[Description of methods used for collection/generation of data] Over a span of 5 years (four complete fruiting seasons; 2018-11-21 to 2023-04-17), we monitored 339 plant individuals belonging to 13 distinct fleshy-fruited plant species with camera traps. Uur approach involved the strategic placement of camera traps in front of specific plant individuals, referred to as focal plants. These cameras were set in video mode, providing valuable insights for identifying animal species, but also annotating specific foraging behaviors and estimating fruit consumption rates for certain species, as well as fruit-picking and fruit handling behaviors. We deployed a total of 70 cameras, including 50 Browning Dark Ops®, 10 Bushnell Trophy® cam Aggressor, and 10 GoPro® cameras. These cameras were strategically positioned towards different plant individuals bearing fleshy fruits. Sensor-triggered camera traps were used for 12 plant species, while continuous recording with GoPro® cameras was used for sampling Pistacia lentiscus. Regular checks of the camera traps were conducted at regular intervals, either weekly or biweekly, depending on the plant species and the specific period of the year, [Methods for processing the data] We developed an image processing workflow specifically designed for integrating video recordings and creating extensive and accurate databases of frugivore interactions using camera traps. We leverage Megadetector and Time-lapse, an AI model for eliminating blank images and enhanced visualisation software respectively., The goal of this repository is to contain and facilitate access to the data associated to the datapaper FRUGIVORY CAMTRAPS: A dataset of plant-animal interactions recorded with camera traps, Ecological interactions are a key component of biodiversity, essential for understanding ecosystem services and functioning. Recording and quantifying ecological interactions is challenging, frequently requiring complex logistics and substantial effort in the field. Camera traps are routinely used in ecology for various applications and have proven to be an excellent method for passive and non-invasive sampling of plant-animal interactions. We implemented a standardized camera trap protocol to monitor frugivory interactions between fleshy-fruited plant species and vertebrate frugivores in Doñana National Park, SW Spain. The central objective was to inventory the diversity of plant-animal ecological interactions providing seed dispersal services. We recorded pairwise interactions between plant individuals and vertebrate frugivore species from 2018 to 2023, reporting both qualitative information (presence-absence of the interaction) and quantitative data (frequency of visits). Each record in the dataset contains information of a visit by an individual animal to an individual plant, resulting in any form of fleshy-fruit use and provides information on visitation phenology, visit length and feeding behavior. The dataset presented here includes 10,659 frugivory interactions for 59 vertebrate species (46 birds, 13 mammals) recorded on 339 plant individuals from 13 different plant species which dominate the fleshy-fruited plant assemblage in the Doñana National Park. The most recorded animal species consuming fruits and playing a legitimate seed dispersal role was Curruca melanocephala (1678 records) among birds and Vulpes vulpes among mammals (751 records). On the other hand, Cervus elaphus was the most recorded mammal species (1508 records), but it is a fruit consumer with a marginal role as legitimate seed disperser. Avian frugivores, particularly those from the Sylviidae and Turdidae families, are widespread in the region, and play a crucial role in maintaining the dispersal service for the fleshy-fruited plant populations in the area. The dataset offers highly versatile quantitative information that can be used to investigate frugivory from the highest resolution scale, the interaction event between pairs of individuals. In addition, information can be extracted about the timing of interactions of animals and plants (their phenological couplings), activity periods of the animals, behavior during the events and preferences for individual plants within populations., This study was funded by MICINN through the European Regional Development Fund [SUMHAL, LIFEWATCH-2019-09-CSIC-4, POPE 2014-2020], with additional funding (PJ) from grant PID2022-136812NB-I00 by MCIN/AEI/10.13039/501100011033. IM was supported by the grant PID2020-115129RJ-I00 by MCIN/AEI/10.13039/501100011033., Peer reviewed

Memoria presentada por Elena Quintero Borrero para optar al Título de Doctora por la Universidad de Sevilla., [EN] Studying interactions among species and their multiple forms and outcomes has become a central question of ecological research. Mutualisms have changed the way ecologists understand the functioning of ecosystems. Generalised mutualistic networks, especially among free-living species, have been observed to include highly heterogeneous assemblages, in which most species establish few interactions while a minority of species establish a wide range of interactions. These highly connected species are referred to as super-generalists. They are characterised for connecting otherwise isolated groups of interacting species or modules and for providing stronger cohesion to ecological networks. Their characteristics make them fundamental for local biodiversity and the structure and stability of interactions.

In this PhD thesis we aim to better comprehend the super-generalist strategy and the mutualistic interactions at the individual level. For this purpose, we use as a model organism Pistacia lentiscus L. (Anacardiaceae), a woody shrub species abundant in the Mediterranean Basin, together with the coterie of frugivorous animals that consume its fruits and disperse its seeds. In Chapter 1 we review sampling methods used to record frugivory interactions, assessing their strengths, caveats, and convenience in different contexts. We discuss different approaches for combining data gathered using different methodologies. In Chapter 2, we explore the effectiveness of the frugivory/seed-dispersal mutualism between individual plants of P. lentiscus and its avian frugivore assemblage, assessing the level of reciprocity in reward exchange and dependence asymmetry among partners. In Chapter 3 we investigate the delayed outcomes of animals’ fruit consumption in early seedling recruitment of P. lentiscus plants. Lastly, in Chapter 4 we analyse individual-based networks of different plant species and regions of the world, we compare their topology and structure with that of species-based networks, and explore the specialisation and interaction profile of individual plants within populations.

We highlight the significant advancements brought by emerging methods like camera traps and molecular tools, which allow recording interactions across large spatiotemporal scales and even tracking the movement of individual seeds by animals. We also show how data integration improves network completeness and representativity, and compare different data merging approaches for adjacency matrices. We document how the effectiveness of interactions established between P. lentiscus individual plants and their frugivores is mainly determined by fruit consumption frequency (the component with highest variation), leading to highly reciprocal exchange of services but still highly asymmetric dependence among partners. Despite being mostly determined by the number of fruits consumed, the dispersal service provided by birds is decoupled from microhabitat suitability: frugivores deposit a smaller number of seeds in the most suitable microhabitats. These results underscore the role of different frugivores in spatial recruitment of heterogeneous landscapes. Finally, we demonstrate that the structure of individualbased networks is very similar to that of species-based networks. Plant individuals presented similar interaction profiles irrespective of the species or regions their population belonged to. Within populations, plants present low to medium levels of specialisation, and few frugivore species contribute most interactions in all studied networks.

Overall, these results suggest that super-generalist species may evolve when they combine sets of traits that make them accessible and attractive to a diversified assemblage of frugivores. Numerical effects, such as abundant fruit crops, facilitate plenty of frugivory interactions, characterised in most cases by high reciprocity. In exchange for a fair nutritious reward, plants secure their recruitment thanks to a vast amount of seeds dispersed. Finally, our results indicate that the variation encountered in the way plant individuals structure their interactions is highly consistent across populations of generalised mutualisms worldwide.

Collectively, the four chapters in this thesis contribute to better understanding the origin and maintenance of super-generalist species within complex ecological networks, by focusing on the biological scale at which interactions actually occur in nature, that is, the individual scale., [ES] Estudiar las interacciones entre especies junto a sus múltiples formas y resultados se ha convertido en una cuestión central de la investigación en ecología. Los mutualismos han cambiado la forma en que los ecólogos entienden el funcionamiento de los ecosistemas. Se ha observado que las redes mutualistas generalizadas, especialmente entre especies de vida libre, incluyen conjuntos muy heterogéneos, en los que la mayoría de las especies establecen pocas interacciones, mientras que una minoría de especies establece una amplia variedad de interacciones. Estas especies altamente conectadas se denominan super-generalistas. Se caracterizan por conectar grupos de especies o módulos que de otro modo estarían aislados, y por proporcionar una mayor cohesión a las redes ecológicas. Sus características las convierten en especies fundamentales para la biodiversidad local así como para la estructura y estabilidad de las interacciones.

En esta tesis doctoral pretendemos comprender mejor la estrategia supergeneralista y las interacciones mutualistas a nivel individual. Para ello, utilizamos como organismo modelo Pistacia lentiscus L. (Anacardiaceae), una especie arbustiva leñosa abundantemente distribuida por la cuenca mediterránea, junto a los animales frugívoros que consumen sus frutos y dispersan sus semillas. En el Capítulo 1 se revisan los métodos de muestreo utilizados para registrar las interacciones de frugivoría, evaluando sus ventajas, inconvenientes y conveniencia en diferentes contextos; y se discuten diferentes enfoques para combinar los datos recogidos utilizando diferentes metodologías. En el Capítulo 2, exploramos la eficacia en el mutualismo frugivoría/semilla-dispersión entre plantas individuales de P. lentiscus y las aves frugívoras que consumen sus frutos, evaluando el nivel de reciprocidad en el intercambio de recompensas y la asimetría de dependencia entre los socios. En el Capítulo 3, investigamos los resultados aplazados del consumo de frutos por animales en plantas de P. lentiscus para el reclutamiento temprano de plántulas. Por último, en el Capítulo 4 analizamos redes basadas en individuos para diferentes especies de plantas y regiones del mundo, comparando su topología y estructura con la de redes basadas en especies, y exploramos la especialización y el perfil de interacción de plantas individuales dentro de sus poblaciones.

Destacamos los importantes avances que han supuesto métodos emergentes como las cámaras trampa y técnicas moleculares, que permiten registrar interacciones a una amplia escala espacio-temporal e incluso rastrear el movimiento de semillas individuales por parte de animales. También mostramos cómo la integración de datos mejora la completitud y representatividad de las redes ecológicas, y comparamos distintos métodos de combinación de datos para matrices de adyacencia. A continuación, documentamos cómo la eficacia de las interacciones establecidas entre plantas individuales y sus frugívoros viene determinada principalmente por la frecuencia de consumo de frutos (el componente con mayor variación), lo que conduce a un intercambio de servicios altamente recíproco, pero manteniendo una dependencia altamente asimétrica entre los socios. A pesar de estar determinado principalmente por el número de frutos consumidos, el servicio de dispersión proporcionado por las aves está desvinculado de la idoneidad del microhábitat: los frugívoros depositan un menor número de semillas en los microhábitats más adecuados. Estos resultados resaltan el papel que los distintos frugívoros tienen en el reclutamiento en paisajes heterogéneos. Por último, demostramos que la estructura de las redes basadas en individuos es muy similar a la de las redes basadas en especies. Las plantas individuales presentaron perfiles de interacción similares independientemente de la especie o región a la que perteneciera su población. Dentro de las poblaciones, las plantas presentan niveles de especialización medio-bajos, y unas pocas especies de frugívoros contribuyen a la mayoría de las interacciones en todos las redes estudiadas.

Estos resultados sugieren que las especies super-generalistas pueden evolucionar cuando combinan conjuntos de rasgos que las hacen accesibles y atractivas para un conjunto diversificado de frugívoros. Efectos numéricos como la alta abundancia de frutos facilitan gran cantidad interacciones frugívoras, caracterizadas en la mayoría de los casos por una elevada reciprocidad. A cambio de la recompensa nutritiva que provee la pulpa de los frutos, las plantas aseguran su reclutamiento gracias a una gran cantidad de semillas dispersadas. Por último, nuestros resultados indican que la variación encontrada en la forma en que las plantas individuales estructuran sus interacciones es muy consistente en las poblaciones de mutualismos generalizados de todo el mundo.

En conjunto, los cuatro capítulos de esta tesis contribuyen a comprender mejor el origen y la persistencia de especies super-generalistas dentro de redes ecológicas complejas, centrándose en la escala biológica a la que se producen realmente las interacciones en la naturaleza, es decir, la escala individual., This thesis received financial support from “la Caixa” Foundation fellowship for Doctoral studies at Spanish Universities or Research Centres (ID 100010434) with code LCF/BQ/DE18/11670007, as well as by the projects CGL2017-82847-P and PID2022-136812NB-I00 from Agencia Estatal de Investigación, Ministerio de Ciencia e Innovación and project PAIDI_2020-P20_00736 from Consejería de Economía, Conocimiento, Empresas y Universidad, Junta de Andalucía. In addition, this thesis received support by a Fulbright fellowship for pre-doctoral research abroad, co-funded by Junta de Andalucia and by project LIFEWATCH-2019-09-CSIC-13 funded by European Fund for Regional Development (FEDER-EU)., Peer reviewed

The consequences of plant–animal interactions often transcend the mere encounter stage, as those encounters are followed by a chain of subsequent stages on the plant's reproductive cycle that ultimately determine fitness. Yet, the dissemination and recruitment stages of animal-mediated seed dispersal are seldom analysed jointly, hindering a full understanding of the ecology of seed dispersal. We analyse the dispersal and recruitment of a fleshy-fruited plant (Pistacia lentiscus), from fruit production to seedling survival up to their second year. We link early reproductive investment of individual plants to seedling recruitment and explore the role played by seed viability, the coterie of frugivores and microhabitat seed deposition. The proportion of viable seeds was generally low (mean = 34%) but highly variable among individual plants (range: 0%–95%). Seed viability did not seem to have a direct effect on individual plant's recruitment. We recorded 28 bird species feeding on P. lentiscus fruits or seeds. Their contribution to plant recruitment was mainly determined by their intensity of fruit consumption and probability to disperse viable seeds. Most frugivores presented non-random microhabitat preferences, delivering uneven seed contributions to different sites. Post-dispersal seed predation by rodents was the most limiting phase in P. lentiscus recruitment. Yet, microhabitats showing the lowest predation rates received the lowest seed rain. Hence, we found a decoupling of the dissemination and recruitment stages: most seeds do not arrive at the most suitable microhabitats. We estimate P. lentiscus plants need to produce c. 5 × 105 fruits to recruit a single seedling that survives to its second summer in our study site. Its success as a prevalent species in Mediterranean lowland landscapes relies on its high fecundity and thorough fruit removal and dispersal by a diversified frugivore assemblage, which compensates for the high seed unviability characteristic of this genus. Synthesis: Measuring the delayed, post-dispersal outcomes of animal frugivory interactions may overturn inferences based on consumption observations only. Seed rain patterns are often decoupled from microhabitats' suitability for seedling recruitment. Hence, more integrative studies that encompass the entire plant reproductive cycle (from fruit production to seedling recruitment) are needed to fully understand frugivores' lasting contribution to plant regeneration in natural populations., EQ received support from “la Caixa” Foundation (ID 100010434), under agreement LCF/BQ/DE18/11670007 and from a Fulbright fellowship co-funded by Junta de Andalucía. E.Q., P.J. and F.R.S. received support from LifeWatch ERIC-SUMHAL project (LIFEWATCH-2019-09-CSIC-13), with FEDER-EU funding, and grant PID2022-136812NB-I00 funded by MCIN/AEI/10.13039/501100011033 and the European Union NextGenerationEU/PRTR. F.R.S. and P.J. were additionally supported by VI Plan Propio de Investigación, Universidad de Sevilla (VI PPIT-US), FEDER 2014–2020 and Consejería de Economía, Conocimiento, Empresas y Universidad, Junta de Andalucía (grant US-1381388, Universidad de Sevilla)., Peer reviewed

Indirect interactions among species within ecological communities govern ecological and evolutionary processes as much as, or even more, than direct effects. In insect-pollinated plant communities, indirect interactions between plants can be mediated by shared pollinators, and may influence plant fitness, population growth and community structure. As individuals are the entities actually interacting in nature, rather than species, downscaling a hierarchical level (i.e. from species to individuals) is essential to understand the underlying processes promoting these indirect interactions. We combined empirical data on plant–pollinator interactions collected in Mediterranean shrublands with a novel modelling framework to assess how the patterns of heterospecific and conspecific pollinator sharing between plant individuals are generated and their fitness implications. We found that the effects of flowering synchrony on pollinator sharing among conspecific and heterospecific plant individuals outperformed those of spatial distance. Our results revealed that plant individuals that shared more pollinators with conspecifics were also involved in a higher pollinator sharing with heterospecifics. For most plant species, the sharing of pollinator species between heterospecific plant individuals produced positive mean fitness outcomes, as long as plants did not share many pollinator interactions, which had negative effects on their fitness. At the level of plant individuals, we found that specific combinations of conspecific and heterospecific pollinator sharing lead to distinct reproductive outcomes that placed each individual along a competition–facilitation continuum. Interestingly, most plant species included a higher proportion of individuals likely experiencing competition compared to those potentially involved in facilitation processes. Our contribution provides novel insights into the factors responsible for local-scale indirect interactions within communities and their individual-level functional consequences. Such intricate patterns of indirect interactions have far-reaching implications, as the transitions of plant individuals along competitive or facilitative processes mediated by shared mutualists may contribute significantly to the functioning and coexistence of ecological communities., BA-C received funding from the Ministry of Universities of the Spanish Government (Ref. FPU19_02552). BA-C and PJ were supported by grants CGL2017-82847-P and PID2022-136812NB-I00, funded by MCIN/AEI/10.13039/501100011033 and by the “European Union NextGenerationEU/PRTR”, grant P20_00736 from Junta de Andalucía, and a LifeWatch ERIC-SUMHAL project (LIFEWATCH-2019-09-CSIC-13), with FEDER-EU funding., Peer reviewed

Ecologists have long recognized that seed dispersal mutualisms trigger natural regeneration and range expansion of animal-dispersed trees. Yet we lack empirical studies addressing whether frugivore activity influences founder effects, which reduce genetic diversity at the colonization front of expanding populations. Here, we evaluate the contribution, from both demographic and genetic perspectives, of animal frugivores dispersing seeds across an expansion gradient. We used DNA barcoding for frugivores identification and highly polymorphic genetic markers (SSRs) for maternal analysis of juniper seeds to investigate how (1) stand maturity, (2) microhabitat types and (3) foraging patterns shape the distribution of the maternal progenies along this gradient. We expect both reduced seed rain density and low numbers of source trees contributing to the seed rain at the colonization front, with limited availability of local fruiting trees. We also anticipated that large-sized frugivore species would promote maternally rich seed rain due to their ability to mix seed progenies during digestive processing and move further distances across the landscape. Contrary to our expectations, we found that all identified frugivores produced dense and genetically diverse seed rains across the expansion gradient, even at the colonization front, characterized by scarce fruiting trees. Synthesis: Our findings shed light on the fundamental and applied implications of plant–frugivore interactions in shaping highly diverse second-growth forests. These results emphasize the necessity of preserving plant–animal mutualistic interactions to ensure the persistence and expansion of natural tree populations, particularly in formerly fragmented landscapes., Our work was funded by grants CGL2017-82847-P and PID2022-136812NB-I00, Spanish Ministry of Science and Innovation and grant SUMHAL, LIFEWATCH-2019-09-CSIC-13, POPE 2014–2020, MICINN through the European Regional Development Fund, as well as funding from the VI Plan Propio de Investigación of the Universidad de Sevilla (VI PPIT-US) (PJ). JI was supported by an FPI grant PRE2018-085916 from the Spanish Ministry of Science and Innovation. MJF was supported by Investigadores por México-CONAHCYT (grant number: 73215). CG was supported by the Research Application Support Programme (RHUL) and the Royal Society (Research Application Support Programme) RaSP., Peer reviewed

Ecological interactions are a key component of biodiversity, essential for understanding ecosystem services and functioning. Recording and quantifying ecological interactions is challenging, frequently requiring complex logistics and substantial effort in the field. Camera traps are routinely used in ecology for various applications, and have proven to be an excellent method for passive and non-invasive sampling of plant-animal interactions. We implemented a standardized camera trap protocol to document vertebrate frugivores-fleshy fruited plants interactions in Doñana National Park, SW Spain, with the central objective of inventorying the diversity of plant-animal ecological interactions providing seed dispersal services. From 2018 to 2023 we recorded pairwise interactions from which we obtained qualitative (presence-absence) and quantitative (frequency of visits) information. Each record in the dataset contains information of a visit by an individual animal to an individual plant, resulting in any form of fleshy-fruit use and provides information on visitation phenology, visit length, and feeding behavior. The dataset presented here includes 10,659 frugivory interaction events for 59 vertebrate species (46 birds, 13 mammals) recorded on 339 plant individuals from 13 different plant species which dominate the fleshy-fruited plant assemblage in the Doñana National Park. The most recorded animal species consuming fruits and playing a legitimate seed dispersal role was Curruca melanocephala (1678 records) among birds and Vulpes vulpes among mammals (751 records). Cervus elaphus, a fruit consumer with a marginal role as legitimate seed disperser, was the most recorded mammal species (1508 records). Avian frugivores, particularly those from the Sylviidae and Turdidae families, are widespread in the region and play a crucial role in maintaining the dispersal service for the fleshy-fruited plant populations in the area. The dataset offers highly versatile quantitative information that can be used to investigate frugivory from the highest resolution scale, the interaction event between pairs of individuals. In addition, other information that can be extracted includes the timing of interactions of animals and plants (their phenological couplings), activity periods of the animals, behavior during the events and preferences for individual plants within populations. There are no copyright restrictions on the data. When using the data from this data paper in publications, we kindly request that you cite the paper accordingly. Additionally, we encourage researchers and educators to inform us about how they are using this data, as we value feedback and would like to be aware of its various applications., Ministerio de Ciencia e Innovación, Grant/Award Numbers: MCIN/AEI/10.13039/501100011033-PID2020-115129RJ-I0, MCIN/AEI/10.13039/501100011033-PID2022-136812NB-I0, LIFEWATCH-2019-09-CSIC-4, POPE 2014-2020, Peer reviewed

Species-level networks emerge as the combination of interactions spanning multiple individuals, and their study has received considerable attention over the past 30 y. However, less is known about the structure of interaction configurations within species, even though individuals are the actual interacting units in nature. We compiled 46 empirical, individual-based, interaction networks on plant-animal seed dispersal mutualisms, comprising 1,037 plant individuals across 29 species from various regions. We compared the structure of individual-based networks to that of species-based networks and, by extending the niche concept to interaction assemblages, we explored individual plant specialization. Using a Bayesian framework to account for uncertainty derived from sampling, we examined how plant individuals "explore" the interaction niche of their populations. Both individual-based and species-based networks exhibited high variability in network properties, lacking remarkable structural and topological differences between them. Within populations, frugivores' interaction allocation among plant individuals was highly heterogeneous, with one to three frugivore species dominating interactions. Regardless of species or bioregion, plant individuals displayed a variety of interaction profiles across populations, with a consistently-small percentage of individuals playing a central role and exhibiting high diversity in their interaction assemblage. Plant populations showed variable mid to low levels of niche specialization; and individuals' interaction niche "breadth" accounted for 70% of the population interaction diversity, on average. Our results highlight how downscaling from species to individual-based networks helps understanding the structuring of interactions within ecological communities and provide an empirical basis for the extension of niche theory to complex mutualistic networks., Funding from LIFEWATCH-2019-09-CSIC-13 project and PID2022-136812NB-I00, Spanish Ministry of Science and Innovation, as well as funding from the VI Plan Propio de Investigación of the Universidad de Sevilla (VI PPIT-US) supported this research (P.J.). E.Q. was supported by “la Caixa” Foundation fellowship LCF/BQ/DE18/1167000. B.A.-C. received funding from the Ministry of Universities of the Spanish Government (Ref. FPU19_02552). J.I. was supported by a Formación de Personal Investigador grant PRE2018-085916 from the Spanish Ministry of Science and Innovation. F.R.-S. was supported by VI PPIT-US and grants US-1381388 from Universidad de Sevilla/Junta de Andalucía/FEDER-UE and CNS2022-135839 funded by MICIU/AEI/10.13039/501100011033 and by European Union NextGenerationEU/PRTR., Peer reviewed

Mutualistic interactions among organisms are fundamental to the origin and maintenance of biodiversity. Yet, the study of community dynamics often relies on values averaged at the species level, ignoring how intraspecific variation can affect those dynamics. We developed a theoretical approach to evaluate the extent to which variation within populations, in terms of interactions, can influence structural stability, a robust measure of species' likelihood of persistence in mutualistic systems. Next, we examine how intraspecific variation in mutualistic interactions affects species' persistence theoretically in a simplified community, which provides a solid foundation for contextualizing empirical results. This theoretical exploration revealed that differences in the benefits received by different individual types by mutualistic partners, as driven by the way interactions are distributed among those types due to individual specialization, strongly influence species persistence. Building on these insights, we move beyond the theoretical framework and work through an empirical case study involving three co-occurring plant species. Drawing from detailed field data on plant–pollinator interactions and plant fitness, we quantify intraspecific variation in the mutualistic benefits received by plants and incorporate this variation into estimations of structural stability. Through explicit consideration of this facet of intraspecific variation, we found that, for all three focal plant species, populations composed of individuals specialized in pollinator use promote the persistence of the plant species they belong to and their associated pollinator community, only in the absence of heterospecific plant competitors. However, more importantly, these positive effects do not hold when plant species compete with a broader, diverse plant community. In this case, two of the focal plant populations are more vulnerable when they comprise more specialized individuals and therefore are less likely to persist. By integrating the proposed theoretical approach with empirical data, this study highlights the importance of individual variation in promoting species persistence in mutualistic systems. In doing so, it not only advances our understanding of basic mechanisms that foster biodiversity maintenance but also provides practical insights for biodiversity conservation in the face of changing environmental conditions., Blanca Arroyo-Correa received funding from the Ministry of Universities of the Spanish Government (FPU19/02552 and EST23/00036) and a project grant from the Spanish Association of Terrestrial Ecology (AEET). Blanca Arroyo-Correa and Pedro Jordano were supported by grants CGL2017-82847-P and PID2022-136812NB-I00, funded by MCIN/AEI/10.13039/501100011033 and by the “European Union NextGenerationEU/PRTR”, grant P20-00736 from Junta de Andalucía, and a LifeWatch ERIC-SUMHAL project (LIFEWATCH-2019-09-CSIC-13), with FEDER-EU funding. E. Fernando Cagua acknowledges the support from the University of Canterbury Doctoral Scholarship, the Meadow Mushrooms Ltd.: University of Canterbury Postgraduate Scholarship, and a Education New Zealand: New Zealand Doctoral Research Scholarship. Daniel B. Stouffer acknowledges the support of a Rutherford Discovery Fellowship (RDF-13-UOC-003), administered by the Royal Society of New Zealand Te Apārangi., Peer reviewed

Recording and quantifying ecological interactions is vital for understanding biodiversity, ecosystem stability, and resilience. Camera traps have become a key tool for documenting plant–animal interactions, especially when combined with computer vision (CV) technology to handle large datasets. However, creating comprehensive ecological interaction databases remains challenging due to labor-intensive processes and a lack of standardization. While CV aids in data processing, it has limitations, including information loss, which can impact subsequent analyses. This study presents a detailed methodology to streamline the creation of robust ecological interaction databases using CV-enhanced tools. It highlights potential pitfalls in applying CV models across different contexts, particularly for specific plant and animal species. The approach aligns with existing camera trap standards and incorporates complex network analysis tools. It also addresses a gap in ecological research by extending the methodology to behavioral studies using video-based image recognition, as most current studies rely on still images. The study evaluates CV's performance in estimating species interaction frequency (PIE) and its ecological implications. Results show that up to 10% of pairwise interactions may be missed with CV, with information loss varying among focal species and individual plants. The loss of information is minimal compared to the vast data CV enables researchers to gather especially if data is intended to be used in community-level approaches where only three out of 344 unique pairwise interactions were missed. In community-level approaches, the overall estimates of both PIEs and interaction strengths remained largely unaffected. The methodology provides a valuable resource for ecologists seeking to document ecological interactions efficiently. It offers guidelines for collecting reliable data while addressing CV's limitations in capturing unbiased species interaction data. Despite its constraints, CV significantly enhances the ability to gather large-scale interaction data, particularly at the community level, making it an indispensable tool for ecological research., This work was supported by Ministerio de Ciencia, Innovación y Universidades-MICIU/AEI/10.13039/501100011033/, PID2022-136812NB-I00. LifeWatch ERIC–SUMHAL–FEDER–EU530, LIFEWATCH–2019–09–CSIC–13. Center for Sustainable Landscapes under Global Change, Naturmål and SustainScapes–NNF20OC0059595. Universidad de Sevilla, Plan Propio de Investigación y Transferencia, US., Peer reviewed

By dispersing seeds, frugivorous animals affect spatial co-occurrence of plants, ultimately influencing plant community dynamics. Frugivorous animals are intrinsically involved in plant community dynamics, by dispersing seeds of fleshy-fruited plants and influencing their spatial co-occurrence. Particularly, generalist avian frugivores forage on them, co-dispersing and co-disseminating their seeds. This dispersal process often promotes spatial clumping and sets the initial spatial template on which subsequent ecological processes operate. Despite this mutualism is key at the early stages of fleshy-fruited species, it remains unknown whether these co-dispersed and co-disseminated plant species maintain their initial pairwise seed co-occurrence along their demographic cycle. If so, this would reflect a lasting signal of these early mutualistic interactions in the plant community assembly. We investigated whether plant species that share avian seed dispersers also co-occur spatially across different life stages, from seed to adult individuals. We combined data on seed rain from nine fleshy-fruited species, dispersed by a community of 21 bird species identified through DNA-barcoding, with spatial co-occurrence patterns among these plant species at the sapling and adult stages, using point pattern analyses. From this, we built matrices showing: 1) the number of frugivore species shared between each pair of plant species, and 2) their spatial co-occurrence at different life stages: seeds (seed–seed), saplings (sapling–sapling and sapling–adult), and adults (adult–adult). We then used structural equation modelling to test whether the level of shared avian frugivores predicts the spatial co-occurrence among adult plants, while also assessing the indirect effects of processes occurring at intermediate demographic stages. Our results revealed both direct and indirect effects of frugivore sharing during the fruit removal and seed dissemination stage on the spatial assembly of established adult species. Specifically, we found that higher levels of shared frugivores increased the co-occurrence among plant species in seed rain (seed–seed), apparent facilitation (sapling–adult) and adult community (adult–adult), but not within the sapling bank (sapling–sapling). Frugivorous birds drive the co-occurrence among fleshy-fruited plant species, from seed to adult stages, although their influence decreases as the plant demographic cycle advances. However, the co-occurrence at adult stages is exclusively achieved when adults apparently facilitate sapling establishment. This reinforces the key role of the plant–plant facilitation in Mediterranean systems, and the deterministic role of mutualistic avian frugivores as drivers of spatial assembly in plant communities., – Funding from LIFEWATCH-2019-09-CSIC-13 project supported this research (EQ, GC, PH, PJ). AJP was supported by “Margarita Salas” fellowship (Acción 5a) in University of Jaén funded by Ministerio de Universidades and European Union (Next Generation EU funds). EQ was supported by “la Caixa” Foundation fellowship LCF/BQ/ DE18/1167000. JI was supported by a Formación de Personal Investigador grant PRE2018-085916 from the Spanish Ministry of Science and Innovation. LAL was supported by a predoctoral contract from the University of Cádiz (UCA/REC13VPCT/2020). BAC received funding from the Ministry of Universities of the Spanish Government (Ref. FPU19_02552). PJ was supported by PID2022-136812NB-I00, Spanish Ministry of Science and Innovation, as well as funding from the VI Plan Propio de Investigación of the Universidad de Sevilla (VI PPIT-US)., Peer reviewed

Mutualistic plant-animal interactions for seed dispersal are crucial for vegetation dynamics, benefiting over half of the world’s plant species. Beyond the tropics, the Mediterranean biome harbors the highest proportion of species adapted to endozoochory, yet major gaps remain in quantifying interaction diversity in these biodiversity-rich areas and their links to ecosystem functioning. High-resolution, quantitative interaction data are essential not only to fill these gaps but also to enable large-scale ecological modeling of species interactions across biomes. Here, we present the FRUGivory INTegration (FRUGINT) dataset — an extensive collection of quantitative frugivory interactions and associated species traits from a Mediterranean biodiversity hotspot in southwestern Spain. By integrating six complementary sampling methods (camera trapping, continuous-monitoring cameras, DNA-barcoding, mist-netting, direct observation and track records) across multiple years, the dataset overcomes limitations of sampling biases, variable effort and spatio-temporal heterogeneity, providing a comprehensive picture of plant-frugivore interactions across the region. Based on a total of 37,923 interaction records and 481 unique pairwise interactions, involving 26 fleshy-fruited plant species present in Doñana and 78 frugivorous vertebrate species, FRUGINT yields estimates of regional-scale plant-frugivore networks based on pairwise interaction probabilities. The dataset encompasses both common and numerous rare interactions, offering a valuable resource for advancing research on plant-animal interactions, network ecology, and biodiversity conservation., Completion of this manuscript was funded by grants: PID2022-136812NB-I00 from the Agencia Estatal de Investigación (Agencia Estatal de Investigación and FEDER (MCIN/AEI/10.13039/501100011033/ FEDER, UE). Plan Propio de Investigación y Transferencia, University of Sevilla (2021-2025). LifeWatch ERIC-SUMHAL project (LIFEWATCH-2019-09-CSIC-13) with FEDER-EU530 funding. PID2020-115129RJ-I00 funded by MICIU/AEI/ 10.13039/501100011033. Marie Sklodowska-Curie grant agreement no. 798269; European Union’s Horizon 2020 research and innovation programme., Peer reviewed

Pairwise plant–frugivore mutualistic interactions build up into mega-diverse networks involving dozens of interacting species, being among the most generalized interactions occurring in free-living species. These mutualisms consist of food provisioning by plants and, their counterpart, plant propagule (seeds) movement by the animals, being crucial for the natural vegetation regeneration in many ecosystems. Yet we are far from understanding which part of this enormous interaction biodiversity is needed for their maintenance. I overview the diversity of interaction modes involved in these mutualisms, the main components of the seed dispersal services, and their functional diversity. I examine how interaction richness covaries with partner species richness at different scales, resulting in variable patterns of species complementarities in terms of seed dispersal effects. The functionality of most generalized plant–frugivore mutualisms relies on complementarity of effects across a high diversity of partners, yet frequently depends on just a distinct subset of them, resulting in high functional redundancy. Two distinct aspects are relevant: 1) variable quantitative effects among species; 2) variable pairwise-interaction outcomes, between the extremes of antagonism and mutualism. Frugivory, occurring at the final stage of each plant reproductive episode, entails a large, cumulative, effect of other biotic interactions occurring at earlier stages (e.g. floral herbivory, pollination, pre-dispersal fruit damage). I examine how plant–frugivore interactions mix-up with the whole biotic interactome of a plant, using the Prunus mahaleb system as a case study. The effects of distinct subsets of frugivores combine with different sets of antagonistic and mutualistic partners in other interactions, yet having a lasting signal on final seed dispersal success., – Completion of this manuscript was funded by grant PID2022-136812NB-I00 (Ministry of Science, Innovation and Universities, Spain–MICIU/AEI/10.13039/501100011033/), the Plan Propio de Investigación y Transferencia, University of Sevilla (2021–2024), and a LifeWatch ERIC-SUMHAL project (LIFEWATCH-2019-09-CSIC-13) with FEDER-EU530 funding., Peer reviewed

Pairwise plant-frugivore mutualistic interactions build up into mega-diverse networks involving dozens of interacting species, being among the most generalized interactions occurring in free-living species. These mutualisms consist of food provisioning by plants and their counterpart, plant propagule (seeds) movement by the animals, being crucial for the natural vegetation regeneration in many ecosystems. Yet we are far from understanding which part of this enormous interaction biodiversity is needed for their maintenance. I overview the diversity of interaction modes involved in these mutualisms, the main components of the seed dispersal services, and their functional diversity. I examine how interaction richness covaries with partner species richness at different scales, resulting in variable patterns of species complementarities in terms of seed dispersal effects. The functionality of most generalized plant-frugivore mutualisms relies on complementarity of effects across a high diversity of partners, yet frequently depends on just a distinct subset of them, resulting in high functional redundancy. Two distinct aspects are relevant: 1) variable quantitative effects among species; 2) variable pairwise-interaction outcomes, between the extremes of antagonism and mutualism. Frugivory, occurring at the final stage of each plant reproductive episode, entails a large, cumulative, effect of other biotic interactions occurring at earlier stages (e.g., floral herbivory, pollination, pre-dispersal fruit damage). I examine how plant-frugivore interactions mix-up with the whole biotic interactome of a plant, using the Prunus mahaleb system as a case study. The effects of distinct subsets of frugivores combine with different sets of antagonistic and mutualistic partners in other interactions, yet having a lasting signal on final seed dispersal success., Ministerio de Ciencia, Innovación y Universidades: PID2022–136812NB–I00, MICIU/AEI/10.13039/501100011033/; Universidad de Sevilla, Plan Propio de Investigación y Transferencia, University of Sevilla (2021-2024), Peer reviewed

[EN] Fire is a natural phenomenon that shapes ecosystems, but climate change and human activities have increased its frequency and magnitude, disrupting essential ecosystem services like pollination. Despite advances in understanding fire's effects on species and interactions, the mechanisms shaping ecological networks and determining their functional outcomes remain poorly understood., To address these gaps, we used individual-based plant-pollinator networks to evaluate how do plant traits, phenology and site-related attributes (i.e. proximity of plant individuals, local temperature) shape interaction patterns before and after a fire. We also examined how these factors, along with each plant's position within the network (i.e. centrality), influence individual plant fitness before and after fire., Our results showed that the frequency of interactions of some pollinator functional groups changed after the fire, with an increase in plant–hummingbird interactions and a decrease in plant–lepidopteran interactions. In addition, before the fire the ecological drivers shaping plant-pollinator networks were the number of flowers, floral buds and distance to conspecifics and heterospecifics. Plant–plant facilitation via pollinator sharing, inferred from significant negative effects of shorter conspecific and also heterospecific distances on shaping network structure before fire, increased individual plant fitness only among aggregated individuals of the same species. In contrast, after the fire, flowering synchrony became the main ecological driver shaping plant-pollinator networks, suggesting stronger intraspecific competition, with plant fitness decreasing with higher spatial proximity of heterospecifics, with diluted effects of conspecific facilitation., Synthesis: Facilitative interactions with conspecifics via pollinator sharing were diluted by fire, with potential delayed effects on population recovery. Our findings emphasize that fire causes a shift from facilitative processes to more competitive scenarios in plant-pollinator networks, with implications for population dynamics and community recovery., [PT] O fogo é um fenômeno natural que impacta os ecossistemas, mas as mudanças climáticas e as atividades humanas aumentaram sua frequência e magnitude, prejudicando serviços ecossistêmicos essenciais, como a polinização. Apesar dos avanços na compreensão dos efeitos do fogo sobre as espécies e suas interações, os mecanismos que moldam as redes ecológicas e determinam suas consequências funcionais continuam sendo pouco compreendidos., Para suprir estas lacunas, utilizamos redes de interação entre plantas e polinizadores, baseadas em indivíduos, para avaliar como as características das plantas, a fenologia e os atributos do local (ex. proximidade entre os indivíduos das plantas e a temperatura local) moldam os padrões de interação antes e depois do fogo. Também examinamos como esses fatores, juntamente com a posição de cada planta na rede (ex. centralidade), influenciam a aptidão individual (fitness) das plantas antes e depois do fogo., Nossos resultados mostraram que a frequência das interações de alguns grupos funcionais de polinizadores mudou após o fogo, com um aumento nas interações entre plantas e beija-flores e uma diminuição nas interações entre plantas e lepidópteros. Além disso, antes do fogo, os fatores ecológicos que moldavam as redes entre plantas e polinizadores eram o número de flores e botões florais, bem como as distâncias entre indivíduos da mesma espécie e entre espécies diferentes. A facilitação entre plantas, por meio do compartilhamento de polinizadores, foi inferida a partir dos efeitos negativos significativos. Ou seja, os efeitos das distâncias mais curtas entre indivíduos da mesma espécie e também entre espécies diferentes influenciaram a estrutura da rede antes do fogo, aumentando a aptidão individual das plantas apenas entre indivíduos agrupados da mesma espécie. Em contrapartida, após o fogo, a sincronia da floração tornou-se o principal fator ecológico que moldou as redes entre plantas e polinizadores, sugerindo uma competição intraespecífica mais intensa, com a aptidão das plantas diminuindo à medida que aumentava a proximidade espacial de espécies diferentes, diluindo os efeitos da facilitação entre indivíduos da mesma espécie., Síntese e aplicações. A facilitação entre indivíduos da mesma espécie de planta por meio do compartilhamento de polinizadores foram enfraquecidas pelo fogo, com efeitos que podem retardar a recuperação da população. Nossas descobertas destacam que o fogo provoca uma mudança de processos facilitadores para cenários mais competitivos nas redes planta-polinizador, com implicações para a dinâmica populacional e a recuperação da comunidade., Larissa Alves-de-Lima thanks UFU and FAPEMIG (Fundação de Amparo à Pesquisa do Estado de Minas Gerais) (11589). Diego Anjos thanks UFU and CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) (88887.466638/2019–00). Pedro Jordano was supported by PID2022-136812NB-I00, Spanish Ministry of Science and Innovation, as well as funding from the VI Plan Propio de Investigación of the Universidad de Sevilla (VI PPIT-US). Helena Maura Torezan-Silingardi thanks UFU and CNPQ (Conselho Nacional de Desenvolvimento Científico e Tecnológico) (403647/2021-5). We thank CAPES for funded the Article Processing Charge. The Article Processing Charge for the publication of this research was funded by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) (ROR identifier: 00x0ma614)., Peer reviewed

Resource availability shapes interactions among free-living species, particularly in plant–pollinator mutualisms, where the composition of floral resource types influences pollinator foraging. Bees, known for utilizing a broad array of floral resources, play a crucial role in these dynamics. However, how the structure of interaction networks vary among plants offering different types of floral resources is still poorly understood, as is the role of bee abundance and resource-use breadth in connecting these networks. We examined plant–bee interactions in an area of the Brazilian Cerrado, focusing on nectar, pollen, and oil as floral resource types. We compared the structure of plant–bee interaction networks mediated by these floral resources and evaluated the roles of bees in connecting and enhancing the structural robustness (i.e. resistance to node loss) of the multilayer network resulting from embedding the full set of floral rewards. Unique bee assemblages were associated with each floral resource, leading to distinct network structures. Oil-mediated interactions were more nested and generalized, while nectar- and pollen-mediated networks were more modular and specialized. Alongside abundant generalist species, the network included less abundant bees that interacted with a wide range of plants offering different floral resources, contributing to multilayer structural robustness by forming interconnected cores that enhance cohesion across layers. These findings suggest that differences in plant–bee network structure arise from constraints and specificities imposed by distinct floral resource types, influencing bees' foraging decisions and interaction patterns. Beyond abundant generalist bees, those collecting different types of floral resources contribute to network robustness. Our study highlights how floral resources, bee abundance, and resource-use breadth shape plant–bee network structure in systems with multiple floral resource types., The Fundação Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES – Finance Code 001; CAPES PrInt – grant no. 88887.839479/2023-00) supported CSB's PhD studies at the Programa de Pós-Graduação em Biologia Vegetal (UNESP - Botucatu) and his PhD sandwich. FWA and CSB are associated with the Center for Research on Biodiversity Dynamics and Climate Change (CEPID FAPESP no. 2021/10639-5; CSB Postdoctoral Research grant no. 2024/02640-1). PJ was supported by grant no. PID2022-136812NB-I00, Spanish Ministry of Science and Innovation, as well as funding from the VI Plan Propio de Investigación of the Universidad de Sevilla (VI PPIT-US)., Peer reviewed