a marginally significant negative correlation, suggesting that in areas with higher ungulate activity, trees tend to form less dense clusters. Although several of the underlying processes varied greatly among the five study sites, the Iberian pear showed a relatively consistent spatial pattern with just quantitative nuances throughout the entire region. This result has significant implications for the reproductive success of the species, management strategies, and ultimately the long-term persistence of populations., This study was partially supported by Spanish Minister of Science, Innovation and Universities (ref. PID2022-142067NB-I00, PGC2018-094808-B-I00) and Generalitat Valenciana through the Santiago Grisolía Program (ref. CIGRIS/2021/038). PJG was supported by Juan de la Cierva Contracts of Spanish Ministry of Science, Innovation and Universities (JDC2022-048287-I)., Peer reviewed

[EN] Fruit traits play a fundamental role in mediating plant-frugivore interactions, shaping seed dispersal dynamics. Given the complexity of multi-species interactions, these traits and, consequently, plant-frugivore interactions can be indirectly influenced by third-party species, leading to trait-mediated effects. While herbivory is demonstrated to alter some fruit traits through resource allocation shifts or compensatory responses, its impact on plant-frugivore interactions remains largely unexplored.
We examined whether herbivory by the invasive palm borer Paysandisia archon (Lepidoptera, Castniidae) influences fruit attractiveness and alters plant-frugivore interactions in the Mediterranean dwarf palm Chamaerops humilis (Arecaceae). Across four sites in Mallorca (Balearic Islands, W. Mediterranean Sea), we characterized fruit and seed traits and monitored visitation and fruit removal rates by functionally diverse frugivores on intact and infested female palms over two consecutive seasons.
Intensely moth-infested palms (i.e. those with 60% of the stems infested) produced 1.7-fold rounder fruits with 1.3-fold lower pulp content than those of intact palms. Surprisingly, the frequency of fruit removal by legitimate seed dispersers was six times higher and overall fruit removal occurred 10 times faster in infested palms compared to intact ones. However, infested palms also experienced a 2.4 and 3-fold increases in pulp removal by invertebrates and seed predation by scolytine beetles, respectively.
Our findings reveal a trade-off whereby herbivory-driven changes in fruit traits (i.e. rounder fruits with lower pulp content) enhance the attraction of both mutualistic (seed dispersers) and antagonistic (pulp feeders and seed predators) frugivores, but with markedly different strength. These contrasting yet asymmetrical responses emphasize the importance of accounting for frugivore functional diversity, as plant fitness ultimately depends on the balance between mutualistic and antagonistic interactions. Herbivory thus has the potential to reshape plant-frugivore interactions and influence the long-term eco-evolutionary dynamics of plant populations., [ES] Los rasgos de los frutos desempeñan un papel fundamental en la mediación de las interacciones planta-frugívoro, modelando la dinámica de la dispersión de semillas. Dada la complejidad de las interacciones multiespecíficas, estos rasgos y, en consecuencia, las interacciones planta-frugívoro pueden verse indirectamente influenciados por terceras especies, dando lugar a efectos mediados por rasgos. Aunque se ha demostrado que la herbivoría puede alterar algunos rasgos de los frutos mediante la re-asignación de recursos o respuestas compensatorias, su impacto sobre las interacciones planta-frugívoro sigue siendo en gran medida desconocido.
En este trabajo, examinamos si la herbivoría causada por la polilla invasora Paysandisia archon (Lepidoptera, Castniidae) influye en el atractivo de los frutos y modifica las interacciones planta-frugívoro en el palmito mediterráneo Chamaerops humilis (Arecaceae). En cuatro localidades de Mallorca (Islas Baleares, Mediterráneo occidental), caracterizamos los rasgos de frutos y semillas, y monitorizamos las tasas de visita y de consumo de frutos por frugívoros funcionalmente diversos en palmeras hembras intactas e infestadas durante dos temporadas consecutivas.
Las palmeras intensamente infestadas (es decir, con el 60% de los troncos infestados) produjeron frutos 1.7 veces más redondeados y con un contenido de pulpa 1.3 veces menor que las palmeras intactas. Sorprendentemente, la frecuencia de consumo de frutos por los dispersores de semillas legítimos fue seis veces mayor y el consumo total de frutos se produjo diez veces más rápido en las palmeras infestadas comparado con las intactas. Sin embargo, estas también experimentaron incrementos de 2.4 y 3 veces en el consumo de pulpa por invertebrados y la depredación de semillas por escarabajos escolítinos, respectivamente.
Nuestros resultados revelan una compensación (trade-off) por la cual los cambios en los frutos (es decir, frutos más redondos y con menos pulpa) inducidos por la herbivoría aumentan la atracción tanto de frugívoros mutualistas (dispersores de semillas) como antagonistas (consumidores de pulpa y depredadores de semillas), aunque con diferente intensidad. Estas respuestas opuestas pero asimétricas resaltan la importancia de considerar la diversidad funcional de los frugívoros, dado que el éxito reproductivo de las plantas depende en última instancia del balance entre interacciones mutualistas y antagonistas. La herbivoría, por tanto, tiene el potencial de reconfigurar las interacciones planta-frugívoro e influir en las dinámicas eco-evolutivas de las poblaciones vegetales a largo plazo., The Spanish Ministry of Science and Innovation financed this study through the MISFAUNATION project (PGC2018-094808-B-I00). Raquel Muñoz-Gallego was funded by a FPU fellowship (FPU17/05463) from the Spanish Ministry of Universities. This study was carried out within the framework of the activities of the Spanish Government through the ‘Maria de Maeztu Centre of Excellence’ accreditation to IMEDEA (CSIC-UIB) (CEX2021-001198)., Peer reviewed

Most research on endozoochorous seed dispersal by ungulates emphasizes long-distance dispersal and its dependence on ungulate species and habitat heterogeneity. In contrast, the role of ungulate community composition in shaping local (around one hundred hectares or less) spatial patterns in the distribution of plant seeds remains largely unexplored. To address such key question, we quantified ungulate seed dispersal at two grasslands in the Doñana National Park (southwestern Spain). One grassland (Martinazo) was used by a mixed community of four ungulate species (deer, wild boar, cattle, and horses), while the other one (Matasgordas) was used almost exclusively by deer. At each site, a plot was established, and ungulate fecal samples were systematically collected and georeferenced from early spring to mid-summer. The seeds contained in the samples were extracted and identified in the laboratory. Our results revealed clear differences between sites in the total number of dispersed herbaceous seeds (1,302 and 606 seeds in Martinazo and Matasgordas respectively) and the most frequently dispersed plant families. Within the four-species community, cattle and deer differed most in the taxonomic composition of the seeds they dispersed, suggesting that herbivore-specific seed selection and dispersal act as key drivers of grassland structure at fine spatial scales, mirroring dynamics typically observed at broader scales. Moreover, only in the four-ungulate community did we detect significant spatial patterns in seed dispersal. These included positive effects of short-distance feces aggregation, spatial covariance in seed content among nearby feces, and a strong correlation between seed content and local feces density. Seed families predominantly dispersed by cattle also exhibited significant spatial structuring. These findings have important implications for biodiversity management in semi-natural and protected ecosystems, as they demonstrate that the management of ungulate species (both wild and domestic) also influences plant communities. This highlights the need to consider the functional differences among herbivores in ecosystem conservation and restoration strategies., This work was funded by the Science, Innovation, and University Ministry of Spain. Ref. PGC2018-094808-B-I00., Peer reviewed

Datasets pertain to a study on the effect of the ungulate community composition on short distance seeds dispersal and the potential generation of spatial pattern of plant species distribution. The study was conducted at two grassland sites (Martinazo and Matasgordas) in the Doñana National Park (SW Spain). The sites were used by: i) a four ungulate species comunity (i.e., deer, wild boar, cattle and horse) and ii) a community highly dominated by deer. Both sites differed in the total number of herbaceous seeds dispersed and in the plant families most abundantly dispersed. In the four-species ungulate community, cow and deer were the most dissimilar pair in taxonomic composition of dispersed seeds. In addition, in this community, but not in the deer dominated one, we found significant spatial patterns of total seed dispersal (significant positive effect of short-distance feces aggregation, spatial covariance in seed content of two nearby feces, and significant correlation between seed content and density of nearby feces) as well as significant spatial pattern of dispersal of the seeds families preferentially associated with cow. These findings highlight a notable effect of the composition of disperser community on seeds dispersal and spatial structure at the short spatial scale of this study, Dataset consists of ten datafiles that include: i) information on seed content and taxonomic composition (families) in fecal samples collected in the two study sites (Matasgordas and Martinazo) and where the ungulate comunities are composed by a single disperser (deer in Matasgordas) or four dispersers (dear, cow, horse, wild board in Martinazo) and ii) information on spatial coordinates of fecal samples collected in the two sites (Martinazo and Matasgordas) as well as the seed content per sample (total seed content and seed content of the three most frequent plant families dispersed at each site)., Ministerio de Ciencia, Innovación y Universidades (MICIU). España, PGC2018-094808-B-I00, Peer reviewed

Extreme weather events are becoming more frequent under climate change. In many plant–frugivore networks, ground-dwelling vertebrates are the primary consumers of ripe fruits after they fall to the ground. Sudden fruit drop pulses triggered by extreme events (e.g. storms and cyclones) may drastically alter the timing and magnitude of fruit availability, potentially redistributing interaction opportunities across functionally diverse frugivores (i.e. seed dispersers and predators). However, the potential impacts of extreme events on plant–seed disperser mutualisms remain largely unexplored.
In this study, we illustrate how concurrent extreme precipitation and wind events alter fruit drop dynamics and plant–frugivore interactions by combining continuous climatic data, fruit drop monitoring, controlled fruit-offering experiments and a mechanistic simulation model. To this end, we focus on the Iberian pear (Pyrus bourgaeana) and its functionally diverse frugivores in an ungulate-dominated landscape in south-western Mediterranean Spain.
Monitoring data showed that extreme events significantly increased the probability of pulsed fruit drop, generating abrupt pulses of fruit availability. Field experiments showed that these pulses enhanced short-term fruit removal by both seed dispersers (mammalian carnivores) and seed predators (ungulates), but predators responded more strongly. Simulations further revealed that as extreme events became more frequent, intense and prolonged, and thus, the number of pulsed fruit drop days increased, the imbalance in fruit removal was amplified as the fruiting season progressed: while each pulsed fruit drop day raised disperser removal, it simultaneously increased predation, resulting in an overall ~4% reduction in seed dispersal throughout the fruiting season.
Synthesis: These results underscore the profound, yet underappreciated, role of extreme weather events in reshaping the ecological dynamics of fruit drop and the balance between seed dispersal and predation in plant–frugivore networks. However, the direction and magnitude of such effects are likely context-dependent. As such, the consequences of increasing extreme events for plant reproductive success will depend not only on changes in fruit supply but also on how species composition and functional structure mediate the redistribution of interaction opportunities., This study was partially supported by the Spanish Minister of Science, Innovation and Universities (ref. PID2022-142067NB-I00, PGC2018-094808-B-I00). Brayan Morera is funded through the Generalitat Valenciana through the Santiago Grisolía Program (ref. CIGRIS/2021/038). Pedro J. Garrote is funded through the Juan de la Cierva Contracts of the Spanish Ministry of Science, Innovation and Universities (ref. JDC2022-048287-I)., Peer reviewed

The spatial distribution pattern of plant species is frequently driven by a combination of biotic and abiotic factors that jointly influence the arrival, establishment, and reproduction of plants. Comparing the spatial distribution of a target plant species in different populations represents a robust approach to identify the underlying mechanisms. We mapped all reproductive individuals of the Iberian pear (Pyrus bourgaeana) in five plots (1.39–8.57 km2) differing in the activity of seed dispersers and vertebrate herbivores in southern Iberian Peninsula. We used Thomas point process models to quantify the consistency in the spatial pattern and the level of spatial aggregation of this mammal-dispersed tree among the five populations. We tested two hypotheses: (i) because the clumped defecation behavior of some dispersers can lead to local tree aggregation, and because denser groups of fruiting trees can limit seed dispersal by attracting frugivores to specific sites, we expected a consistent small-scale aggregation pattern across all populations; and (ii) because ungulates reduce recruitment by preying on seeds and seedlings, we hypothesize that ungulate activity will show negative relationships with tree density and level of aggregation. Our spatial analysis revealed consistent and highly aggregated small-scale patterns of all Iberian pear populations, with one critical scale aggregation, a low density of clusters and high variability in the number of trees per cluster. Ungulate activity and the number of trees per cluster showed a marginally significant negative correlation, suggesting that in areas with higher ungulate activity, trees tend to form less dense clusters. Although several of the underlying processes varied greatly among the five study sites, the Iberian pear showed a relatively consistent spatial pattern with just quantitative nuances throughout the entire region. This result has significant implications for the reproductive success of the species, management strategies, and ultimately the long-term persistence of populations.

Landscape fragmentation and defaunation have major impacts on plant dispersal and dynamics. However, whether the impact of such perturbations on seed dispersal and recruitment change in sign and strength across habitats and spatial scales, and whether they amplify or buffer each other, remains largely unknown. To evaluate, for the first time, the joint impact of fragmentation and defaunation on seed dispersal effectiveness (SDE) across spatial scales (e.g., short- and long-distance seed dispersal), we utilized the long-term field data of a mammal-dispersed tree (Pyrus bourgaeana) in a spatially explicit individual-based model. By means of simulation experiments, we evaluated the effects of different levels of landscape fragmentation and defaunation on SDE and tree recruitment. Our simulation results revealed that the direction and magnitude of the fragmentation effect on SDE depended on its strength (mild, severe) and the spatial scale considered. Severe fragmentation decreased SDE for short- and intermediate-distance seed dispersal. Interestingly, mild and severe fragmentation increased SDE of long-distance seed dispersal, suggesting a positive effect of such perturbations (i.e., an increase in the proportion of successful long-distance dispersal events). Though defaunation had a consistently negative effect on overall SDE, its magnitude was highly species- and spatial–scale-dependent. The impact of defaunation on seed dispersal distance was also species-specific: the proportion of long-distance dispersal increased under total badger (Meles meles) defaunation but decreased under total fox (Vulpes vulpes) defaunation. A pervasive integration of seed dispersal distance into the SDE framework is essential to most comprehensively understand the scale-dependent nature of human activity impacts on plant dynamics. In the long term, strong perturbations (landscape fragmentation and defaunation) could select either for or against long-distance seed dispersal, altering in different ways the ability of plants to cope with climate change.